Review ArticleActinomycetes A Source of Lignocellulolytic Enzymes

Anita Saini1 Neeraj K Aggarwal1 Anuja Sharma1 and Anita Yadav2

1Department of Microbiology Kurukshetra University Kurukshetra Haryana 136119 India2Department of Biotechnology Kurukshetra University Kurukshetra Haryana 136119 India

Correspondence should be addressed to Neeraj K Aggarwal neerajkuk26rediffmailcom

Received 30 September 2015 Accepted 1 December 2015

Academic Editor Jean-Marie Dupret

Copyright copy 2015 Anita Saini et al This is an open access article distributed under the Creative Commons Attribution Licensewhich permits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

Lignocellulose is the most abundant biomass on earth Agricultural forest and agroindustrial activities generate tons oflignocellulosic wastes annually which present readily procurable economically affordable and renewable feedstock for variouslignocelluloses based applications Lignocelluloses are the focus of present decade researchers globally in an attempt to developtechnologies based onnatural biomass for reducing dependence on expensive and exhaustible substrates Lignocellulolytic enzymesthat is cellulases hemicellulases and lignolytic enzymes play very important role in the processing of lignocelluloses whichis prerequisite for their utilization in various processes These enzymes are obtained from microorganisms distributed in bothprokaryotic and eukaryotic domains including bacteria fungi and actinomycetes Actinomycetes are an attractive microbialgroup for production of lignocellulose degrading enzymes Various studies have evaluated the lignocellulose degrading ability ofactinomycetes which can be potentially implemented in the production of different value added productsThis paper is an overviewof the diversity of cellulolytic hemicellulolytic and lignolytic actinomycetes along with brief discussion of their hydrolytic enzymesystems involved in biomass modification

1 Introduction

Actinomycetes a separate taxonomic group within domainbacteria are members of the order Actinomycetales [1]They are Gram positive bacteria primarily aerobic and sporeformers with high G+C content [2] As their name reflects(in Greek ldquoatkisrdquo means ray and ldquomykesrdquo means fungus)they share some morphological features with fungi [3]They show filamentous growth producing aerial or substratemycelium Actinomycetes are responsible for earthy smell ofthe soil [1] They are ubiquitous in nature found both interrestrial and aquatic habitats [1 4] including mangrovesand sea sediments [5] They belong to both mesophilic andthermophilic groups [6] which broaden the range of habitatsinhabited by them Actinomycetes are known to produce anextensive range of bioactive compounds including variousenzymes having multiple biotechnological applications

Lignocellulolytic enzymes one of the potent enzymesproduced by actinomycetes can be exploited widely in var-ious lignocelluloses based industries [7] Lignocellulases arehydrolytic enzymes capable of degrading tough lignocellulose

in the plant biomass and include cellulases hemicellu-lases and lignolytic enzymes [8] Lignocellulose is the mostabundant renewable biomass on earth [9] It refers to themain constituents of the plant matter that is cellulosehemicellulose and lignin [10] Hydrolysis of lignocellulosicbiomass is accomplished by lignocellulolytic enzymes whichare used in diverse applications [11] Cellulases are usedin production of bioethanol and biomethane in ligandbinding studies [12] textile industry pulp and paper makingdetergents industry animal feed and food and so forth [13]Hemicellulases are employed in biobleaching deinking ofpaper waste clarification of fruit juices upgradation of feedfodder and fibres and saccharification of hemicelluloses toxylose sugars [14] Applications of lignin-degrading enzymesinvolve pretreatment of recalcitrant lignocellulosic biomassfor biofuel production use in paper industry textile industryfood industry wastewater treatment bioremediation organicsynthesis and cosmetic and pharmaceutical industries [15]

Lignocellulolytic enzymes can be obtained from diversetypes of microorganisms including bacteria and fungi [16]Among bacteria actinomycetes are an attractive group being

Hindawi Publishing CorporationEnzyme ResearchVolume 2015 Article ID 279381 15 pageshttpdxdoiorg1011552015279381

2 Enzyme Research

O

OH OH

OHOH

HOHOHO O

OO O

OH OH

OHOH

HOHOO

OO

O

OH OH

OHOH

HOHOO

OOH

Cellobiose unitn

(a) Cellulose polymer

HO OH

HOO

OHHO OH

OHO

OHHO

OHHOO

OH

CH2OH CH2OHCH2O

Glucose Galactose Mannose

HO OH

HOO

HO

OHHO OH

HOOCOOH

OH

OHO

OH

CH2OH

Xylose Arabinose Glucuronic acid

(b) Hemicellulose monomers

OH

CH

CH

p-coumaryl alcohol

CH2OH

Coniferyl alcohol

OH

CH

CH

CH2OH

OCH3

Sinapyl alcohol

OH

CH

CH

CH2OH

OCH3CH3O

1 2 3

(c) Constituents of lignin

Figure 1 Chemical structure of lignocellulose

tapped for production of lignocellulases [7 17ndash20] In thisreview the diversity and applications of lignocellulolytic acti-nomycetes have been discussed along with description oftheir lignocellulases enzyme systems involved in biomassdegradation

2 Lignocellulose Structure and Uses

Lignocellulose is comprised of three main componentsthat is cellulose hemicellulose and lignin [21] (Figure 1)Cellulose is the high molecular weight linear polymer of D-glucopyranose units linked together by 120573-(lrarr 4)-glycosidicbonds with cellobiose dimer being the repeating unit Thecellulose chains are hydrogen bonded to each other makinga bundle of microfibrils which further aggregate together

to make cellulose fibrils [22] The structure shows variationsfrom amorphous to crystalline regions The cellulose fibrilsare packed in the cell wall in a matrix of hemicelluloses andlignin [23] Hemicelluloses are linear or branched hetero-polysaccharides composed of D-xylose L-arabinose D-gal-actose D-glucose or D-mannose sugars with or withoutdifferent uronic acids and may include xylans mannans glu-cans glucuronoxylans arabinoxylans glucomannans galac-tomannans galactoglucomannans 120573-glucans and xyloglu-cans [10 24] The branching and composition vary amongdifferent plant sources Xylan the polymer of xylose is themost abundant hemicellulosic component [10] Lignin is avery complex polyphenolic heteropolymer primarily madeup of three phenyl propionic alcohol monomers that is p-coumaryl alcohol coniferyl alcohol and sinapyl alcohol [25]

Enzyme Research 3

Celluloses hemicelluloses and lignin are packed closelyin a crisscross network and glued with the help of a variety ofnoncovalent and covalent linkages Lignin acts as a cement-ing agent and provides structural support It also providesimpermeable barrier to the enzymes making whole structurerobust and resistant [22] Cellulose is the most abundantpolymer of plant cell walls (35ndash50) followed with hemicel-lulose (20ndash35) and lignin (10ndash25) [10]

Cellulose is used for many purposes such as in manufac-turing of paper and textile fabric in production of biofuel(from fermentable sugar glucose) as inert packing and insu-lating material and in food and drugs as base and stabilizer[26] Hemicelluloses are utilized in a range of applicationsBiodegradability and nontoxicity of hemicelluloses enable itto be used as dietary fibre [27] and as edible coating overfoodstuffs for their stabilization Mannans have cholesterollowering properties [28] Xyloglucans and 120573-glucans are usedin ice-creams and other foods due to their gelling stabilizingand thickening effects Adhesive properties are used in papermaking Hemicelluloses are rich source of xylose sugarswhich are fermented to ethanol fuel [29] Characteristics oflignin also make it an important component used in the pro-duction of a wide range of value added products [30] Lignincan be used as a source of activated carbon and phenol usefulin the synthesis of wide array of chemicals [31] and to producebiocomposites [32] or polymers [33] which can replace plasticbased products either partially or completely It can be usedas binder in small scale processes and as filler in place ofoil based carbon fibres Biosorption properties of lignin [34]are also foreseen as a potent application Furthermore ligninis a rich source of energy [35] Thus lignocellulose is arepertoire of industrially useful components But the struc-tural characteristics of the cell wall or lignocellulose are suchthat the extraction of individual component to homogeneityand purity is a difficult task requiring specific chemical orbiological procedures Also many applications are based ondegradation products of the cellulose hemicellulose or ligninmaking hydrolysis an essential requirement Hydrolysis canbe mediated both chemically and enzymatically Chemicalmethods involve use of expensive chemicals harsh condi-tions and have certain other limitations Enzymatic methodsare therefore preferred and lignocellulolytic enzymes playvery important role in these applications Presently biomassutilization is executed through biorefinery which aims atsignificant utilization of all components generating leastpossible amount of waste When one constituent of biomassis utilized others are simultaneously used for production ofvaluable products During biofuel production sugars fromhemicellulose and cellulose components are used in ethanolformation and lignin residue left is used for power generationor in other uses of lignin Lignocellulases perform importantfunctions in biorefinery processes also

3 Lignocellulolytic EnzymeSystems in Actinomycetes

31 Cellulases Cellulolytic enzymes are a group of glycosylhydrolases classified into different families depending on

their sequence homologies The mechanisms of action andsubstrate specificities vary among different cellulases butthey are generally divided into exoglucanases (EC 32174)endoglucanases (EC 3214) cellobiohydrolases (EC 32191)and 120573-glucosidases (EC 32121) [36 37] Exoglucanasesact on reducing or nonreducing ends of cellulose chainsreleasing glucose units whereas endoglucanases hydrolyse120573-14-glycosidic bonds randomly inside the cellulose chainsreleasing dextrans of variable lengths [38] Cellobiohydro-lases cleave glycosidic bonds at nonreducing ends andrelease cellobiose units [39] These enzymes are particularlyimportant in hydrolysing crystalline cellulose because oftheir processivity [36] 120573-glucosidases enzymes take part inhydrolysis of cellobiose units to monomeric glucose [38]Complete hydrolysis of cellulose involves synergistic effect ofall these enzymes showing synergy between endoglucanasesand exoglucanases (endo-exo synergy) exoglucanases actingon the reducing and nonreducing ends (exo-exo synergy)between cellobiohydrolases and 120573-glucosidases and betweencatalytic and carbohydrate binding domains [39] Figure 2shows schematic presentation of enzymatic hydrolysis of cel-lulose polymer

Microbial cellulase systems are either complexed ornoncomplexed [39] Complexed systems known as cellulo-somes are characteristics of anaerobic bacteria consisting ofmultienzyme complex protuberances from cell surface stabi-lized by dockerin and adhesion proteins In aerobic bacteriaincluding most of the actinomycetes cellulases are noncom-plexed or free and are secreted extracellularly using specificsecretion pathways

Among cellulase producing actinomycetes CellulomonasfimiMicrobispora bispora andThermobifida fusca have beenstudied extensively [6 39] Thermobifida fusca is a ther-mophilic spore forming actinomycete [6] The genome ofT fusca consists of 36 billion bp in a single circular chro-mosome with 3117 coding sequences and has 675 G+Ccontent which stabilizes DNA in extreme temperature condi-tions [40] The genome encodes for 36 glycoside hydrolasesdistributed in 22 GH (glycoside hydrolases) families [40]Cellulase system of T fusca is comprised of six extracellularcellulases (4 endocellulases and 2 exocellulases) and oneintracellular 120573-glucosidase [6 40ndash42] Each enzyme has aseparate catalytic and carbohydrate binding domain bothlinked together with a linker peptide [36 41] Carbohydratebinding domains in all six cellulases belong to the samefamily that is 2CBD [41] The catalytic domains howeverare different in all enzymes belonging to different familieswith Cel5A and Cel5B from GH family 5 Cel6A and Cel6Bfrom GH family 6 Cel9A and Cel9B from GH family 9 andCel48A from family 48 [43] Cel5A Cel5B Cel6A and Cel9Bare endocellulases and do not show processivity [36 40 41]Cel6B and Cel48A are processive exocellulases which act atnonreducing and reducing ends respectively Cel9A is a novelprocessive cellulase with both exo- and endocellulase actionsstarting exohydrolysis from the nonreducing end [36] Theposition of CBD varies in different cellulases which is N-terminus in Cel5A Cel6B and Cel48A and C-terminus inCel6A Cel9A and Cel9B [41] Sequence studies of catalyticdomains have revealed less than 31 similarity between

4 Enzyme Research

Glucose

Cellobiohydrolase

Cellobiohydrolase

Cellobiose units

Cellulose polymer

Oligosaccharides ofvariable length

120573-glucosidase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

Figure 2 Scheme of cellulose hydrolysis

cellulases from taxonomically similar as well as dissimilarmicrobes which indicates development of cellulase systemas a result of horizontal gene transfer compared to the geneduplication [44] The three-dimensional structure of Cel6Ashows 120572-120573 barrel with a deep active site cleft formed by oneshorter and one turned loop consisting of four conservedAsp residues [42 45] Active site of GH48 is also in a cleft[42] Structural characteristics of exocellulases allow them tobind processively whereas open active sites of endocellulasesenable them to bind cellulose internally at random sites [42]Structural elucidation of Cel9A has shown that it gives activ-ity between exo- and endocellulases because its weak bindingdomain 3c CBD is aligned with the active site in the catalyticdomain allowing processive hydrolysis by the enzyme [42]Tfusca also produces cellulose and chitin binding proteins E7and E8 the CBM33 proteins which improve cellulose hydrol-ysis mediated through exoglucanases [36 40 42] Cellulasesfrom T bifida have also been found showing synergism withendocellulases and Trichoderma reesei CBHI [6]

Cellulomonas fimi is a facultative anaerobe but it does notconsist of cellulosomes of cellulolytic anaerobes rather it pro-duces free cellulases [46] Similarly facultatively anaerobicCellulomonas flavigena also secrets free cellulases [46] Bothcarry out efficient hydrolysis of celluloses and hemicelluloses

The cellulase enzyme systems in Cellulomonas fimi also con-sist of six cellulases [39] that is three endocellulases (CenACenB and CenD) two exocellulases (CbhA and CbhB) anda processive endocellulase CenC [41 46] All these enzymeshave activities similar to that inT bifida with same families ofCBDs (2CBD) and catalytic domains but different sequences[41 47] These cellulases are primarily secreted by sec depen-dent pathway [46] and therefore do not require intracellularfolding or cofactors for their activity InC fimiATCC484 andC flavigena ATCC 482 another enzyme GH94 (cellobiosephosphorylase) has also been discovered [46] Microbisporabispora also shows synthesis of six different cellulases show-ing exo-exo and endo-exo synergism [6] Genomic studiesof Streptomyces sp SirexAA-E (ActE) isolated from pine-boring woodwasp Sirex noctilio have also shown genes forGH48 (CBH activity) GH74 (endocellulase) and CDB33[48] Streptomyces coelicolor consisted of 221 carbohydrateactive enzymes (CAZy) or 154 glycosyl hydrolases (GHs)encoded within 86 billion bp long genome [48]

The expression of cellulolytic genes in T bifida is inducedby cellobiose whereas easily utilizable sugar glucose showscatabolite repression as in most of the other cellulolyticmicrobes [49]The regulation of cellulolytic genes ismediatedby the CelR repressor which binds to a 9ndash14 bp palindrome

Enzyme Research 5

(51015840-TGGGAGCGCTCCCA-31015840) in region 51015840-upstream of thecellulase genes in the absence of cellobiose or cellulose [50]During induction cellulose is hydrolysed to cellobiose byconstitutive cellulases Cellobiose binds to CelR releasing itfrom the promoter allowing transcription of the downstreamgenes [47] In addition to the presence of glucose cellulasegenes expression is also regulated by cAMP levels

The extracellular cellulases are secreted by actinomycetesusing either one or both of the common bacterial systemsfor secretion of extracellular proteins that is sec generalsecretion system and sec independent twin-arginine translo-cation (TAT) systems The general Secretion route catalysestransmembrane translocation of proteins in their unfoldedconformation whereas Twin-arginine (TAT) system translo-cates secretory proteins in their native folded state InT bifidaboth of these systems were discovered whereas S coelicolormainly utilizes TAT systems for protein export [40]

32 Hemicellulases Hemicellulases are generally synthesisedalong with cellulases [36 39] Xylan and mannan are mostabundant components of hemicelluloses Complete hydroly-sis of xylan involves an enzyme system consisting of endo-14-120573-xylanases (EC 3218) 120573-D-xylosidases (EC 32137)120572-L-arabinofuranosidases (EC 32155) 120572-glucuronidases(EC 321139) acetyl xylan esterases (EC 31172) and fer-uliccoumaric acid esterases (EC 31173) Mannan is hydrol-ysed primarily by synergistic action of mannanases (EC32178) 120573-mannosidases (EC 32125) and 120572-galactosidases(EC 32122) [36 51] Mannanases hydrolyze 120573-14-glycosidicbonds internally 120573-mannosidase cleave 120573-14 linked man-nose from nonreducing ends and 120572-galactosidase removesterminal D-galactosyl residues linked by 120572-16 linkages [52]Degradation of mannan and xylan also enhances cellulosehydrolysis as they are known to inhibit cellulase activities[36]

Most of the hemicellulases belong to glycosyl hydrolasesfamilies however some enzymes involved in hemicellulosehydrolysis belong to glycosyltransferases (EC 241x) [39]Xylanases hydrolysing internal 120573-14-glycosidic bonds areclassified into GH families 5 7 8 10 11 and 43 120573-D-xylos-idases hydrolyse xylose monomers from nonreducing endsof xylan oligosaccharides and belong to GH families 3 3943 52 and 54 [53] Studies have indicated production ofseveral xylanases by T bifida and other actinomycetes Tbifida has been found to be producing 120573-14-endoxylan-ases (xyl10A xyl10B and xil11A) xylosidases 120572-L-arabino-furanosidases xyloglucanases 120573-13-glucanases (GH81) and120572-N-arabinofuranosidases (xil43) [36 40] Cellulomonas fimisynthesises extracellular endo- as well as exo-xylanases xylanbinding domain CBM4 120573-mannanase mannosidase andxel74 (xyloglucan specific 120573-14-glucanase) [36 46] Cellu-lomonas flavigena ATCC 482 are known to synthesize anunusual mixture of 19 endoxylanases along with GH10GH11 and GH30 xylanases GH43 (120573-xylosidase) GH51 120572-arabinofuranosidase and 120572-glucuronidase GH26 and GH13mannans and GH16 and GH81 120573-glucanase [46] Strepto-myces flavogriseus has shown production of 120573-14-glucan glu-canohydrolase [54] Xylanase genes GH5 (120573-mannosidase)

GH10 (beta xylanase) GH11 (beta xylanase) CE4 (acetylxy-lan esterase) and GH6 (CBH) and GH9 (CBH) have alsobeen found in Streptomyces sp SirexAA-E (ActE) [48]

33 Lignolytic Enzymes Lignin degradation is mediated bya complex of enzymes containing three principal enzymeslaccases (EC 11032) manganese peroxidases (MnP EC111113) and lignin peroxidases (LiP EC 111114) [55 56]Laccases are the oxidoreductases which degrade polyphenolthe principal recalcitrant component in the lignocellulose [1557] They are extracellular inducible enzymes which employsimple oxygen as an oxidizing agent as well as cofactorThey are multicopper oxidases having four copper atoms intheir active sites taking part in oxygen reduction [58] Lowsubstrate specificity of laccases enables them to degrade widevariety of compounds Manganese and lignin peroxidasesare together known as heme peroxidases containing proto-porphyrin IX as a prosthetic group Lignin peroxidases canspecifically degrade high redox potential compounds and areknown to oxidize phenolic as well as nonphenolic aromaticrings which make up around 90 of the lignin polymerThey require H

2O2for their activity Veratryl alcohol is an

attractive substrate for LiP which oxidises other substratesby acting as the redox mediator for indirect oxidation Man-ganese peroxidases are low redox potential heme peroxidasesrequiring H

2O2for their activity They can be manganese

dependent or versatile peroxidases [56 58]Laccases or Laccase-likemulticopper oxidases containing

(LMCO) four copper atoms are classified in types 1 2 and3 [57 59] The four copper atoms are distributed in threedomains in most of the bacterial and fungal laccases [60]Structural studies in several actinomycetes however haverevealed presence of two Cu-binding domains rather thanthree [61 62] The two-domain structure has been named assmall laccase or small LMCO [59 61] LMCOs in Strepto-myces griseus Streptomyces cyaneus Streptomyces coelicolorStreptomyces ipomoea Streptomyces sviceus Streptomyces spand Thermobifida fusca are active as dimers or trimmers[61 63 64]

4 Genetic Engineering

The genes from several lignocellulolytic actinomycetes havebeen successfully cloned to show heterologous expression indifferent microbes GH1 and GH3 enzymes of C fimi ATCC484 expressed in E coli have shown efficient hydrolysis ofcelluloses and xylanases [65] CelStrep gene from cellulolyticStreptomyces sp G12 cloned and expressed in E coli wasfound to belong to GH12 family and catalysed hydrolysis ofcarboxymethylcellulose following aMichaelis-Menten kinet-ics with a 119870

119898of 913mgmL and a 119881max of 3469 120583Mminminus1

[66] Streptomyces reticuli consists of Cel1 gene encodingfor avicelase enzyme which alone can hydrolyse crystallinecellulose effectively [67] When this gene was cloned andexpressed in E coli Bacillus subtilis and Streptomyces sppenzyme was produced but in lower amounts probably dueto the absence of genes encoding for essential regulatoryfactors [68] From xylanolytic Actinomadura sp strain FC7two genes xylI and xylII have been cloned expressed and

6 Enzyme Research

well characterized in Streptomyces lividans [69] Xylanasegene xylBS27 belonging to GH11 from Streptomyces sp S27has been successfully cloned and expressed in Pichia pastorishydrolysing xylan to xylobiose [70] Similarly expression oflaccase gene from Streptomyces coelicolor (SLAC) in Strepto-myces lividans produced large amount of high purity laccase(350mg Lminus1) [71] The gene for a thermostable laccase fromStreptomyces lavendulae REN-7 was successfully cloned andexpressed in E coli [72] Cloning of a lignin peroxidase fromStreptomyces viridosporus T7A into Streptomyces lividansTK64 has resulted in better lignocellulose degradation bygenetically engineered S lividans compared to S lividansTK64 [73] Thus genetic engineering techniques can be andare being used for constructing industrially valuable strainswith potent applications based on actinomycetes lignocel-lulolytic enzymes

5 Diversity of Lignocellulolytic Actinomycetes

51 Cellulolytic Actinomycetes Cellulolytic potential of acti-nomycetes has been explored since inspection of othermicroorganisms for cellulase production Various researchstudies support high cellulose degradation potential ofmicrobes from actinomycetales Table 1 represents the diver-sity of actinomycetes producing cellulase enzymes

52 Hemicellulolytic Actinomycetes Diverse types of actino-mycetes belonging to wide range of habitats and active indifferent environmental conditions are known to producehemicellulolytic enzymes Streptomyces have been found tobe the most abundant hemicellulases producer among acti-nomycetes In a study by Boroujeni et al [74] all of theisolated hemicellulolytic actinomycetes were found to belongto Streptomyces genus Xylanase has been successfully puri-fied from Streptomyces sp E-86 and characterized for itsxylanolytic activity [75] Optimization studies were carriedout for endoxylanase production by Streptomyces sp F2621isolated from Turkey [76] 120573-xylosidase activity of Strepto-myces has been used in saccharification of ball milled wheatstraw [77] A thermostable xylanase was obtained from Strep-tomyces sp QG-11-3 which has shown biobleaching effects ineucalyptus kraft pulp [78] Xylanase has also been producedfrom other strains of Streptomyces sp such as Streptomycessp strain C1-3 [79] Streptomyces sp CD3 [80] Streptomycessp 7b [81] Streptomyces sp PC22 Streptomyces sp 234P-16ISWU-10 Streptomyces sp MDS [82] and Streptomyces sp[83] In Streptomyces rochei and Streptomyces chromofuscusxylanase production has been achieved using treated Papyrusand cotton stalk pulp The obtained xylanase when used forstudying bleaching effects has shown enhanced brightness inthe presence of EDTA [84] Xylanases have also been obtainedfrom Streptomyces albus and Streptomyces hygroscopicus andhave shown successful production of biogas using oil cakeand straw waste [85]

In a study by Ninawe et al [86] three Streptomyces iso-lates that is Streptomyces cyaneus S tendae and S caelestiswere found to be xylanolytic and the enzyme from Strep-tomyces cyaneus was successfully purified followed with its

characterization [87] Streptomyces thermoviolaceus OPC-520 exhibited production of acetyl xylan esterases and 120572-L-arabinofuranosidases enzymes [88] Extracellular xylanaseproduction has also been observed in Streptomyces aureofa-ciens [89] and in Streptomyces coelicolor grown on differentagricultural wastes such as sugarcane bagasse pineappleorange and pomegranate peels [90] Xylanases from Strep-tomyces albus and Streptomyces chromofuscus have indicatedpositive bleaching effects in rice straw pulp [91] Bhosale et al[92] have shown production of 326 IUmL of xylanases fromStreptomyces rameus using sugarcane bagasse along withpeptone and dextrose [92] Studies have indicated productionof cellulase free xylanases from Streptomyces roseiscleroticus[93 94] and Saccharomonospora viridis [95] Improvement inxylanase production has been documented in Streptomycespseudogriseolus subjected to UV mutagenesis [96] Xylanaseproduction has also been seen in Thermomonospora fusca[77 97]Thermomonospora curvataThermomonospora albaMicromonospora Microbispora bispora Nocardia Saccha-romonospora viridis and Thermoactinomyces have shownproduction of 120573-xylosidases acetylesterases and arabinofu-ranosidases [77] Extracellular xylanases have been partiallypurified and characterized in Microbispora siamensis in astudy by Boondaeng et al [98] Xylanase production has alsobeen observed inMicrotetraspora flexuosa [99] Streptomyceschattanoogensis UAH 23 [100] Streptomyces chattanoogensisCECT 3336 [101] Streptomyces violaceoruber [102] Ther-moactinomyces thalophilus [103]Thermomonospora sp [104]Streptomyces thermocyanaeviolaceus [105] and Streptomyceslividans [106] The enzyme from Streptomyces lividans waspurified and characterized by different researchers [107 108]Microbispora sp has been found to be producing hemicellu-lolytic mannanase enzyme [109] whereas other studies haveindicated production of 120573-xylosidases by Streptomyces albo-griseolus S nitrosporeus and Micromonospora melanosporea[110]

53 Lignolytic Actinomycetes Lignolytic activity is exhibitedby diverse range of actinomycetes which play importantrole in biodegradation processes in the environment Searchis in progress for more actinomycetes with high lignolyticpotential using advanced techniques combinedwith conven-tional methods A study by Fernandes et al [111] have usedspecifically designed primers for detection of laccase-likegeneswithin actinomycetes andhas identified gene fragmentsundetectable by known primers which corresponded tosuperfamilies I and K based on laccase and multicopperoxidase engineering database Arias et al [112] have shownproduction of laccase by Streptomyces cyaneus CECT 3335using soya flour Laccase was purified and characterizedand was found to show increase in brightness of eucalyptuskraft pulp in biobleaching studies The enzyme was able tooxidize veratryl alcohol suggesting potential of the strainin industrial applications Veratryl alcohol oxidation andother lignolytic activities have also been demonstrated inStreptomyces viridosporus [113 114]

Streptomyces sp strain EC-22 strain EC1 Streptomycesbadius Streptomyces cyaneus MT813 Thermomonosporafusca Thermomonospora chromogena Thermomonospora

Enzyme Research 7

Table1Cellulase

prod

ucingactin

omycetes

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Streptom

ycessp

120573-glucosid

ases

Saccharifi

catio

nof

rices

traw

[77]

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109130]

Endo

glucanases

Enzymep

rodu

ction

[131]

Cellulases

Aid

incompo

sting

[132]

Streptom

ycesspstrain

AT7

Carboxym

ethylcellulose

(Cx)

andAv

icelase(C1

)enzyme

Prod

uctio

nof

enzymes

[133]

Streptom

ycesspstrainsM

7aandM7bF2621

LIPIMC-A-

194LIPIMC-A-

251andLIPIMC-A-

278

Endo

glucanasesCMCa

se(carbo

xymethylcellulose)

CMCa

seprod

uctio

n[76134135]

Streptom

ycesspT3-1

Scale-up

ofenzymep

rodu

ction

[136]

Streptom

ycesspS7

Enzymep

rodu

ctionusingfruitw

aste

[137]

Streptom

ycesalbogrise

olus

Streptom

ycesnitro

sporeus

120573-glucosid

aseendo

glucanaseandavicelase

Optim

izationstu

dies

forp

rodu

ctionof

cellu

lases

[110]

Streptom

yceslividan

sCellulases

Enzymec

haracterization

Enzymep

rodu

ctionop

timization

[106]

[138]

Streptom

ycesflavogrise

usStreptom

ycesnitro

sporus

Cellulases

Optim

izationof

enzymep

rodu

ction

[138]

Streptom

ycesalbadu

ncus

Exoglucanase120573

-glucosid

aseandendo

glucanase

Catabo

liter

epressionstu

dies

Characteriz

ationof

enzyme

[139]

[140]

Streptom

ycesreticuli

Avicelase

Genee

xpressionstu

dies

[68]

Streptom

ycescellu

lolyticus

Extracellularc

ellulases

Cellulose

decompo

sition

[141]

Streptom

ycesdrozdowicz

iiEn

doglucanase

Successfu

lapp

licationin

detergentand

textile

processin

g[14

2]Streptom

ycesmalachitofuscus

Streptom

ycesglo

meratusand

Streptom

ycesstram

ineus

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109]

Streptom

ycesgancidicu

sEx

tracellularc

ellulaseC

MCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[109143]

Streptom

ycesactuosus

Endo

glucanase

Optim

izationof

CMCa

seprod

uctio

n[14

4]Streptom

ycesbobiliLIPIMC-A-

283

Streptom

ycesolivochrom

ogenesLIPIMC-A-

247

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[135]

Streptom

ycesglo

bosus

Streptom

ycesalan

osinicu

sCM

Caseend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]Streptom

ycesruber

Optim

izationstu

dies

Streptom

ycesvirid

obrunn

eusS

CPE-09

Endo

glucanase

Enzymep

rodu

ctionfro

magroindu

strialresidues

[145]

Streptom

ycesvirid

iochromogenes

AvicelaseCM

Caseand

totalcellulase

Saccharifi

catio

nof

rices

trawandethano

lprodu

ction

[146]

Streptom

ycesalbospinus

Extracellularc

ellulases

Degradatio

nof

cellu

losic

materials

[147]

Streptom

ycesgrise

orub

ens

Cellulases

Enzymep

rodu

ctionop

timization

[148]

Streptom

ycesmatensis

Endo

glucanasea

ndexoglucanase

Enzymep

rodu

ctionop

timization

[149]

Streptom

yceslongisp

ororub

erCa

rboxym

ethylcellulose

Prod

uctio

nandpu

rificatio

nof

enzyme

[150]

Cellu

lomonas

fmi

Cellulase

Activ

ityob

served

[151]

[152]

FPase(totalcellulases)

saccharifi

catio

nof

rices

traw

andethano

lprodu

ction

andop

timizationof

enzymep

rodu

ctionusingwheat

straw

[153]

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 3

Celluloses hemicelluloses and lignin are packed closelyin a crisscross network and glued with the help of a variety ofnoncovalent and covalent linkages Lignin acts as a cement-ing agent and provides structural support It also providesimpermeable barrier to the enzymes making whole structurerobust and resistant [22] Cellulose is the most abundantpolymer of plant cell walls (35ndash50) followed with hemicel-lulose (20ndash35) and lignin (10ndash25) [10]

Cellulose is used for many purposes such as in manufac-turing of paper and textile fabric in production of biofuel(from fermentable sugar glucose) as inert packing and insu-lating material and in food and drugs as base and stabilizer[26] Hemicelluloses are utilized in a range of applicationsBiodegradability and nontoxicity of hemicelluloses enable itto be used as dietary fibre [27] and as edible coating overfoodstuffs for their stabilization Mannans have cholesterollowering properties [28] Xyloglucans and 120573-glucans are usedin ice-creams and other foods due to their gelling stabilizingand thickening effects Adhesive properties are used in papermaking Hemicelluloses are rich source of xylose sugarswhich are fermented to ethanol fuel [29] Characteristics oflignin also make it an important component used in the pro-duction of a wide range of value added products [30] Lignincan be used as a source of activated carbon and phenol usefulin the synthesis of wide array of chemicals [31] and to producebiocomposites [32] or polymers [33] which can replace plasticbased products either partially or completely It can be usedas binder in small scale processes and as filler in place ofoil based carbon fibres Biosorption properties of lignin [34]are also foreseen as a potent application Furthermore ligninis a rich source of energy [35] Thus lignocellulose is arepertoire of industrially useful components But the struc-tural characteristics of the cell wall or lignocellulose are suchthat the extraction of individual component to homogeneityand purity is a difficult task requiring specific chemical orbiological procedures Also many applications are based ondegradation products of the cellulose hemicellulose or ligninmaking hydrolysis an essential requirement Hydrolysis canbe mediated both chemically and enzymatically Chemicalmethods involve use of expensive chemicals harsh condi-tions and have certain other limitations Enzymatic methodsare therefore preferred and lignocellulolytic enzymes playvery important role in these applications Presently biomassutilization is executed through biorefinery which aims atsignificant utilization of all components generating leastpossible amount of waste When one constituent of biomassis utilized others are simultaneously used for production ofvaluable products During biofuel production sugars fromhemicellulose and cellulose components are used in ethanolformation and lignin residue left is used for power generationor in other uses of lignin Lignocellulases perform importantfunctions in biorefinery processes also

3 Lignocellulolytic EnzymeSystems in Actinomycetes

31 Cellulases Cellulolytic enzymes are a group of glycosylhydrolases classified into different families depending on

their sequence homologies The mechanisms of action andsubstrate specificities vary among different cellulases butthey are generally divided into exoglucanases (EC 32174)endoglucanases (EC 3214) cellobiohydrolases (EC 32191)and 120573-glucosidases (EC 32121) [36 37] Exoglucanasesact on reducing or nonreducing ends of cellulose chainsreleasing glucose units whereas endoglucanases hydrolyse120573-14-glycosidic bonds randomly inside the cellulose chainsreleasing dextrans of variable lengths [38] Cellobiohydro-lases cleave glycosidic bonds at nonreducing ends andrelease cellobiose units [39] These enzymes are particularlyimportant in hydrolysing crystalline cellulose because oftheir processivity [36] 120573-glucosidases enzymes take part inhydrolysis of cellobiose units to monomeric glucose [38]Complete hydrolysis of cellulose involves synergistic effect ofall these enzymes showing synergy between endoglucanasesand exoglucanases (endo-exo synergy) exoglucanases actingon the reducing and nonreducing ends (exo-exo synergy)between cellobiohydrolases and 120573-glucosidases and betweencatalytic and carbohydrate binding domains [39] Figure 2shows schematic presentation of enzymatic hydrolysis of cel-lulose polymer

Microbial cellulase systems are either complexed ornoncomplexed [39] Complexed systems known as cellulo-somes are characteristics of anaerobic bacteria consisting ofmultienzyme complex protuberances from cell surface stabi-lized by dockerin and adhesion proteins In aerobic bacteriaincluding most of the actinomycetes cellulases are noncom-plexed or free and are secreted extracellularly using specificsecretion pathways

Among cellulase producing actinomycetes CellulomonasfimiMicrobispora bispora andThermobifida fusca have beenstudied extensively [6 39] Thermobifida fusca is a ther-mophilic spore forming actinomycete [6] The genome ofT fusca consists of 36 billion bp in a single circular chro-mosome with 3117 coding sequences and has 675 G+Ccontent which stabilizes DNA in extreme temperature condi-tions [40] The genome encodes for 36 glycoside hydrolasesdistributed in 22 GH (glycoside hydrolases) families [40]Cellulase system of T fusca is comprised of six extracellularcellulases (4 endocellulases and 2 exocellulases) and oneintracellular 120573-glucosidase [6 40ndash42] Each enzyme has aseparate catalytic and carbohydrate binding domain bothlinked together with a linker peptide [36 41] Carbohydratebinding domains in all six cellulases belong to the samefamily that is 2CBD [41] The catalytic domains howeverare different in all enzymes belonging to different familieswith Cel5A and Cel5B from GH family 5 Cel6A and Cel6Bfrom GH family 6 Cel9A and Cel9B from GH family 9 andCel48A from family 48 [43] Cel5A Cel5B Cel6A and Cel9Bare endocellulases and do not show processivity [36 40 41]Cel6B and Cel48A are processive exocellulases which act atnonreducing and reducing ends respectively Cel9A is a novelprocessive cellulase with both exo- and endocellulase actionsstarting exohydrolysis from the nonreducing end [36] Theposition of CBD varies in different cellulases which is N-terminus in Cel5A Cel6B and Cel48A and C-terminus inCel6A Cel9A and Cel9B [41] Sequence studies of catalyticdomains have revealed less than 31 similarity between

4 Enzyme Research

Glucose

Cellobiohydrolase

Cellobiohydrolase

Cellobiose units

Cellulose polymer

Oligosaccharides ofvariable length

120573-glucosidase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

Figure 2 Scheme of cellulose hydrolysis

cellulases from taxonomically similar as well as dissimilarmicrobes which indicates development of cellulase systemas a result of horizontal gene transfer compared to the geneduplication [44] The three-dimensional structure of Cel6Ashows 120572-120573 barrel with a deep active site cleft formed by oneshorter and one turned loop consisting of four conservedAsp residues [42 45] Active site of GH48 is also in a cleft[42] Structural characteristics of exocellulases allow them tobind processively whereas open active sites of endocellulasesenable them to bind cellulose internally at random sites [42]Structural elucidation of Cel9A has shown that it gives activ-ity between exo- and endocellulases because its weak bindingdomain 3c CBD is aligned with the active site in the catalyticdomain allowing processive hydrolysis by the enzyme [42]Tfusca also produces cellulose and chitin binding proteins E7and E8 the CBM33 proteins which improve cellulose hydrol-ysis mediated through exoglucanases [36 40 42] Cellulasesfrom T bifida have also been found showing synergism withendocellulases and Trichoderma reesei CBHI [6]

Cellulomonas fimi is a facultative anaerobe but it does notconsist of cellulosomes of cellulolytic anaerobes rather it pro-duces free cellulases [46] Similarly facultatively anaerobicCellulomonas flavigena also secrets free cellulases [46] Bothcarry out efficient hydrolysis of celluloses and hemicelluloses

The cellulase enzyme systems in Cellulomonas fimi also con-sist of six cellulases [39] that is three endocellulases (CenACenB and CenD) two exocellulases (CbhA and CbhB) anda processive endocellulase CenC [41 46] All these enzymeshave activities similar to that inT bifida with same families ofCBDs (2CBD) and catalytic domains but different sequences[41 47] These cellulases are primarily secreted by sec depen-dent pathway [46] and therefore do not require intracellularfolding or cofactors for their activity InC fimiATCC484 andC flavigena ATCC 482 another enzyme GH94 (cellobiosephosphorylase) has also been discovered [46] Microbisporabispora also shows synthesis of six different cellulases show-ing exo-exo and endo-exo synergism [6] Genomic studiesof Streptomyces sp SirexAA-E (ActE) isolated from pine-boring woodwasp Sirex noctilio have also shown genes forGH48 (CBH activity) GH74 (endocellulase) and CDB33[48] Streptomyces coelicolor consisted of 221 carbohydrateactive enzymes (CAZy) or 154 glycosyl hydrolases (GHs)encoded within 86 billion bp long genome [48]

The expression of cellulolytic genes in T bifida is inducedby cellobiose whereas easily utilizable sugar glucose showscatabolite repression as in most of the other cellulolyticmicrobes [49]The regulation of cellulolytic genes ismediatedby the CelR repressor which binds to a 9ndash14 bp palindrome

Enzyme Research 5

(51015840-TGGGAGCGCTCCCA-31015840) in region 51015840-upstream of thecellulase genes in the absence of cellobiose or cellulose [50]During induction cellulose is hydrolysed to cellobiose byconstitutive cellulases Cellobiose binds to CelR releasing itfrom the promoter allowing transcription of the downstreamgenes [47] In addition to the presence of glucose cellulasegenes expression is also regulated by cAMP levels

The extracellular cellulases are secreted by actinomycetesusing either one or both of the common bacterial systemsfor secretion of extracellular proteins that is sec generalsecretion system and sec independent twin-arginine translo-cation (TAT) systems The general Secretion route catalysestransmembrane translocation of proteins in their unfoldedconformation whereas Twin-arginine (TAT) system translo-cates secretory proteins in their native folded state InT bifidaboth of these systems were discovered whereas S coelicolormainly utilizes TAT systems for protein export [40]

32 Hemicellulases Hemicellulases are generally synthesisedalong with cellulases [36 39] Xylan and mannan are mostabundant components of hemicelluloses Complete hydroly-sis of xylan involves an enzyme system consisting of endo-14-120573-xylanases (EC 3218) 120573-D-xylosidases (EC 32137)120572-L-arabinofuranosidases (EC 32155) 120572-glucuronidases(EC 321139) acetyl xylan esterases (EC 31172) and fer-uliccoumaric acid esterases (EC 31173) Mannan is hydrol-ysed primarily by synergistic action of mannanases (EC32178) 120573-mannosidases (EC 32125) and 120572-galactosidases(EC 32122) [36 51] Mannanases hydrolyze 120573-14-glycosidicbonds internally 120573-mannosidase cleave 120573-14 linked man-nose from nonreducing ends and 120572-galactosidase removesterminal D-galactosyl residues linked by 120572-16 linkages [52]Degradation of mannan and xylan also enhances cellulosehydrolysis as they are known to inhibit cellulase activities[36]

Most of the hemicellulases belong to glycosyl hydrolasesfamilies however some enzymes involved in hemicellulosehydrolysis belong to glycosyltransferases (EC 241x) [39]Xylanases hydrolysing internal 120573-14-glycosidic bonds areclassified into GH families 5 7 8 10 11 and 43 120573-D-xylos-idases hydrolyse xylose monomers from nonreducing endsof xylan oligosaccharides and belong to GH families 3 3943 52 and 54 [53] Studies have indicated production ofseveral xylanases by T bifida and other actinomycetes Tbifida has been found to be producing 120573-14-endoxylan-ases (xyl10A xyl10B and xil11A) xylosidases 120572-L-arabino-furanosidases xyloglucanases 120573-13-glucanases (GH81) and120572-N-arabinofuranosidases (xil43) [36 40] Cellulomonas fimisynthesises extracellular endo- as well as exo-xylanases xylanbinding domain CBM4 120573-mannanase mannosidase andxel74 (xyloglucan specific 120573-14-glucanase) [36 46] Cellu-lomonas flavigena ATCC 482 are known to synthesize anunusual mixture of 19 endoxylanases along with GH10GH11 and GH30 xylanases GH43 (120573-xylosidase) GH51 120572-arabinofuranosidase and 120572-glucuronidase GH26 and GH13mannans and GH16 and GH81 120573-glucanase [46] Strepto-myces flavogriseus has shown production of 120573-14-glucan glu-canohydrolase [54] Xylanase genes GH5 (120573-mannosidase)

GH10 (beta xylanase) GH11 (beta xylanase) CE4 (acetylxy-lan esterase) and GH6 (CBH) and GH9 (CBH) have alsobeen found in Streptomyces sp SirexAA-E (ActE) [48]

33 Lignolytic Enzymes Lignin degradation is mediated bya complex of enzymes containing three principal enzymeslaccases (EC 11032) manganese peroxidases (MnP EC111113) and lignin peroxidases (LiP EC 111114) [55 56]Laccases are the oxidoreductases which degrade polyphenolthe principal recalcitrant component in the lignocellulose [1557] They are extracellular inducible enzymes which employsimple oxygen as an oxidizing agent as well as cofactorThey are multicopper oxidases having four copper atoms intheir active sites taking part in oxygen reduction [58] Lowsubstrate specificity of laccases enables them to degrade widevariety of compounds Manganese and lignin peroxidasesare together known as heme peroxidases containing proto-porphyrin IX as a prosthetic group Lignin peroxidases canspecifically degrade high redox potential compounds and areknown to oxidize phenolic as well as nonphenolic aromaticrings which make up around 90 of the lignin polymerThey require H

2O2for their activity Veratryl alcohol is an

attractive substrate for LiP which oxidises other substratesby acting as the redox mediator for indirect oxidation Man-ganese peroxidases are low redox potential heme peroxidasesrequiring H

2O2for their activity They can be manganese

dependent or versatile peroxidases [56 58]Laccases or Laccase-likemulticopper oxidases containing

(LMCO) four copper atoms are classified in types 1 2 and3 [57 59] The four copper atoms are distributed in threedomains in most of the bacterial and fungal laccases [60]Structural studies in several actinomycetes however haverevealed presence of two Cu-binding domains rather thanthree [61 62] The two-domain structure has been named assmall laccase or small LMCO [59 61] LMCOs in Strepto-myces griseus Streptomyces cyaneus Streptomyces coelicolorStreptomyces ipomoea Streptomyces sviceus Streptomyces spand Thermobifida fusca are active as dimers or trimmers[61 63 64]

4 Genetic Engineering

The genes from several lignocellulolytic actinomycetes havebeen successfully cloned to show heterologous expression indifferent microbes GH1 and GH3 enzymes of C fimi ATCC484 expressed in E coli have shown efficient hydrolysis ofcelluloses and xylanases [65] CelStrep gene from cellulolyticStreptomyces sp G12 cloned and expressed in E coli wasfound to belong to GH12 family and catalysed hydrolysis ofcarboxymethylcellulose following aMichaelis-Menten kinet-ics with a 119870

119898of 913mgmL and a 119881max of 3469 120583Mminminus1

[66] Streptomyces reticuli consists of Cel1 gene encodingfor avicelase enzyme which alone can hydrolyse crystallinecellulose effectively [67] When this gene was cloned andexpressed in E coli Bacillus subtilis and Streptomyces sppenzyme was produced but in lower amounts probably dueto the absence of genes encoding for essential regulatoryfactors [68] From xylanolytic Actinomadura sp strain FC7two genes xylI and xylII have been cloned expressed and

6 Enzyme Research

well characterized in Streptomyces lividans [69] Xylanasegene xylBS27 belonging to GH11 from Streptomyces sp S27has been successfully cloned and expressed in Pichia pastorishydrolysing xylan to xylobiose [70] Similarly expression oflaccase gene from Streptomyces coelicolor (SLAC) in Strepto-myces lividans produced large amount of high purity laccase(350mg Lminus1) [71] The gene for a thermostable laccase fromStreptomyces lavendulae REN-7 was successfully cloned andexpressed in E coli [72] Cloning of a lignin peroxidase fromStreptomyces viridosporus T7A into Streptomyces lividansTK64 has resulted in better lignocellulose degradation bygenetically engineered S lividans compared to S lividansTK64 [73] Thus genetic engineering techniques can be andare being used for constructing industrially valuable strainswith potent applications based on actinomycetes lignocel-lulolytic enzymes

5 Diversity of Lignocellulolytic Actinomycetes

51 Cellulolytic Actinomycetes Cellulolytic potential of acti-nomycetes has been explored since inspection of othermicroorganisms for cellulase production Various researchstudies support high cellulose degradation potential ofmicrobes from actinomycetales Table 1 represents the diver-sity of actinomycetes producing cellulase enzymes

52 Hemicellulolytic Actinomycetes Diverse types of actino-mycetes belonging to wide range of habitats and active indifferent environmental conditions are known to producehemicellulolytic enzymes Streptomyces have been found tobe the most abundant hemicellulases producer among acti-nomycetes In a study by Boroujeni et al [74] all of theisolated hemicellulolytic actinomycetes were found to belongto Streptomyces genus Xylanase has been successfully puri-fied from Streptomyces sp E-86 and characterized for itsxylanolytic activity [75] Optimization studies were carriedout for endoxylanase production by Streptomyces sp F2621isolated from Turkey [76] 120573-xylosidase activity of Strepto-myces has been used in saccharification of ball milled wheatstraw [77] A thermostable xylanase was obtained from Strep-tomyces sp QG-11-3 which has shown biobleaching effects ineucalyptus kraft pulp [78] Xylanase has also been producedfrom other strains of Streptomyces sp such as Streptomycessp strain C1-3 [79] Streptomyces sp CD3 [80] Streptomycessp 7b [81] Streptomyces sp PC22 Streptomyces sp 234P-16ISWU-10 Streptomyces sp MDS [82] and Streptomyces sp[83] In Streptomyces rochei and Streptomyces chromofuscusxylanase production has been achieved using treated Papyrusand cotton stalk pulp The obtained xylanase when used forstudying bleaching effects has shown enhanced brightness inthe presence of EDTA [84] Xylanases have also been obtainedfrom Streptomyces albus and Streptomyces hygroscopicus andhave shown successful production of biogas using oil cakeand straw waste [85]

In a study by Ninawe et al [86] three Streptomyces iso-lates that is Streptomyces cyaneus S tendae and S caelestiswere found to be xylanolytic and the enzyme from Strep-tomyces cyaneus was successfully purified followed with its

characterization [87] Streptomyces thermoviolaceus OPC-520 exhibited production of acetyl xylan esterases and 120572-L-arabinofuranosidases enzymes [88] Extracellular xylanaseproduction has also been observed in Streptomyces aureofa-ciens [89] and in Streptomyces coelicolor grown on differentagricultural wastes such as sugarcane bagasse pineappleorange and pomegranate peels [90] Xylanases from Strep-tomyces albus and Streptomyces chromofuscus have indicatedpositive bleaching effects in rice straw pulp [91] Bhosale et al[92] have shown production of 326 IUmL of xylanases fromStreptomyces rameus using sugarcane bagasse along withpeptone and dextrose [92] Studies have indicated productionof cellulase free xylanases from Streptomyces roseiscleroticus[93 94] and Saccharomonospora viridis [95] Improvement inxylanase production has been documented in Streptomycespseudogriseolus subjected to UV mutagenesis [96] Xylanaseproduction has also been seen in Thermomonospora fusca[77 97]Thermomonospora curvataThermomonospora albaMicromonospora Microbispora bispora Nocardia Saccha-romonospora viridis and Thermoactinomyces have shownproduction of 120573-xylosidases acetylesterases and arabinofu-ranosidases [77] Extracellular xylanases have been partiallypurified and characterized in Microbispora siamensis in astudy by Boondaeng et al [98] Xylanase production has alsobeen observed inMicrotetraspora flexuosa [99] Streptomyceschattanoogensis UAH 23 [100] Streptomyces chattanoogensisCECT 3336 [101] Streptomyces violaceoruber [102] Ther-moactinomyces thalophilus [103]Thermomonospora sp [104]Streptomyces thermocyanaeviolaceus [105] and Streptomyceslividans [106] The enzyme from Streptomyces lividans waspurified and characterized by different researchers [107 108]Microbispora sp has been found to be producing hemicellu-lolytic mannanase enzyme [109] whereas other studies haveindicated production of 120573-xylosidases by Streptomyces albo-griseolus S nitrosporeus and Micromonospora melanosporea[110]

53 Lignolytic Actinomycetes Lignolytic activity is exhibitedby diverse range of actinomycetes which play importantrole in biodegradation processes in the environment Searchis in progress for more actinomycetes with high lignolyticpotential using advanced techniques combinedwith conven-tional methods A study by Fernandes et al [111] have usedspecifically designed primers for detection of laccase-likegeneswithin actinomycetes andhas identified gene fragmentsundetectable by known primers which corresponded tosuperfamilies I and K based on laccase and multicopperoxidase engineering database Arias et al [112] have shownproduction of laccase by Streptomyces cyaneus CECT 3335using soya flour Laccase was purified and characterizedand was found to show increase in brightness of eucalyptuskraft pulp in biobleaching studies The enzyme was able tooxidize veratryl alcohol suggesting potential of the strainin industrial applications Veratryl alcohol oxidation andother lignolytic activities have also been demonstrated inStreptomyces viridosporus [113 114]

Streptomyces sp strain EC-22 strain EC1 Streptomycesbadius Streptomyces cyaneus MT813 Thermomonosporafusca Thermomonospora chromogena Thermomonospora

Enzyme Research 7

Table1Cellulase

prod

ucingactin

omycetes

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Streptom

ycessp

120573-glucosid

ases

Saccharifi

catio

nof

rices

traw

[77]

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109130]

Endo

glucanases

Enzymep

rodu

ction

[131]

Cellulases

Aid

incompo

sting

[132]

Streptom

ycesspstrain

AT7

Carboxym

ethylcellulose

(Cx)

andAv

icelase(C1

)enzyme

Prod

uctio

nof

enzymes

[133]

Streptom

ycesspstrainsM

7aandM7bF2621

LIPIMC-A-

194LIPIMC-A-

251andLIPIMC-A-

278

Endo

glucanasesCMCa

se(carbo

xymethylcellulose)

CMCa

seprod

uctio

n[76134135]

Streptom

ycesspT3-1

Scale-up

ofenzymep

rodu

ction

[136]

Streptom

ycesspS7

Enzymep

rodu

ctionusingfruitw

aste

[137]

Streptom

ycesalbogrise

olus

Streptom

ycesnitro

sporeus

120573-glucosid

aseendo

glucanaseandavicelase

Optim

izationstu

dies

forp

rodu

ctionof

cellu

lases

[110]

Streptom

yceslividan

sCellulases

Enzymec

haracterization

Enzymep

rodu

ctionop

timization

[106]

[138]

Streptom

ycesflavogrise

usStreptom

ycesnitro

sporus

Cellulases

Optim

izationof

enzymep

rodu

ction

[138]

Streptom

ycesalbadu

ncus

Exoglucanase120573

-glucosid

aseandendo

glucanase

Catabo

liter

epressionstu

dies

Characteriz

ationof

enzyme

[139]

[140]

Streptom

ycesreticuli

Avicelase

Genee

xpressionstu

dies

[68]

Streptom

ycescellu

lolyticus

Extracellularc

ellulases

Cellulose

decompo

sition

[141]

Streptom

ycesdrozdowicz

iiEn

doglucanase

Successfu

lapp

licationin

detergentand

textile

processin

g[14

2]Streptom

ycesmalachitofuscus

Streptom

ycesglo

meratusand

Streptom

ycesstram

ineus

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109]

Streptom

ycesgancidicu

sEx

tracellularc

ellulaseC

MCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[109143]

Streptom

ycesactuosus

Endo

glucanase

Optim

izationof

CMCa

seprod

uctio

n[14

4]Streptom

ycesbobiliLIPIMC-A-

283

Streptom

ycesolivochrom

ogenesLIPIMC-A-

247

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[135]

Streptom

ycesglo

bosus

Streptom

ycesalan

osinicu

sCM

Caseend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]Streptom

ycesruber

Optim

izationstu

dies

Streptom

ycesvirid

obrunn

eusS

CPE-09

Endo

glucanase

Enzymep

rodu

ctionfro

magroindu

strialresidues

[145]

Streptom

ycesvirid

iochromogenes

AvicelaseCM

Caseand

totalcellulase

Saccharifi

catio

nof

rices

trawandethano

lprodu

ction

[146]

Streptom

ycesalbospinus

Extracellularc

ellulases

Degradatio

nof

cellu

losic

materials

[147]

Streptom

ycesgrise

orub

ens

Cellulases

Enzymep

rodu

ctionop

timization

[148]

Streptom

ycesmatensis

Endo

glucanasea

ndexoglucanase

Enzymep

rodu

ctionop

timization

[149]

Streptom

yceslongisp

ororub

erCa

rboxym

ethylcellulose

Prod

uctio

nandpu

rificatio

nof

enzyme

[150]

Cellu

lomonas

fmi

Cellulase

Activ

ityob

served

[151]

[152]

FPase(totalcellulases)

saccharifi

catio

nof

rices

traw

andethano

lprodu

ction

andop

timizationof

enzymep

rodu

ctionusingwheat

straw

[153]

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

4 Enzyme Research

Glucose

Cellobiohydrolase

Cellobiohydrolase

Cellobiose units

Cellulose polymer

Oligosaccharides ofvariable length

120573-glucosidase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-endoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

120573-14-exoglucanase

Figure 2 Scheme of cellulose hydrolysis

cellulases from taxonomically similar as well as dissimilarmicrobes which indicates development of cellulase systemas a result of horizontal gene transfer compared to the geneduplication [44] The three-dimensional structure of Cel6Ashows 120572-120573 barrel with a deep active site cleft formed by oneshorter and one turned loop consisting of four conservedAsp residues [42 45] Active site of GH48 is also in a cleft[42] Structural characteristics of exocellulases allow them tobind processively whereas open active sites of endocellulasesenable them to bind cellulose internally at random sites [42]Structural elucidation of Cel9A has shown that it gives activ-ity between exo- and endocellulases because its weak bindingdomain 3c CBD is aligned with the active site in the catalyticdomain allowing processive hydrolysis by the enzyme [42]Tfusca also produces cellulose and chitin binding proteins E7and E8 the CBM33 proteins which improve cellulose hydrol-ysis mediated through exoglucanases [36 40 42] Cellulasesfrom T bifida have also been found showing synergism withendocellulases and Trichoderma reesei CBHI [6]

Cellulomonas fimi is a facultative anaerobe but it does notconsist of cellulosomes of cellulolytic anaerobes rather it pro-duces free cellulases [46] Similarly facultatively anaerobicCellulomonas flavigena also secrets free cellulases [46] Bothcarry out efficient hydrolysis of celluloses and hemicelluloses

The cellulase enzyme systems in Cellulomonas fimi also con-sist of six cellulases [39] that is three endocellulases (CenACenB and CenD) two exocellulases (CbhA and CbhB) anda processive endocellulase CenC [41 46] All these enzymeshave activities similar to that inT bifida with same families ofCBDs (2CBD) and catalytic domains but different sequences[41 47] These cellulases are primarily secreted by sec depen-dent pathway [46] and therefore do not require intracellularfolding or cofactors for their activity InC fimiATCC484 andC flavigena ATCC 482 another enzyme GH94 (cellobiosephosphorylase) has also been discovered [46] Microbisporabispora also shows synthesis of six different cellulases show-ing exo-exo and endo-exo synergism [6] Genomic studiesof Streptomyces sp SirexAA-E (ActE) isolated from pine-boring woodwasp Sirex noctilio have also shown genes forGH48 (CBH activity) GH74 (endocellulase) and CDB33[48] Streptomyces coelicolor consisted of 221 carbohydrateactive enzymes (CAZy) or 154 glycosyl hydrolases (GHs)encoded within 86 billion bp long genome [48]

The expression of cellulolytic genes in T bifida is inducedby cellobiose whereas easily utilizable sugar glucose showscatabolite repression as in most of the other cellulolyticmicrobes [49]The regulation of cellulolytic genes ismediatedby the CelR repressor which binds to a 9ndash14 bp palindrome

Enzyme Research 5

(51015840-TGGGAGCGCTCCCA-31015840) in region 51015840-upstream of thecellulase genes in the absence of cellobiose or cellulose [50]During induction cellulose is hydrolysed to cellobiose byconstitutive cellulases Cellobiose binds to CelR releasing itfrom the promoter allowing transcription of the downstreamgenes [47] In addition to the presence of glucose cellulasegenes expression is also regulated by cAMP levels

The extracellular cellulases are secreted by actinomycetesusing either one or both of the common bacterial systemsfor secretion of extracellular proteins that is sec generalsecretion system and sec independent twin-arginine translo-cation (TAT) systems The general Secretion route catalysestransmembrane translocation of proteins in their unfoldedconformation whereas Twin-arginine (TAT) system translo-cates secretory proteins in their native folded state InT bifidaboth of these systems were discovered whereas S coelicolormainly utilizes TAT systems for protein export [40]

32 Hemicellulases Hemicellulases are generally synthesisedalong with cellulases [36 39] Xylan and mannan are mostabundant components of hemicelluloses Complete hydroly-sis of xylan involves an enzyme system consisting of endo-14-120573-xylanases (EC 3218) 120573-D-xylosidases (EC 32137)120572-L-arabinofuranosidases (EC 32155) 120572-glucuronidases(EC 321139) acetyl xylan esterases (EC 31172) and fer-uliccoumaric acid esterases (EC 31173) Mannan is hydrol-ysed primarily by synergistic action of mannanases (EC32178) 120573-mannosidases (EC 32125) and 120572-galactosidases(EC 32122) [36 51] Mannanases hydrolyze 120573-14-glycosidicbonds internally 120573-mannosidase cleave 120573-14 linked man-nose from nonreducing ends and 120572-galactosidase removesterminal D-galactosyl residues linked by 120572-16 linkages [52]Degradation of mannan and xylan also enhances cellulosehydrolysis as they are known to inhibit cellulase activities[36]

Most of the hemicellulases belong to glycosyl hydrolasesfamilies however some enzymes involved in hemicellulosehydrolysis belong to glycosyltransferases (EC 241x) [39]Xylanases hydrolysing internal 120573-14-glycosidic bonds areclassified into GH families 5 7 8 10 11 and 43 120573-D-xylos-idases hydrolyse xylose monomers from nonreducing endsof xylan oligosaccharides and belong to GH families 3 3943 52 and 54 [53] Studies have indicated production ofseveral xylanases by T bifida and other actinomycetes Tbifida has been found to be producing 120573-14-endoxylan-ases (xyl10A xyl10B and xil11A) xylosidases 120572-L-arabino-furanosidases xyloglucanases 120573-13-glucanases (GH81) and120572-N-arabinofuranosidases (xil43) [36 40] Cellulomonas fimisynthesises extracellular endo- as well as exo-xylanases xylanbinding domain CBM4 120573-mannanase mannosidase andxel74 (xyloglucan specific 120573-14-glucanase) [36 46] Cellu-lomonas flavigena ATCC 482 are known to synthesize anunusual mixture of 19 endoxylanases along with GH10GH11 and GH30 xylanases GH43 (120573-xylosidase) GH51 120572-arabinofuranosidase and 120572-glucuronidase GH26 and GH13mannans and GH16 and GH81 120573-glucanase [46] Strepto-myces flavogriseus has shown production of 120573-14-glucan glu-canohydrolase [54] Xylanase genes GH5 (120573-mannosidase)

GH10 (beta xylanase) GH11 (beta xylanase) CE4 (acetylxy-lan esterase) and GH6 (CBH) and GH9 (CBH) have alsobeen found in Streptomyces sp SirexAA-E (ActE) [48]

33 Lignolytic Enzymes Lignin degradation is mediated bya complex of enzymes containing three principal enzymeslaccases (EC 11032) manganese peroxidases (MnP EC111113) and lignin peroxidases (LiP EC 111114) [55 56]Laccases are the oxidoreductases which degrade polyphenolthe principal recalcitrant component in the lignocellulose [1557] They are extracellular inducible enzymes which employsimple oxygen as an oxidizing agent as well as cofactorThey are multicopper oxidases having four copper atoms intheir active sites taking part in oxygen reduction [58] Lowsubstrate specificity of laccases enables them to degrade widevariety of compounds Manganese and lignin peroxidasesare together known as heme peroxidases containing proto-porphyrin IX as a prosthetic group Lignin peroxidases canspecifically degrade high redox potential compounds and areknown to oxidize phenolic as well as nonphenolic aromaticrings which make up around 90 of the lignin polymerThey require H

2O2for their activity Veratryl alcohol is an

attractive substrate for LiP which oxidises other substratesby acting as the redox mediator for indirect oxidation Man-ganese peroxidases are low redox potential heme peroxidasesrequiring H

2O2for their activity They can be manganese

dependent or versatile peroxidases [56 58]Laccases or Laccase-likemulticopper oxidases containing

(LMCO) four copper atoms are classified in types 1 2 and3 [57 59] The four copper atoms are distributed in threedomains in most of the bacterial and fungal laccases [60]Structural studies in several actinomycetes however haverevealed presence of two Cu-binding domains rather thanthree [61 62] The two-domain structure has been named assmall laccase or small LMCO [59 61] LMCOs in Strepto-myces griseus Streptomyces cyaneus Streptomyces coelicolorStreptomyces ipomoea Streptomyces sviceus Streptomyces spand Thermobifida fusca are active as dimers or trimmers[61 63 64]

4 Genetic Engineering

The genes from several lignocellulolytic actinomycetes havebeen successfully cloned to show heterologous expression indifferent microbes GH1 and GH3 enzymes of C fimi ATCC484 expressed in E coli have shown efficient hydrolysis ofcelluloses and xylanases [65] CelStrep gene from cellulolyticStreptomyces sp G12 cloned and expressed in E coli wasfound to belong to GH12 family and catalysed hydrolysis ofcarboxymethylcellulose following aMichaelis-Menten kinet-ics with a 119870

119898of 913mgmL and a 119881max of 3469 120583Mminminus1

[66] Streptomyces reticuli consists of Cel1 gene encodingfor avicelase enzyme which alone can hydrolyse crystallinecellulose effectively [67] When this gene was cloned andexpressed in E coli Bacillus subtilis and Streptomyces sppenzyme was produced but in lower amounts probably dueto the absence of genes encoding for essential regulatoryfactors [68] From xylanolytic Actinomadura sp strain FC7two genes xylI and xylII have been cloned expressed and

6 Enzyme Research

well characterized in Streptomyces lividans [69] Xylanasegene xylBS27 belonging to GH11 from Streptomyces sp S27has been successfully cloned and expressed in Pichia pastorishydrolysing xylan to xylobiose [70] Similarly expression oflaccase gene from Streptomyces coelicolor (SLAC) in Strepto-myces lividans produced large amount of high purity laccase(350mg Lminus1) [71] The gene for a thermostable laccase fromStreptomyces lavendulae REN-7 was successfully cloned andexpressed in E coli [72] Cloning of a lignin peroxidase fromStreptomyces viridosporus T7A into Streptomyces lividansTK64 has resulted in better lignocellulose degradation bygenetically engineered S lividans compared to S lividansTK64 [73] Thus genetic engineering techniques can be andare being used for constructing industrially valuable strainswith potent applications based on actinomycetes lignocel-lulolytic enzymes

5 Diversity of Lignocellulolytic Actinomycetes

51 Cellulolytic Actinomycetes Cellulolytic potential of acti-nomycetes has been explored since inspection of othermicroorganisms for cellulase production Various researchstudies support high cellulose degradation potential ofmicrobes from actinomycetales Table 1 represents the diver-sity of actinomycetes producing cellulase enzymes

52 Hemicellulolytic Actinomycetes Diverse types of actino-mycetes belonging to wide range of habitats and active indifferent environmental conditions are known to producehemicellulolytic enzymes Streptomyces have been found tobe the most abundant hemicellulases producer among acti-nomycetes In a study by Boroujeni et al [74] all of theisolated hemicellulolytic actinomycetes were found to belongto Streptomyces genus Xylanase has been successfully puri-fied from Streptomyces sp E-86 and characterized for itsxylanolytic activity [75] Optimization studies were carriedout for endoxylanase production by Streptomyces sp F2621isolated from Turkey [76] 120573-xylosidase activity of Strepto-myces has been used in saccharification of ball milled wheatstraw [77] A thermostable xylanase was obtained from Strep-tomyces sp QG-11-3 which has shown biobleaching effects ineucalyptus kraft pulp [78] Xylanase has also been producedfrom other strains of Streptomyces sp such as Streptomycessp strain C1-3 [79] Streptomyces sp CD3 [80] Streptomycessp 7b [81] Streptomyces sp PC22 Streptomyces sp 234P-16ISWU-10 Streptomyces sp MDS [82] and Streptomyces sp[83] In Streptomyces rochei and Streptomyces chromofuscusxylanase production has been achieved using treated Papyrusand cotton stalk pulp The obtained xylanase when used forstudying bleaching effects has shown enhanced brightness inthe presence of EDTA [84] Xylanases have also been obtainedfrom Streptomyces albus and Streptomyces hygroscopicus andhave shown successful production of biogas using oil cakeand straw waste [85]

In a study by Ninawe et al [86] three Streptomyces iso-lates that is Streptomyces cyaneus S tendae and S caelestiswere found to be xylanolytic and the enzyme from Strep-tomyces cyaneus was successfully purified followed with its

characterization [87] Streptomyces thermoviolaceus OPC-520 exhibited production of acetyl xylan esterases and 120572-L-arabinofuranosidases enzymes [88] Extracellular xylanaseproduction has also been observed in Streptomyces aureofa-ciens [89] and in Streptomyces coelicolor grown on differentagricultural wastes such as sugarcane bagasse pineappleorange and pomegranate peels [90] Xylanases from Strep-tomyces albus and Streptomyces chromofuscus have indicatedpositive bleaching effects in rice straw pulp [91] Bhosale et al[92] have shown production of 326 IUmL of xylanases fromStreptomyces rameus using sugarcane bagasse along withpeptone and dextrose [92] Studies have indicated productionof cellulase free xylanases from Streptomyces roseiscleroticus[93 94] and Saccharomonospora viridis [95] Improvement inxylanase production has been documented in Streptomycespseudogriseolus subjected to UV mutagenesis [96] Xylanaseproduction has also been seen in Thermomonospora fusca[77 97]Thermomonospora curvataThermomonospora albaMicromonospora Microbispora bispora Nocardia Saccha-romonospora viridis and Thermoactinomyces have shownproduction of 120573-xylosidases acetylesterases and arabinofu-ranosidases [77] Extracellular xylanases have been partiallypurified and characterized in Microbispora siamensis in astudy by Boondaeng et al [98] Xylanase production has alsobeen observed inMicrotetraspora flexuosa [99] Streptomyceschattanoogensis UAH 23 [100] Streptomyces chattanoogensisCECT 3336 [101] Streptomyces violaceoruber [102] Ther-moactinomyces thalophilus [103]Thermomonospora sp [104]Streptomyces thermocyanaeviolaceus [105] and Streptomyceslividans [106] The enzyme from Streptomyces lividans waspurified and characterized by different researchers [107 108]Microbispora sp has been found to be producing hemicellu-lolytic mannanase enzyme [109] whereas other studies haveindicated production of 120573-xylosidases by Streptomyces albo-griseolus S nitrosporeus and Micromonospora melanosporea[110]

53 Lignolytic Actinomycetes Lignolytic activity is exhibitedby diverse range of actinomycetes which play importantrole in biodegradation processes in the environment Searchis in progress for more actinomycetes with high lignolyticpotential using advanced techniques combinedwith conven-tional methods A study by Fernandes et al [111] have usedspecifically designed primers for detection of laccase-likegeneswithin actinomycetes andhas identified gene fragmentsundetectable by known primers which corresponded tosuperfamilies I and K based on laccase and multicopperoxidase engineering database Arias et al [112] have shownproduction of laccase by Streptomyces cyaneus CECT 3335using soya flour Laccase was purified and characterizedand was found to show increase in brightness of eucalyptuskraft pulp in biobleaching studies The enzyme was able tooxidize veratryl alcohol suggesting potential of the strainin industrial applications Veratryl alcohol oxidation andother lignolytic activities have also been demonstrated inStreptomyces viridosporus [113 114]

Streptomyces sp strain EC-22 strain EC1 Streptomycesbadius Streptomyces cyaneus MT813 Thermomonosporafusca Thermomonospora chromogena Thermomonospora

Enzyme Research 7

Table1Cellulase

prod

ucingactin

omycetes

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Streptom

ycessp

120573-glucosid

ases

Saccharifi

catio

nof

rices

traw

[77]

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109130]

Endo

glucanases

Enzymep

rodu

ction

[131]

Cellulases

Aid

incompo

sting

[132]

Streptom

ycesspstrain

AT7

Carboxym

ethylcellulose

(Cx)

andAv

icelase(C1

)enzyme

Prod

uctio

nof

enzymes

[133]

Streptom

ycesspstrainsM

7aandM7bF2621

LIPIMC-A-

194LIPIMC-A-

251andLIPIMC-A-

278

Endo

glucanasesCMCa

se(carbo

xymethylcellulose)

CMCa

seprod

uctio

n[76134135]

Streptom

ycesspT3-1

Scale-up

ofenzymep

rodu

ction

[136]

Streptom

ycesspS7

Enzymep

rodu

ctionusingfruitw

aste

[137]

Streptom

ycesalbogrise

olus

Streptom

ycesnitro

sporeus

120573-glucosid

aseendo

glucanaseandavicelase

Optim

izationstu

dies

forp

rodu

ctionof

cellu

lases

[110]

Streptom

yceslividan

sCellulases

Enzymec

haracterization

Enzymep

rodu

ctionop

timization

[106]

[138]

Streptom

ycesflavogrise

usStreptom

ycesnitro

sporus

Cellulases

Optim

izationof

enzymep

rodu

ction

[138]

Streptom

ycesalbadu

ncus

Exoglucanase120573

-glucosid

aseandendo

glucanase

Catabo

liter

epressionstu

dies

Characteriz

ationof

enzyme

[139]

[140]

Streptom

ycesreticuli

Avicelase

Genee

xpressionstu

dies

[68]

Streptom

ycescellu

lolyticus

Extracellularc

ellulases

Cellulose

decompo

sition

[141]

Streptom

ycesdrozdowicz

iiEn

doglucanase

Successfu

lapp

licationin

detergentand

textile

processin

g[14

2]Streptom

ycesmalachitofuscus

Streptom

ycesglo

meratusand

Streptom

ycesstram

ineus

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109]

Streptom

ycesgancidicu

sEx

tracellularc

ellulaseC

MCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[109143]

Streptom

ycesactuosus

Endo

glucanase

Optim

izationof

CMCa

seprod

uctio

n[14

4]Streptom

ycesbobiliLIPIMC-A-

283

Streptom

ycesolivochrom

ogenesLIPIMC-A-

247

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[135]

Streptom

ycesglo

bosus

Streptom

ycesalan

osinicu

sCM

Caseend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]Streptom

ycesruber

Optim

izationstu

dies

Streptom

ycesvirid

obrunn

eusS

CPE-09

Endo

glucanase

Enzymep

rodu

ctionfro

magroindu

strialresidues

[145]

Streptom

ycesvirid

iochromogenes

AvicelaseCM

Caseand

totalcellulase

Saccharifi

catio

nof

rices

trawandethano

lprodu

ction

[146]

Streptom

ycesalbospinus

Extracellularc

ellulases

Degradatio

nof

cellu

losic

materials

[147]

Streptom

ycesgrise

orub

ens

Cellulases

Enzymep

rodu

ctionop

timization

[148]

Streptom

ycesmatensis

Endo

glucanasea

ndexoglucanase

Enzymep

rodu

ctionop

timization

[149]

Streptom

yceslongisp

ororub

erCa

rboxym

ethylcellulose

Prod

uctio

nandpu

rificatio

nof

enzyme

[150]

Cellu

lomonas

fmi

Cellulase

Activ

ityob

served

[151]

[152]

FPase(totalcellulases)

saccharifi

catio

nof

rices

traw

andethano

lprodu

ction

andop

timizationof

enzymep

rodu

ctionusingwheat

straw

[153]

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 5

(51015840-TGGGAGCGCTCCCA-31015840) in region 51015840-upstream of thecellulase genes in the absence of cellobiose or cellulose [50]During induction cellulose is hydrolysed to cellobiose byconstitutive cellulases Cellobiose binds to CelR releasing itfrom the promoter allowing transcription of the downstreamgenes [47] In addition to the presence of glucose cellulasegenes expression is also regulated by cAMP levels

The extracellular cellulases are secreted by actinomycetesusing either one or both of the common bacterial systemsfor secretion of extracellular proteins that is sec generalsecretion system and sec independent twin-arginine translo-cation (TAT) systems The general Secretion route catalysestransmembrane translocation of proteins in their unfoldedconformation whereas Twin-arginine (TAT) system translo-cates secretory proteins in their native folded state InT bifidaboth of these systems were discovered whereas S coelicolormainly utilizes TAT systems for protein export [40]

32 Hemicellulases Hemicellulases are generally synthesisedalong with cellulases [36 39] Xylan and mannan are mostabundant components of hemicelluloses Complete hydroly-sis of xylan involves an enzyme system consisting of endo-14-120573-xylanases (EC 3218) 120573-D-xylosidases (EC 32137)120572-L-arabinofuranosidases (EC 32155) 120572-glucuronidases(EC 321139) acetyl xylan esterases (EC 31172) and fer-uliccoumaric acid esterases (EC 31173) Mannan is hydrol-ysed primarily by synergistic action of mannanases (EC32178) 120573-mannosidases (EC 32125) and 120572-galactosidases(EC 32122) [36 51] Mannanases hydrolyze 120573-14-glycosidicbonds internally 120573-mannosidase cleave 120573-14 linked man-nose from nonreducing ends and 120572-galactosidase removesterminal D-galactosyl residues linked by 120572-16 linkages [52]Degradation of mannan and xylan also enhances cellulosehydrolysis as they are known to inhibit cellulase activities[36]

Most of the hemicellulases belong to glycosyl hydrolasesfamilies however some enzymes involved in hemicellulosehydrolysis belong to glycosyltransferases (EC 241x) [39]Xylanases hydrolysing internal 120573-14-glycosidic bonds areclassified into GH families 5 7 8 10 11 and 43 120573-D-xylos-idases hydrolyse xylose monomers from nonreducing endsof xylan oligosaccharides and belong to GH families 3 3943 52 and 54 [53] Studies have indicated production ofseveral xylanases by T bifida and other actinomycetes Tbifida has been found to be producing 120573-14-endoxylan-ases (xyl10A xyl10B and xil11A) xylosidases 120572-L-arabino-furanosidases xyloglucanases 120573-13-glucanases (GH81) and120572-N-arabinofuranosidases (xil43) [36 40] Cellulomonas fimisynthesises extracellular endo- as well as exo-xylanases xylanbinding domain CBM4 120573-mannanase mannosidase andxel74 (xyloglucan specific 120573-14-glucanase) [36 46] Cellu-lomonas flavigena ATCC 482 are known to synthesize anunusual mixture of 19 endoxylanases along with GH10GH11 and GH30 xylanases GH43 (120573-xylosidase) GH51 120572-arabinofuranosidase and 120572-glucuronidase GH26 and GH13mannans and GH16 and GH81 120573-glucanase [46] Strepto-myces flavogriseus has shown production of 120573-14-glucan glu-canohydrolase [54] Xylanase genes GH5 (120573-mannosidase)

GH10 (beta xylanase) GH11 (beta xylanase) CE4 (acetylxy-lan esterase) and GH6 (CBH) and GH9 (CBH) have alsobeen found in Streptomyces sp SirexAA-E (ActE) [48]

33 Lignolytic Enzymes Lignin degradation is mediated bya complex of enzymes containing three principal enzymeslaccases (EC 11032) manganese peroxidases (MnP EC111113) and lignin peroxidases (LiP EC 111114) [55 56]Laccases are the oxidoreductases which degrade polyphenolthe principal recalcitrant component in the lignocellulose [1557] They are extracellular inducible enzymes which employsimple oxygen as an oxidizing agent as well as cofactorThey are multicopper oxidases having four copper atoms intheir active sites taking part in oxygen reduction [58] Lowsubstrate specificity of laccases enables them to degrade widevariety of compounds Manganese and lignin peroxidasesare together known as heme peroxidases containing proto-porphyrin IX as a prosthetic group Lignin peroxidases canspecifically degrade high redox potential compounds and areknown to oxidize phenolic as well as nonphenolic aromaticrings which make up around 90 of the lignin polymerThey require H

2O2for their activity Veratryl alcohol is an

attractive substrate for LiP which oxidises other substratesby acting as the redox mediator for indirect oxidation Man-ganese peroxidases are low redox potential heme peroxidasesrequiring H

2O2for their activity They can be manganese

dependent or versatile peroxidases [56 58]Laccases or Laccase-likemulticopper oxidases containing

(LMCO) four copper atoms are classified in types 1 2 and3 [57 59] The four copper atoms are distributed in threedomains in most of the bacterial and fungal laccases [60]Structural studies in several actinomycetes however haverevealed presence of two Cu-binding domains rather thanthree [61 62] The two-domain structure has been named assmall laccase or small LMCO [59 61] LMCOs in Strepto-myces griseus Streptomyces cyaneus Streptomyces coelicolorStreptomyces ipomoea Streptomyces sviceus Streptomyces spand Thermobifida fusca are active as dimers or trimmers[61 63 64]

4 Genetic Engineering

The genes from several lignocellulolytic actinomycetes havebeen successfully cloned to show heterologous expression indifferent microbes GH1 and GH3 enzymes of C fimi ATCC484 expressed in E coli have shown efficient hydrolysis ofcelluloses and xylanases [65] CelStrep gene from cellulolyticStreptomyces sp G12 cloned and expressed in E coli wasfound to belong to GH12 family and catalysed hydrolysis ofcarboxymethylcellulose following aMichaelis-Menten kinet-ics with a 119870

119898of 913mgmL and a 119881max of 3469 120583Mminminus1

[66] Streptomyces reticuli consists of Cel1 gene encodingfor avicelase enzyme which alone can hydrolyse crystallinecellulose effectively [67] When this gene was cloned andexpressed in E coli Bacillus subtilis and Streptomyces sppenzyme was produced but in lower amounts probably dueto the absence of genes encoding for essential regulatoryfactors [68] From xylanolytic Actinomadura sp strain FC7two genes xylI and xylII have been cloned expressed and

6 Enzyme Research

well characterized in Streptomyces lividans [69] Xylanasegene xylBS27 belonging to GH11 from Streptomyces sp S27has been successfully cloned and expressed in Pichia pastorishydrolysing xylan to xylobiose [70] Similarly expression oflaccase gene from Streptomyces coelicolor (SLAC) in Strepto-myces lividans produced large amount of high purity laccase(350mg Lminus1) [71] The gene for a thermostable laccase fromStreptomyces lavendulae REN-7 was successfully cloned andexpressed in E coli [72] Cloning of a lignin peroxidase fromStreptomyces viridosporus T7A into Streptomyces lividansTK64 has resulted in better lignocellulose degradation bygenetically engineered S lividans compared to S lividansTK64 [73] Thus genetic engineering techniques can be andare being used for constructing industrially valuable strainswith potent applications based on actinomycetes lignocel-lulolytic enzymes

5 Diversity of Lignocellulolytic Actinomycetes

51 Cellulolytic Actinomycetes Cellulolytic potential of acti-nomycetes has been explored since inspection of othermicroorganisms for cellulase production Various researchstudies support high cellulose degradation potential ofmicrobes from actinomycetales Table 1 represents the diver-sity of actinomycetes producing cellulase enzymes

52 Hemicellulolytic Actinomycetes Diverse types of actino-mycetes belonging to wide range of habitats and active indifferent environmental conditions are known to producehemicellulolytic enzymes Streptomyces have been found tobe the most abundant hemicellulases producer among acti-nomycetes In a study by Boroujeni et al [74] all of theisolated hemicellulolytic actinomycetes were found to belongto Streptomyces genus Xylanase has been successfully puri-fied from Streptomyces sp E-86 and characterized for itsxylanolytic activity [75] Optimization studies were carriedout for endoxylanase production by Streptomyces sp F2621isolated from Turkey [76] 120573-xylosidase activity of Strepto-myces has been used in saccharification of ball milled wheatstraw [77] A thermostable xylanase was obtained from Strep-tomyces sp QG-11-3 which has shown biobleaching effects ineucalyptus kraft pulp [78] Xylanase has also been producedfrom other strains of Streptomyces sp such as Streptomycessp strain C1-3 [79] Streptomyces sp CD3 [80] Streptomycessp 7b [81] Streptomyces sp PC22 Streptomyces sp 234P-16ISWU-10 Streptomyces sp MDS [82] and Streptomyces sp[83] In Streptomyces rochei and Streptomyces chromofuscusxylanase production has been achieved using treated Papyrusand cotton stalk pulp The obtained xylanase when used forstudying bleaching effects has shown enhanced brightness inthe presence of EDTA [84] Xylanases have also been obtainedfrom Streptomyces albus and Streptomyces hygroscopicus andhave shown successful production of biogas using oil cakeand straw waste [85]

In a study by Ninawe et al [86] three Streptomyces iso-lates that is Streptomyces cyaneus S tendae and S caelestiswere found to be xylanolytic and the enzyme from Strep-tomyces cyaneus was successfully purified followed with its

characterization [87] Streptomyces thermoviolaceus OPC-520 exhibited production of acetyl xylan esterases and 120572-L-arabinofuranosidases enzymes [88] Extracellular xylanaseproduction has also been observed in Streptomyces aureofa-ciens [89] and in Streptomyces coelicolor grown on differentagricultural wastes such as sugarcane bagasse pineappleorange and pomegranate peels [90] Xylanases from Strep-tomyces albus and Streptomyces chromofuscus have indicatedpositive bleaching effects in rice straw pulp [91] Bhosale et al[92] have shown production of 326 IUmL of xylanases fromStreptomyces rameus using sugarcane bagasse along withpeptone and dextrose [92] Studies have indicated productionof cellulase free xylanases from Streptomyces roseiscleroticus[93 94] and Saccharomonospora viridis [95] Improvement inxylanase production has been documented in Streptomycespseudogriseolus subjected to UV mutagenesis [96] Xylanaseproduction has also been seen in Thermomonospora fusca[77 97]Thermomonospora curvataThermomonospora albaMicromonospora Microbispora bispora Nocardia Saccha-romonospora viridis and Thermoactinomyces have shownproduction of 120573-xylosidases acetylesterases and arabinofu-ranosidases [77] Extracellular xylanases have been partiallypurified and characterized in Microbispora siamensis in astudy by Boondaeng et al [98] Xylanase production has alsobeen observed inMicrotetraspora flexuosa [99] Streptomyceschattanoogensis UAH 23 [100] Streptomyces chattanoogensisCECT 3336 [101] Streptomyces violaceoruber [102] Ther-moactinomyces thalophilus [103]Thermomonospora sp [104]Streptomyces thermocyanaeviolaceus [105] and Streptomyceslividans [106] The enzyme from Streptomyces lividans waspurified and characterized by different researchers [107 108]Microbispora sp has been found to be producing hemicellu-lolytic mannanase enzyme [109] whereas other studies haveindicated production of 120573-xylosidases by Streptomyces albo-griseolus S nitrosporeus and Micromonospora melanosporea[110]

53 Lignolytic Actinomycetes Lignolytic activity is exhibitedby diverse range of actinomycetes which play importantrole in biodegradation processes in the environment Searchis in progress for more actinomycetes with high lignolyticpotential using advanced techniques combinedwith conven-tional methods A study by Fernandes et al [111] have usedspecifically designed primers for detection of laccase-likegeneswithin actinomycetes andhas identified gene fragmentsundetectable by known primers which corresponded tosuperfamilies I and K based on laccase and multicopperoxidase engineering database Arias et al [112] have shownproduction of laccase by Streptomyces cyaneus CECT 3335using soya flour Laccase was purified and characterizedand was found to show increase in brightness of eucalyptuskraft pulp in biobleaching studies The enzyme was able tooxidize veratryl alcohol suggesting potential of the strainin industrial applications Veratryl alcohol oxidation andother lignolytic activities have also been demonstrated inStreptomyces viridosporus [113 114]

Streptomyces sp strain EC-22 strain EC1 Streptomycesbadius Streptomyces cyaneus MT813 Thermomonosporafusca Thermomonospora chromogena Thermomonospora

Enzyme Research 7

Table1Cellulase

prod

ucingactin

omycetes

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Streptom

ycessp

120573-glucosid

ases

Saccharifi

catio

nof

rices

traw

[77]

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109130]

Endo

glucanases

Enzymep

rodu

ction

[131]

Cellulases

Aid

incompo

sting

[132]

Streptom

ycesspstrain

AT7

Carboxym

ethylcellulose

(Cx)

andAv

icelase(C1

)enzyme

Prod

uctio

nof

enzymes

[133]

Streptom

ycesspstrainsM

7aandM7bF2621

LIPIMC-A-

194LIPIMC-A-

251andLIPIMC-A-

278

Endo

glucanasesCMCa

se(carbo

xymethylcellulose)

CMCa

seprod

uctio

n[76134135]

Streptom

ycesspT3-1

Scale-up

ofenzymep

rodu

ction

[136]

Streptom

ycesspS7

Enzymep

rodu

ctionusingfruitw

aste

[137]

Streptom

ycesalbogrise

olus

Streptom

ycesnitro

sporeus

120573-glucosid

aseendo

glucanaseandavicelase

Optim

izationstu

dies

forp

rodu

ctionof

cellu

lases

[110]

Streptom

yceslividan

sCellulases

Enzymec

haracterization

Enzymep

rodu

ctionop

timization

[106]

[138]

Streptom

ycesflavogrise

usStreptom

ycesnitro

sporus

Cellulases

Optim

izationof

enzymep

rodu

ction

[138]

Streptom

ycesalbadu

ncus

Exoglucanase120573

-glucosid

aseandendo

glucanase

Catabo

liter

epressionstu

dies

Characteriz

ationof

enzyme

[139]

[140]

Streptom

ycesreticuli

Avicelase

Genee

xpressionstu

dies

[68]

Streptom

ycescellu

lolyticus

Extracellularc

ellulases

Cellulose

decompo

sition

[141]

Streptom

ycesdrozdowicz

iiEn

doglucanase

Successfu

lapp

licationin

detergentand

textile

processin

g[14

2]Streptom

ycesmalachitofuscus

Streptom

ycesglo

meratusand

Streptom

ycesstram

ineus

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109]

Streptom

ycesgancidicu

sEx

tracellularc

ellulaseC

MCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[109143]

Streptom

ycesactuosus

Endo

glucanase

Optim

izationof

CMCa

seprod

uctio

n[14

4]Streptom

ycesbobiliLIPIMC-A-

283

Streptom

ycesolivochrom

ogenesLIPIMC-A-

247

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[135]

Streptom

ycesglo

bosus

Streptom

ycesalan

osinicu

sCM

Caseend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]Streptom

ycesruber

Optim

izationstu

dies

Streptom

ycesvirid

obrunn

eusS

CPE-09

Endo

glucanase

Enzymep

rodu

ctionfro

magroindu

strialresidues

[145]

Streptom

ycesvirid

iochromogenes

AvicelaseCM

Caseand

totalcellulase

Saccharifi

catio

nof

rices

trawandethano

lprodu

ction

[146]

Streptom

ycesalbospinus

Extracellularc

ellulases

Degradatio

nof

cellu

losic

materials

[147]

Streptom

ycesgrise

orub

ens

Cellulases

Enzymep

rodu

ctionop

timization

[148]

Streptom

ycesmatensis

Endo

glucanasea

ndexoglucanase

Enzymep

rodu

ctionop

timization

[149]

Streptom

yceslongisp

ororub

erCa

rboxym

ethylcellulose

Prod

uctio

nandpu

rificatio

nof

enzyme

[150]

Cellu

lomonas

fmi

Cellulase

Activ

ityob

served

[151]

[152]

FPase(totalcellulases)

saccharifi

catio

nof

rices

traw

andethano

lprodu

ction

andop

timizationof

enzymep

rodu

ctionusingwheat

straw

[153]

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

6 Enzyme Research

well characterized in Streptomyces lividans [69] Xylanasegene xylBS27 belonging to GH11 from Streptomyces sp S27has been successfully cloned and expressed in Pichia pastorishydrolysing xylan to xylobiose [70] Similarly expression oflaccase gene from Streptomyces coelicolor (SLAC) in Strepto-myces lividans produced large amount of high purity laccase(350mg Lminus1) [71] The gene for a thermostable laccase fromStreptomyces lavendulae REN-7 was successfully cloned andexpressed in E coli [72] Cloning of a lignin peroxidase fromStreptomyces viridosporus T7A into Streptomyces lividansTK64 has resulted in better lignocellulose degradation bygenetically engineered S lividans compared to S lividansTK64 [73] Thus genetic engineering techniques can be andare being used for constructing industrially valuable strainswith potent applications based on actinomycetes lignocel-lulolytic enzymes

5 Diversity of Lignocellulolytic Actinomycetes

51 Cellulolytic Actinomycetes Cellulolytic potential of acti-nomycetes has been explored since inspection of othermicroorganisms for cellulase production Various researchstudies support high cellulose degradation potential ofmicrobes from actinomycetales Table 1 represents the diver-sity of actinomycetes producing cellulase enzymes

52 Hemicellulolytic Actinomycetes Diverse types of actino-mycetes belonging to wide range of habitats and active indifferent environmental conditions are known to producehemicellulolytic enzymes Streptomyces have been found tobe the most abundant hemicellulases producer among acti-nomycetes In a study by Boroujeni et al [74] all of theisolated hemicellulolytic actinomycetes were found to belongto Streptomyces genus Xylanase has been successfully puri-fied from Streptomyces sp E-86 and characterized for itsxylanolytic activity [75] Optimization studies were carriedout for endoxylanase production by Streptomyces sp F2621isolated from Turkey [76] 120573-xylosidase activity of Strepto-myces has been used in saccharification of ball milled wheatstraw [77] A thermostable xylanase was obtained from Strep-tomyces sp QG-11-3 which has shown biobleaching effects ineucalyptus kraft pulp [78] Xylanase has also been producedfrom other strains of Streptomyces sp such as Streptomycessp strain C1-3 [79] Streptomyces sp CD3 [80] Streptomycessp 7b [81] Streptomyces sp PC22 Streptomyces sp 234P-16ISWU-10 Streptomyces sp MDS [82] and Streptomyces sp[83] In Streptomyces rochei and Streptomyces chromofuscusxylanase production has been achieved using treated Papyrusand cotton stalk pulp The obtained xylanase when used forstudying bleaching effects has shown enhanced brightness inthe presence of EDTA [84] Xylanases have also been obtainedfrom Streptomyces albus and Streptomyces hygroscopicus andhave shown successful production of biogas using oil cakeand straw waste [85]

In a study by Ninawe et al [86] three Streptomyces iso-lates that is Streptomyces cyaneus S tendae and S caelestiswere found to be xylanolytic and the enzyme from Strep-tomyces cyaneus was successfully purified followed with its

characterization [87] Streptomyces thermoviolaceus OPC-520 exhibited production of acetyl xylan esterases and 120572-L-arabinofuranosidases enzymes [88] Extracellular xylanaseproduction has also been observed in Streptomyces aureofa-ciens [89] and in Streptomyces coelicolor grown on differentagricultural wastes such as sugarcane bagasse pineappleorange and pomegranate peels [90] Xylanases from Strep-tomyces albus and Streptomyces chromofuscus have indicatedpositive bleaching effects in rice straw pulp [91] Bhosale et al[92] have shown production of 326 IUmL of xylanases fromStreptomyces rameus using sugarcane bagasse along withpeptone and dextrose [92] Studies have indicated productionof cellulase free xylanases from Streptomyces roseiscleroticus[93 94] and Saccharomonospora viridis [95] Improvement inxylanase production has been documented in Streptomycespseudogriseolus subjected to UV mutagenesis [96] Xylanaseproduction has also been seen in Thermomonospora fusca[77 97]Thermomonospora curvataThermomonospora albaMicromonospora Microbispora bispora Nocardia Saccha-romonospora viridis and Thermoactinomyces have shownproduction of 120573-xylosidases acetylesterases and arabinofu-ranosidases [77] Extracellular xylanases have been partiallypurified and characterized in Microbispora siamensis in astudy by Boondaeng et al [98] Xylanase production has alsobeen observed inMicrotetraspora flexuosa [99] Streptomyceschattanoogensis UAH 23 [100] Streptomyces chattanoogensisCECT 3336 [101] Streptomyces violaceoruber [102] Ther-moactinomyces thalophilus [103]Thermomonospora sp [104]Streptomyces thermocyanaeviolaceus [105] and Streptomyceslividans [106] The enzyme from Streptomyces lividans waspurified and characterized by different researchers [107 108]Microbispora sp has been found to be producing hemicellu-lolytic mannanase enzyme [109] whereas other studies haveindicated production of 120573-xylosidases by Streptomyces albo-griseolus S nitrosporeus and Micromonospora melanosporea[110]

53 Lignolytic Actinomycetes Lignolytic activity is exhibitedby diverse range of actinomycetes which play importantrole in biodegradation processes in the environment Searchis in progress for more actinomycetes with high lignolyticpotential using advanced techniques combinedwith conven-tional methods A study by Fernandes et al [111] have usedspecifically designed primers for detection of laccase-likegeneswithin actinomycetes andhas identified gene fragmentsundetectable by known primers which corresponded tosuperfamilies I and K based on laccase and multicopperoxidase engineering database Arias et al [112] have shownproduction of laccase by Streptomyces cyaneus CECT 3335using soya flour Laccase was purified and characterizedand was found to show increase in brightness of eucalyptuskraft pulp in biobleaching studies The enzyme was able tooxidize veratryl alcohol suggesting potential of the strainin industrial applications Veratryl alcohol oxidation andother lignolytic activities have also been demonstrated inStreptomyces viridosporus [113 114]

Streptomyces sp strain EC-22 strain EC1 Streptomycesbadius Streptomyces cyaneus MT813 Thermomonosporafusca Thermomonospora chromogena Thermomonospora

Enzyme Research 7

Table1Cellulase

prod

ucingactin

omycetes

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Streptom

ycessp

120573-glucosid

ases

Saccharifi

catio

nof

rices

traw

[77]

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109130]

Endo

glucanases

Enzymep

rodu

ction

[131]

Cellulases

Aid

incompo

sting

[132]

Streptom

ycesspstrain

AT7

Carboxym

ethylcellulose

(Cx)

andAv

icelase(C1

)enzyme

Prod

uctio

nof

enzymes

[133]

Streptom

ycesspstrainsM

7aandM7bF2621

LIPIMC-A-

194LIPIMC-A-

251andLIPIMC-A-

278

Endo

glucanasesCMCa

se(carbo

xymethylcellulose)

CMCa

seprod

uctio

n[76134135]

Streptom

ycesspT3-1

Scale-up

ofenzymep

rodu

ction

[136]

Streptom

ycesspS7

Enzymep

rodu

ctionusingfruitw

aste

[137]

Streptom

ycesalbogrise

olus

Streptom

ycesnitro

sporeus

120573-glucosid

aseendo

glucanaseandavicelase

Optim

izationstu

dies

forp

rodu

ctionof

cellu

lases

[110]

Streptom

yceslividan

sCellulases

Enzymec

haracterization

Enzymep

rodu

ctionop

timization

[106]

[138]

Streptom

ycesflavogrise

usStreptom

ycesnitro

sporus

Cellulases

Optim

izationof

enzymep

rodu

ction

[138]

Streptom

ycesalbadu

ncus

Exoglucanase120573

-glucosid

aseandendo

glucanase

Catabo

liter

epressionstu

dies

Characteriz

ationof

enzyme

[139]

[140]

Streptom

ycesreticuli

Avicelase

Genee

xpressionstu

dies

[68]

Streptom

ycescellu

lolyticus

Extracellularc

ellulases

Cellulose

decompo

sition

[141]

Streptom

ycesdrozdowicz

iiEn

doglucanase

Successfu

lapp

licationin

detergentand

textile

processin

g[14

2]Streptom

ycesmalachitofuscus

Streptom

ycesglo

meratusand

Streptom

ycesstram

ineus

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109]

Streptom

ycesgancidicu

sEx

tracellularc

ellulaseC

MCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[109143]

Streptom

ycesactuosus

Endo

glucanase

Optim

izationof

CMCa

seprod

uctio

n[14

4]Streptom

ycesbobiliLIPIMC-A-

283

Streptom

ycesolivochrom

ogenesLIPIMC-A-

247

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[135]

Streptom

ycesglo

bosus

Streptom

ycesalan

osinicu

sCM

Caseend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]Streptom

ycesruber

Optim

izationstu

dies

Streptom

ycesvirid

obrunn

eusS

CPE-09

Endo

glucanase

Enzymep

rodu

ctionfro

magroindu

strialresidues

[145]

Streptom

ycesvirid

iochromogenes

AvicelaseCM

Caseand

totalcellulase

Saccharifi

catio

nof

rices

trawandethano

lprodu

ction

[146]

Streptom

ycesalbospinus

Extracellularc

ellulases

Degradatio

nof

cellu

losic

materials

[147]

Streptom

ycesgrise

orub

ens

Cellulases

Enzymep

rodu

ctionop

timization

[148]

Streptom

ycesmatensis

Endo

glucanasea

ndexoglucanase

Enzymep

rodu

ctionop

timization

[149]

Streptom

yceslongisp

ororub

erCa

rboxym

ethylcellulose

Prod

uctio

nandpu

rificatio

nof

enzyme

[150]

Cellu

lomonas

fmi

Cellulase

Activ

ityob

served

[151]

[152]

FPase(totalcellulases)

saccharifi

catio

nof

rices

traw

andethano

lprodu

ction

andop

timizationof

enzymep

rodu

ctionusingwheat

straw

[153]

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 7

Table1Cellulase

prod

ucingactin

omycetes

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Streptom

ycessp

120573-glucosid

ases

Saccharifi

catio

nof

rices

traw

[77]

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109130]

Endo

glucanases

Enzymep

rodu

ction

[131]

Cellulases

Aid

incompo

sting

[132]

Streptom

ycesspstrain

AT7

Carboxym

ethylcellulose

(Cx)

andAv

icelase(C1

)enzyme

Prod

uctio

nof

enzymes

[133]

Streptom

ycesspstrainsM

7aandM7bF2621

LIPIMC-A-

194LIPIMC-A-

251andLIPIMC-A-

278

Endo

glucanasesCMCa

se(carbo

xymethylcellulose)

CMCa

seprod

uctio

n[76134135]

Streptom

ycesspT3-1

Scale-up

ofenzymep

rodu

ction

[136]

Streptom

ycesspS7

Enzymep

rodu

ctionusingfruitw

aste

[137]

Streptom

ycesalbogrise

olus

Streptom

ycesnitro

sporeus

120573-glucosid

aseendo

glucanaseandavicelase

Optim

izationstu

dies

forp

rodu

ctionof

cellu

lases

[110]

Streptom

yceslividan

sCellulases

Enzymec

haracterization

Enzymep

rodu

ctionop

timization

[106]

[138]

Streptom

ycesflavogrise

usStreptom

ycesnitro

sporus

Cellulases

Optim

izationof

enzymep

rodu

ction

[138]

Streptom

ycesalbadu

ncus

Exoglucanase120573

-glucosid

aseandendo

glucanase

Catabo

liter

epressionstu

dies

Characteriz

ationof

enzyme

[139]

[140]

Streptom

ycesreticuli

Avicelase

Genee

xpressionstu

dies

[68]

Streptom

ycescellu

lolyticus

Extracellularc

ellulases

Cellulose

decompo

sition

[141]

Streptom

ycesdrozdowicz

iiEn

doglucanase

Successfu

lapp

licationin

detergentand

textile

processin

g[14

2]Streptom

ycesmalachitofuscus

Streptom

ycesglo

meratusand

Streptom

ycesstram

ineus

Extracellularc

ellulases

Zone

ofhydrolysisin

platea

ssay

metho

d[109]

Streptom

ycesgancidicu

sEx

tracellularc

ellulaseC

MCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[109143]

Streptom

ycesactuosus

Endo

glucanase

Optim

izationof

CMCa

seprod

uctio

n[14

4]Streptom

ycesbobiliLIPIMC-A-

283

Streptom

ycesolivochrom

ogenesLIPIMC-A-

247

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[135]

Streptom

ycesglo

bosus

Streptom

ycesalan

osinicu

sCM

Caseend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]Streptom

ycesruber

Optim

izationstu

dies

Streptom

ycesvirid

obrunn

eusS

CPE-09

Endo

glucanase

Enzymep

rodu

ctionfro

magroindu

strialresidues

[145]

Streptom

ycesvirid

iochromogenes

AvicelaseCM

Caseand

totalcellulase

Saccharifi

catio

nof

rices

trawandethano

lprodu

ction

[146]

Streptom

ycesalbospinus

Extracellularc

ellulases

Degradatio

nof

cellu

losic

materials

[147]

Streptom

ycesgrise

orub

ens

Cellulases

Enzymep

rodu

ctionop

timization

[148]

Streptom

ycesmatensis

Endo

glucanasea

ndexoglucanase

Enzymep

rodu

ctionop

timization

[149]

Streptom

yceslongisp

ororub

erCa

rboxym

ethylcellulose

Prod

uctio

nandpu

rificatio

nof

enzyme

[150]

Cellu

lomonas

fmi

Cellulase

Activ

ityob

served

[151]

[152]

FPase(totalcellulases)

saccharifi

catio

nof

rices

traw

andethano

lprodu

ction

andop

timizationof

enzymep

rodu

ctionusingwheat

straw

[153]

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

8 Enzyme Research

Table1Con

tinued

Actin

omyceteisolate

Cellulolytic

enzyme

Observedresults

Reference

Cellu

lomonas

fimi

Cellu

lomonas

sp

Cellu

lomonas

sp(str

ains

ATCC

21399andJH

HY3

5)CM

CaseFPase

CMCdepo

lymerisa

tionandfilterp

aper

disin

tegration

[154]

Cellu

lomonas

cartae

Cellu

lomonas

cellu

lase

Cellu

lomonas

cellu

lase

Cellu

lomonas

flavigena

Cellu

lomonas

subalbus

Degradatio

nof

flaxsis

aland

cotto

nfib

res

Cellu

lomonas

uda

Cellu

lomonas

biazotea

Cellu

lomonas

gelid

a[155]

Cellu

lomonas

biazotea

FPaseendo

-120573-glucanase

and120573-glucosid

ase

endo

-and

exoglucanasesandcello

biases

Enzymep

rodu

ctionusingcellu

losic

substrates

Cellulosic

bioh

ydrogenprod

uctio

n[156]

[157]

Cellu

lomonas

cellu

lans

CMCa

seDegradatio

nof

flaxsis

aland

cotto

nfib

res

[155]

Cellu

lomonas

cellu

lans

NRR

LB4567

Stud

yof

substrateinh

ibition

kinetics

[158]

Cellu

lomonas

Endo

glucanaseCM

Case

Enzymep

rodu

ctionusingcassavab

agasse

[159]

cellu

lans

Isolationfro

msilverfi

shandcellu

lase

characteriz

ation

[160]

Intra

sporangium

Saccharopolyspora

StreptosporangiumR

hodococcusSaccharom

onospora

andNo

cardia

Microm

onospora

Extracellularc

ellulase

Zone

ofhydrolysisin

platea

ssay

metho

d[130]

Microm

onospora

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

Microm

onospora

chalcea

120573-glucosid

asea

ndCM

Case

Dem

onstr

ationof

activ

ity[161]

Enzymep

rodu

ction

[162]

Microbisp

orabispora

Microbisp

orasp

Microm

onospora

mela

nosporea

Microbisp

orabispora

120573-glucosid

ase

Cellulase

120573-glucosid

aseendo

glucanaseamp

Avicelase

Endo

glucanaseCM

Case

Saccharifi

catio

nof

rices

traw

Help

incompo

sting

Optim

izationof

cellu

lase

prod

uctio

nEn

zymep

rodu

ction

[77]

[132]

[110]

[163]

Thermom

onospora

sp

120573-glucosid

ase

Saccharifi

catio

nof

rices

traw

[77]

CMCa

seEn

zymep

rodu

ction

purifi

catio

nandcharacteriz

ation

[164

]Th

ermom

onospora

curvata

Extracellularc

ellulase

Prod

uctio

nandpartialpurificatio

nof

enzyme

[165]

Actin

oplanesm

inutisp

orangius

LIPIMC-A-

269and279

CMCa

seend

oglucanase

CMCa

seprod

uctio

n[137]

Streptoverticillium

morookaense

Nocardiopsisaegyptia

CMCa

seend

oglucanase

Zone

ofhydrolysisin

platea

ssay

metho

d[14

3]

Pseudonocardiathermophila

120573-glucosid

asea

ndCM

Case

Activ

ities

demon

strated

[161]

Actin

opolyspora

halophila

CMCa

seand120573-glucosid

ase

Detectio

nof

activ

ity[166]

Thermoactinom

ycessp

Endo

glucanase120573-glucosid

aseandavicelase

Degradatio

nof

microcrystalline

cellu

lose

[167]

Thermoactinom

ycesspstrain

TA3

Extracellularc

ellulase

Higheffi

ciency

andbioactivity

observed

durin

gcompo

siting

[168]

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 9

mesophila Amncolata autotirophica and Micromonosporasp have shown significant activities against lignin relatedcompounds [115 116] Pasti et al [117] have shown lignolyticactivity in Streptomyces chromofuscus Streptomyces diastati-cus and Streptomyces rochei Several actinomycetes such asStreptomyces coelicolor Streptomyces griseus and Nocardiaand several strains of Streptomvces sp isolated from termiteAmitermes hastatus have indicated production of laccaseslignin peroxidases or manganese peroxidases enzymes bythem [118] Laccase and lignin peroxidase activities have alsobeen observed in Streptomyces cinnamomeus [119] Laccaseenzyme studies were carried out including their structuralelucidation in Streptomyces lavendulae Streptomyces psam-moticus Streptomyces ipomoeae and Streptomyces sviceus[61] In a study by Escudero et al [120] Tsukamurella andCellulosimicrobium actinomycetes showed ABTS (221015840-azino-bis-3-ethylbenzothiazoline-6-sulphonic acid) oxidation rateof 108UL and 056UL respectively [120] Submergedfermentation in marine actinomycete Streptomyces lydicusisolated from Egypt Red Sea grown in medium supple-mented with peanut shell produced 1625UmL laccaseunder optimized conditions [121]Thermoalkali stable laccasefrom Thermobifida fusca has shown oxidation of several dyeintermediates including 26-dimethylphenylalanine and p-aminophenol [64] Streptomyces sviceus was also found tobe showing lignolytic activity [63] Streptomyces psammoticushas shown enhanced production of laccase under solidstate fermentation conditions in the presence of pyrogallolinducer which was taken to the level of scale-up studiesusing a packed bed bioreactor [122] Study by Niladevi andPrema [123] has shown production of all three enzymes thatis laccase manganese peroxidase and lignin peroxidase byStreptomyces psammoticus Actinomycete Rhodococcus ruberhas shown oxidation and degradation of polyethylene as aresult of laccase production by it [124] A study by Aoyamaet al [125] demonstrated laccase production by Streptomycesatratus In search of the genes involved in lignocellulosedegradation during composting of agricultural wastes two-domain laccase-like multicopper oxidase genes were iden-tified in Streptomyces violaceusniger [62] Rhodococcus jostiiwas found to produce lignin peroxidases capable of modi-fying lignin [126] Several other studies have shown lignindegradation ability in many other actinomycetes includingStreptomyces flavovirens [127] Streptomyces setoni [128] Acti-nomadura spp [55] and Streptomyces thermoviolaceus [129]

6 Future Prospects

Owing to the abundance and renewability of lignocellulosicbiomass it is considered as most appropriate and econom-ical feedstock for production of various industrially usefulproducts Lignocellulases enzymes are therefore critical inprocesses associated with bioconversion of lignocellulosesPresently most of the commercially exploited lignocellulasesrely on fungal or bacterial microorganisms Actinomycetesare relatively less explored for their biomass hydrolysispotential The studies can be elaborated in search of newactinomycetes producing lignocellulose degrading enzymesystemsDifferent feedstock shows variation in their chemical

composition The production of enzymes needs to be opti-mized for different biomass The production of lignocellu-lases from all microbial sources is still quite expensiveEfforts can be made for reducing the cost of productionof these enzymes using high potency actinomycete enzymesystems with broader range of tolerance and active at diverseenvironmental conditions Genetic engineering techniquescan be used to construct enzyme systems with desirablecharacteristics Also the studies can be expanded gradually toscale up to the industrial levels for their subsequent adoptionin commercial processes

7 Conclusion

Actinomycetes are an important source of lignocellulosehydrolysing enzymes They constitute considerable propor-tion of the soil or aquatic microflora responsible for biomassdegradation in nature The research studies on search oflignocellulose hydrolysing actinomycetes revealed the abun-dance and diversity of these microbes in different ecologicalniches The genetic and protein studies on their hydrolyticenzymes lead to the elucidation of structural and mecha-nism details of enzymes and their relatedness with otherknown lignocellulose producers and their enzyme systemsRelatively scanty information is available on lignocellulolyticactinomycetes The research studies therefore need to beelaborated in view of utilization of lignocellulolytic potentialof actinomycetes applicable in different industrial sectors

Conflict of Interests

The authors declare that there is no conflict of interestsregarding publication of this review article

References

[1] H S Chaudhary B Soni A R Shrivastava and S ShrivastavaldquoDiversity and versatility of actinomycetes and its role in antibi-otic productionrdquo Journal of Applied Pharmaceutical Science vol3 no 8 supplement 1 pp S83ndashS94 2013

[2] L S H Jeffrey ldquoIsolation characterization and identificationof actinomycetes from agriculture soils at Semongok SarawakrdquoAfrican Journal of Biotechnology vol 7 no 20 pp 3700ndash37052008

[3] A Das K Hamedani M Soudbakhsh K Prashanthi S Bhat-tacharya and S Suryan ldquoEnzymatic screening antibacterialpotential and molecular characterization of Streptomyces iso-lated from Wayanad District in Kerala Indiardquo InternationalJournal of Pharma and Bio Sciences vol 2012 no 2 pp 201ndash2102012

[4] P Das R Solanki and M Khanna ldquoIsolation and screening ofcellulolytic actinomycetes from diverse habitatsrdquo InternationalJournal of Advanced Biotechnology and Research vol l5 no 3pp 438ndash451 2014

[5] M Veiga A Esparis and J Fabregas ldquoIsolation of cellulolyticactinomycetes from marine sedimentsrdquo Applied and Environ-mental Microbiology vol 46 no 1 pp 286ndash287 1983

[6] D B Wilson ldquoBiochemistry and genetics of actinomycete cel-lulasesrdquo Critical Reviews in Biotechnology vol 12 no 1-2 pp45ndash63 1992

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

10 Enzyme Research

[7] D Prakash N Nawani M Prakash et al ldquoActinomycetes arepertory of green catalysts with a potential revenue resourcerdquoBioMed Research International vol 2013 Article ID 264020 8pages 2013

[8] G Y S Mtui ldquoLignocellulolytic enzymes from tropical fungitypes substrates and applicationsrdquo Scientific Research andEssays vol 7 no 15 pp 1544ndash1555 2012

[9] F H Isikgor and C R Becer ldquoLignocellulosic biomass a sus-tainable platform for the production of bio-based chemicals andpolymersrdquo Polymer Chemistry vol 6 no 25 pp 4497ndash45592015

[10] A Limayem and S C Ricke ldquoLignocellulosic biomass for bio-ethanol production current perspectives potential issues andfuture prospectsrdquo Progress in Energy and Combustion Sciencevol 38 no 4 pp 449ndash467 2012

[11] D Deswal A Sharma R Gupta and R C Kuhad ldquoApplicationof lignocellulolytic enzymes produced under solid state cultiva-tion conditionsrdquo Bioresource Technology vol 115 pp 249ndash2542012

[12] P Gupta K Samant and A Sahu ldquoIsolation of cellulose-degrading bacteria and determination of their cellulolyticpotentialrdquo International Journal of Microbiology vol 2012Article ID 578925 5 pages 2012

[13] R K Sukumaran R R Singhania and A Pandey ldquoMicrobialcellulasesmdashproduction applications and challengesrdquo Journal ofScientific amp Industrial Research vol 64 no 11 pp 832ndash8442005

[14] H Soni and N Kango ldquoHemicellulases in lignocellulose bio-technology recent patentsrdquo Recent Patents on Biotechnologyvol 7 no 3 pp 207ndash218 2013

[15] A M Abdel-Hamid J O Solbiati and I K O Cann ldquoInsightsinto lignin degradation and its potential industrial applicationsrdquoAdvances in Applied Microbiology vol 82 pp 1ndash28 2013

[16] W R de-Souza ldquoMicrobial degradation of lignocellulosic bio-massrdquo in Sustainable Degradation of Lignocellulosic BiomassmdashTechniques Applications and Commercialization A ChandelEd InTech 2013

[17] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 3 no 2 pp 145ndash163 1987

[18] A J McCarthy and S T Williams ldquoActinomycetes as agents ofbiodegradation in the environmentmdasha reviewrdquo Gene vol 115no 1-2 pp 189ndash192 1992

[19] R Kumar K Biswas V Soalnki P Kumar and A TarafdarldquoActinomycetes potential bioresource for human welfare areviewrdquo Research Journal of Chemical and Environmental Sci-ences vol 2 no 3 pp 5ndash16 2014

[20] T Vetrovsky K T Steffen and P Baldrian ldquoPotential of comet-abolic transformation of polysaccharides and lignin in lignocel-lulose by soil Actinobacteriardquo PLoSONE vol 9 no 2 Article IDe89108 2014

[21] L J Jonsson B Alriksson and N-O Nilvebrant ldquoBioconver-sion of lignocellulose inhibitors and detoxificationrdquoBiotechnol-ogy for Biofuels vol 6 no 16 pp 1ndash10 2013

[22] G Brodeur E Yau K Badal J Collier K B Ramachandran andS Ramakrishnan ldquoChemical and physicochemical pretreat-ment of lignocellulosic biomass a reviewrdquo Enzyme Researchvol 2011 Article ID 787532 17 pages 2011

[23] J Perez J Munoz-Dorado T de La Rubia and J MartınezldquoBiodegradation and biological treatments of cellulose hemi-cellulose and lignin an overviewrdquo International Microbiologyvol 5 no 2 pp 53ndash63 2002

[24] H V Scheller and P Ulvskov ldquoHemicellulosesrdquo Annual Reviewof Plant Biology vol 61 pp 263ndash289 2010

[25] V B Agbor N Cicek R Sparling A Berlin and D B LevinldquoBiomass pretreatment fundamentals toward applicationrdquo Bio-technology Advances vol 29 no 6 pp 675ndash685 2011

[26] J Shokri and K Adibkia ldquoApplication of cellulose and cellu-lose derivatives in pharmaceutical industriesrdquo in CellulosemdashMedical Pharmaceutical and Electronic Applications T G MVan De Ven Ed InTech Rijeka Croatia 2013

[27] DDhingraMMichaelH Rajput andR T Patil ldquoDietary fibrein foods a reviewrdquo Journal of Food Science and Technology vol49 no 3 pp 255ndash266 2012

[28] C M Gallaher J Munion R Hesslink Jr J Wise and D DGallaher ldquoCholesterol reduction by glucomannan and chitosanis mediated by changes in cholesterol absorption and bile acidand fat excretion in ratsrdquo Journal of Nutrition vol 130 no 11pp 2753ndash2759 2000

[29] FMGırio C Fonseca F Carvalheiro L C Duarte SMarquesand R Bogel-Łukasik ldquoHemicelluloses for fuel ethanol areviewrdquo Bioresource Technology vol 101 no 13 pp 4775ndash48002010

[30] J H Lora and W G Glasser ldquoRecent industrial applicationsof lignin a sustainable alternative to nonrenewable materialsrdquoJournal of Polymers and the Environment vol 10 no 1 pp 39ndash48 2002

[31] Suhas P J M Carrott and M M L Ribeiro Carrott ldquoLigninmdashfrom natural adsorbent to activated carbon a reviewrdquo Biore-source Technology vol 98 no 12 pp 2301ndash2312 2007

[32] F Le Digabel and L Averous ldquoEffects of lignin content onthe properties of lignocellulose-based biocompositesrdquoCarbohy-drate Polymers vol 66 no 4 pp 537ndash545 2006

[33] D Stewart ldquoLignin as a base material for materials applicationschemistry application and economicsrdquo Industrial Crops andProducts vol 27 no 2 pp 202ndash207 2008

[34] A B Albadarin A H Al-Muhtaseb N A Al-laqtah G MWalker S J Allen and M N M Ahmad ldquoBiosorption of toxicchromium from aqueous phase by lignin mechanism effect ofother metal ions and saltsrdquo Chemical Engineering Journal vol169 no 1ndash3 pp 20ndash30 2011

[35] G Dorez L Ferry R Sonnier A Taguet and J-M Lopez-Cuesta ldquoEffect of cellulose hemicellulose and lignin contentson pyrolysis and combustion of natural fibersrdquo Journal ofAnalytical and Applied Pyrolysis vol 107 pp 323ndash331 2014

[36] E M G del-Pulgar and A Saadeddin ldquoThe cellulolytic systemofThermobifida fuscardquo Critical Reviews inMicrobiology vol 40no 3 pp 236ndash247 2014

[37] S Sadhu and T K Maiti ldquoCellulase production by bacteria areviewrdquo British Microbiology Research Journal vol 3 no 3 pp235ndash258 2013

[38] R C Kuhad R Gupta and A Singh ldquoMicrobial cellulases andtheir industrial applicationsrdquoEnzymeResearch vol 2011 ArticleID 280696 10 pages 2011

[39] L R Lynd P J Weimer W H Van Zyl and I S PretoriusldquoMicrobial cellulose utilization fundamentals and biotechnol-ogyrdquoMicrobiology andMolecular Biology Reviews vol 66 no 3pp 506ndash577 2002

[40] A Lykidis KMavromatis N Ivanova et al ldquoGenome sequenceand analysis of the soil cellulolytic actinomycete ThermobifidafuscaYXrdquo Journal of Bacteriology vol 189 no 6 pp 2477ndash24862007

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 11

[41] D B Wilson ldquoStudies of Thermobifida fusca plant cell walldegrading enzymesrdquoThe Chemical Record vol 4 no 2 pp 72ndash82 2004

[42] D B Wilson ldquoProcessive and nonprocessive cellulases for bio-fuel productionmdashlessons from bacterial genomes and struc-tural analysisrdquo Applied Microbiology and Biotechnology vol 93no 2 pp 497ndash502 2012

[43] P Tomme A J Warren R C Miller Jr D G Kilburn and NR Gilkes ldquoCellulose-binding domains classification and prop-ertiesrdquo in Enzymatic Degradation of Insoluble CarbohydratesJ N Saddler and M H Penner Eds pp 142ndash163 AmericanChemical Society Washington DC USA 1995

[44] K Posta E Beki D B Wilson J Kukolya and L HornokldquoCloning characterization and phylogenetic relationships ofcel5B a new endoglucanase encoding gene from Thermobifidafuscardquo Journal of Basic Microbiology vol 44 no 5 pp 383ndash3992004

[45] D B Wilson and M Kostylev ldquoCellulase processivityrdquoMethodsin Molecular Biology vol 908 pp 93ndash99 2012

[46] M R Christopherson G Suen S Bramhacharya et al ldquoThegenome sequences of Cellulomonas fimi and lsquoCellvibrio gilvusrsquoreveal the cellulolytic strategies of two facultative anaerobestransfer of lsquoCellvibrio gilvusrsquo to the genus Cellulomonas andproposal of Cellulomonas gilvus sp novrdquo PLoS ONE vol 8 no1 Article ID e53954 2013

[47] I Anderson B Abt A Lykidis H-P Klenk N Kyrpides andN Ivanova ldquoGenomics of aerobic cellulose utilization systemsin actinobacteriardquo PLoS ONE vol 7 no 6 Article ID e39331 pp1ndash10 2012

[48] T E Takasuka A J Book G R Lewin C R Currie and B GFox ldquoAerobic deconstruction of cellulosic biomass by an insect-associated Streptomycesrdquo Scientific Reports vol 3 article 10302013

[49] E Lin and D B Wilson ldquoRegulation of 120573-14-endoglucanasesynthesis inThermomonospora fuscardquoApplied and Environmen-tal Microbiology vol 53 no 6 pp 1352ndash1357 1987

[50] N A Spiridonov and D B Wilson ldquoA celR mutation affectingtranscription of cellulase genes in Thermobifida fuscardquo Journalof Bacteriology vol 182 no 1 pp 252ndash255 2000

[51] S S Adav C S Ng M Arulmani and S K Sze ldquoQuantitativeiTRAQ secretome analysis of cellulolytic Thermobifida fuscardquoJournal of Proteome Research vol 9 no 6 pp 3016ndash3024 2010

[52] H J Gilbert ldquoThe biochemistry and structural biology of plantcell wall deconstructionrdquo Plant Physiology vol 153 no 2 pp444ndash455 2010

[53] B Henrissat ldquoA classification of glycosyl hydrolases based onamino acid sequence similaritiesrdquoBiochemical Journal vol 280no 2 pp 309ndash316 1991

[54] M Ishaque and D Kluepfel ldquoCellulase complex of a mesophilicStreptomyces strainrdquo Canadian Journal of Microbiology vol 26no 2 pp 183ndash189 1980

[55] M G Mason A S Ball B J Reeder G Silkstone P Nichollsand M T Wilson ldquoExtracellular heme peroxidases in actino-mycetes a case ofmistaken identityrdquoApplied andEnvironmentalMicrobiology vol 67 no 10 pp 4512ndash4519 2001

[56] J Placido and S Capareda ldquoLigninolytic enzymes a biotechno-logical alternative for bioethanol productionrdquo Bioresources andBioprocessing vol 2 no 23 pp 1ndash12 2015

[57] V Madhavi and S S Lele ldquoLaccase properties and applica-tionsrdquo BioResources vol 4 no 4 pp 1694ndash1717 2009

[58] A B Fisher and S S Fong ldquoLignin biodegradation and indus-trial implicationsrdquoAIMSBioengineering vol 1 no 2 pp 92ndash1122014

[59] D Sirim F Wagner L Wang R D Schmid and J PleissldquoThe laccase engineering database a classification and analysissystem for laccases and relatedmulticopper oxidasesrdquoDatabasevol 2011 Article ID bar006 7 pages 2011

[60] B Valderrama P Oliver A Medrano-Soto and R Vazquez-Duhalt ldquoEvolutionary and structural diversity of fungal lac-casesrdquo Antonie van Leeuwenhoek vol 84 no 4 pp 289ndash2992003

[61] T A R Fernandes W B da Silveira F M L Passos and T DZucchi ldquoLaccases fromActinobacteriamdashwhat we have andwhatto expectrdquo Advances in Microbiology vol 4 no 6 pp 285ndash2962014

[62] L Lu G Zeng C Fan et al ldquoDiversity of two-domain laccase-like multicopper oxidase genes in Streptomyces spp identifica-tion of genes potentially involved in extracellular activities andlignocellulose degradation during composting of agriculturalwasterdquo Applied and Environmental Microbiology vol 80 no 11pp 3305ndash3314 2014

[63] M Gunne and V B Urlacher ldquoCharacterization of the alkalinelaccase Ssl1 from Streptomyces sviceus with unusual propertiesdiscovered by genomeminingrdquo PLoS ONE vol 7 no 12 ArticleID e52360 pp 1ndash8 2012

[64] C-Y Chen Y-C Huang C-M Wei M Meng W-H Liuand C-H Yang ldquoProperties of the newly isolated extracellularthermo-alkali-stable laccase from thermophilic actinomycetesThermobifida fusca and its application in dye intermediatesoxidationrdquo AMB Express vol 3 article 49 9 pages 2013

[65] J Gao andWWakarchuk ldquoCharacterization of five 120573-glycosidehydrolases from Cellulomonas fimi ATCC 484rdquo Journal ofBacteriology vol 196 no 23 pp 4103ndash4110 2014

[66] A Amore O Pepe V Ventorino L Birolo C Giangrande andV Faraco ldquoCloning and recombinant expression of a cellulasefrom the cellulolytic strain Streptomyces sp G12 isolated fromcompostrdquoMicrobial Cell Factories vol 11 no 164 pp 1ndash12 2012

[67] H Schrempf and S Walter ldquoThe cellulolytic system of Strep-tomyces reticulirdquo International Journal of Biological Macro-molecules vol 17 no 6 pp 353ndash355 1995

[68] S Walter and H Schrempf ldquoStudies of Streptomyces reticuli cel-1 (cellulase) gene expression in Streptomyces strains Escherichiacoli and Bacillus subtilisrdquo Applied and Environmental Microbi-ology vol 61 no 2 pp 487ndash494 1995

[69] J-F Ethier S Harpin C Girard C Beaulieu C V Deryand R Brzezinski ldquoCloning of two xylanase genes from thenewly isolated actinomycete Actinomadura sp strain FC7 andcharacterization of the gene productsrdquo Canadian Journal ofMicrobiology vol 40 no 5 pp 362ndash368 1994

[70] N Li P Shi P Yang et al ldquoCloning expression and char-acterization of a new Streptomyces sp S27 xylanase for whichxylobiose is the main hydrolysis productrdquo Applied Biochemistryand Biotechnology vol 159 no 2 pp 521ndash531 2009

[71] E Dube F Shareck Y Hurtubise C Daneault and M Beaure-gard ldquoHomologous cloning expression and characterisation ofa laccase from Streptomyces coelicolor and enzymatic decolouri-sation of an indigo dyerdquo Applied Microbiology amp Biotechnologyvol 79 no 4 pp 597ndash603 2008

[72] T Suzuki K Endo M Ito H Tsujibo K Miyamoto and YInamori ldquoA thermostable laccase from Streptomyces lavendulaeREN-7 purification characterization nucleotide sequence and

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

12 Enzyme Research

expressionrdquo Bioscience Biotechnology and Biochemistry vol 67no 10 pp 2167ndash2175 2003

[73] Z M Wang B H Bleakley D L Crawford G Hertel and FRafii ldquoCloning and expression of a lignin peroxidase gene fromStreptomyces viridosporus in Streptomyces lividansrdquo Journal ofBiotechnology vol 13 no 2-3 pp 131ndash144 1990

[74] M E Boroujeni A Das K Prashanthi S Suryan and S Bhat-tacharya ldquoEnzymatic screening and random amplified poly-morphic DNA fingerprinting of soil Streptomycetes isolatedfrom Wayanad District in Kerala Indiardquo Journal of BiologicalSciences vol 12 no 1 pp 43ndash50 2012

[75] I Kusakabie M Kawaguchi T Yasui and T Kobayashi ldquoPurifi-cation and some properties of extracellular xylanase fromStreptomyces sp E-86rdquo Nippon Nogei Kagaku Kaishi vol 51 no7 pp 429ndash437 1977

[76] M Tuncer A Kuru M Isikli N Sahin and F G CelenkldquoOptimization of extracellular endoxylanase endoglucanaseand peroxidase production by Streptomyces sp F2621 isolatedin Turkeyrdquo Journal of Applied Microbiology vol 97 no 4 pp783ndash791 2004

[77] A S Ball and A J McCarthy ldquoSaccharification of straw byactinomycete enzymesrdquo Journal of General Microbiology vol134 pp 2139ndash2147 1988

[78] Q K Beg B Bhushan M Kapoor and G S HoondalldquoEnhanced production of a thermostable xylanase from Strep-tomyces sp QG- 11-3 and its application in biobleaching ofeucalyptus kraft pulprdquo Enzyme and Microbial Technology vol27 no 7 pp 459ndash466 2000

[79] A Maryandani ldquoCharacterization of xylanase from Strepto-myces sp strain C1-3rdquoHAYATI Journal of Biosciences vol 14 no3 pp 115ndash118 2007

[80] P Sharma and B K Bajaj ldquoProduction and partial character-ization of alkali-tolerant xylanase from an alkalophilic Strepto-myces sp CD3rdquo Journal of Scientific and Industrial Research vol64 no 9 pp 688ndash697 2005

[81] B K Bajaj and N P Singh ldquoProduction of xylanase from analkali tolerant Streptomyces sp 7b under solid-state fermenta-tion its purification and characterizationrdquo Applied Biochem-istry and Biotechnology vol 162 no 6 pp 1804ndash1818 2010

[82] L Thomas A Joseph M Arumugam and A Pandey ldquoPro-duction purification characterization and over-expression ofxylanases from actinomycetesrdquo Indian Journal of ExperimentalBiology vol 51 no 11 pp 875ndash884 2013

[83] L Thomas R Sindhu and A Pandey ldquoIdentification and char-acterization of a highly alkaline and thermotolerant novelxylanase from Streptomyces sprdquo Biologia vol 68 no 6 pp1022ndash1027 2013

[84] Z A Nagieb M G E Milegy H M R Faat and K B IsisldquoEffect of EDTA on production of xylanase by Streptomycesspecies and their bleaching effect on papyrus and cotton stalkpulprdquo International Journal of Forestry andWood Science vol 1no 1 pp 2ndash9 2014

[85] B S Priya T Stalin and K Selvam ldquoEfficient utilization ofxylanase and lipase producing thermophilic marine actino-mycetes (Streptomyces albus and Streptomyces hygroscopicus) inthe production of ecofriendly alternative energy from wasterdquoAfrican Journal of Biotechnology vol 11 no 78 pp 14320ndash143252012

[86] S Ninawe R Lal and R C Kuhad ldquoIsolation of three xylanase-producing strains of actinomycetes and their identificationusing molecular methodsrdquo Current Microbiology vol 53 no 3pp 178ndash182 2006

[87] S Ninawe M Kapoor and R C Kuhad ldquoPurification andcharacterization of extracellular xylanase from Streptomycescyaneus SN32rdquo Bioresource Technology vol 99 no 5 pp 1252ndash1258 2008

[88] H Tsujibo M Kosaka S Ikenishi T Sato K Miyamotoand Y Inamori ldquoMolecular characterization of a high-affinityxylobiose transporter of Streptomyces thermoviolaceusOPC-520and its transcriptional regulationrdquo Journal of Bacteriology vol186 no 4 pp 1029ndash1037 2004

[89] L S H Jeffrey A N Norzaimawati andH Rosnah ldquoPrescreen-ing of bioactivities from actinomycetes isolated from forestpeat soil of Sarawakrdquo Journal of Tropical Agriculture and FoodScience vol 39 no 2 pp 245ndash253 2011

[90] K Padmavathi M Thiyagarajan N N Ahamed and T Pal-vannan ldquoProduction optimization and partial purification ofxylanase from streptomyces coelicolor using agriculture wasterdquoInternational Journal of Chemical and Pharmaceutical Sciencesvol 2 no 1 pp 18ndash24 2011

[91] H M Rifaat Z A Nagieb and Y M Ahmed ldquoProduction ofxylanases by Streptomyces species and their bleaching effect onrice straw pulprdquo Applied Ecology and Environmental Researchvol 4 no 1 pp 151ndash160 2005

[92] H J Bhosale S R Sukalkar S M Z Uzma and T A KadamldquoProduction of xylanase by Streptomyces rameus grown onagricultural wastesrdquo Biotechnology Bioinformatics and Bioengi-neering vol 1 no 4 pp 505ndash512 2011

[93] A C Grabski and T W Jeffries ldquoProduction purification andcharacterization of 120573-14-endoxylanse of Streptomyces roseiscle-roticusrdquo Applied and Environmental Microbiology vol 57 pp987ndash992 1991

[94] A C Grabski I T Forrester R Patel and T W Jeffries ldquoChar-acterization and N-terminal amino acid sequences of 120573-(1ndash4)endoxylanases from Streptomycesroseiscleroticus purifica-tion incorporating a bioprocessing agentrdquo Protein Expressionand Purification vol 4 no 2 pp 120ndash129 1993

[95] J C Roberts A J McCarthy N J Flynn and P Broda ldquoMod-ification of paper properties by the pretreatment of pulp withSaccharomonospora viridis xylanaserdquo Enzyme and MicrobialTechnology vol 12 no 3 pp 210ndash213 1990

[96] M S Abdel-Aziz F N Talkhan M Fadel A A AbouZied andA S Abdel-Razik ldquoImprovement of xylanase production fromStreptomyces pseudogriseolus via UV mutagenesisrdquo AustralianJournal of Basic andApplied Sciences vol 5 no 5 pp 1045ndash10502011

[97] A J McCarthy E Peace and P Broda ldquoStudies on the extra-cellular xylanase activity of some thermophilic actinomycetesrdquoApplied Microbiology and Biotechnology vol 21 no 3-4 pp238ndash244 1985

[98] A Boondaeng S Tokuyama and V Kitpreechavanich ldquoXylan-ase fromanovel strain ofMicrobispora siemensisDMKUA245Tenzyme production and characterizationrdquo in Proceedings of the49th Kasetsart University Annual Conference vol 7 pp 308ndash315Kasetsart University February 2011

[99] S Berens H Kaspari and J-H Klemme ldquoPurification andcharacterization of two different xylanases from the ther-mophilic actinomycete Microtetraspora flexuosa SIIXrdquo Antonievan Leeuwenhoek vol 69 no 3 pp 235ndash241 1996

[100] L C L Fernandez J Rodrigez J Soliveri J L Copa-Patina MJ Perez-Leblic and M E Arias ldquoThe effect of culture mediaon the production of xylan-degrading enzymes by StreptomyceschattanoogensisUAH 23rdquo Journal of Basic Microbiology vol 35pp 405ndash412 1995

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 13

[101] C L Lopez-Fernandez J Rodrıguez A S Ball J L Copa-Patino M I Perez-Leblic and M E Arias ldquoApplication of theaffinity binding of xylanases to oat-spelt xylan in the purifica-tion of endoxylanase CM-2 from Streptomyces chattanoogensisCECT 3336rdquo Applied Microbiology and Biotechnology vol 50no 2 pp 284ndash287 1998

[102] S Khurana M Kapoor S Gupta and R C Kuhad ldquoStatisticaloptimization of alkaline xylanase production from Streptomycesviolaceoruber under submerged fermentation using responsesurface methodologyrdquo Indian Journal of Microbiology vol 47no 2 pp 144ndash152 2007

[103] U Kohli P NigamD Singh andK Chaudhary ldquoThermostablealkalophilic and cellulase free xylanase production by Ther-moactinomyces thalophilus subgroup Crdquo Enzyme and MicrobialTechnology vol 28 no 7-8 pp 606ndash610 2001

[104] D L Ristroph and A E Humphrey ldquoKinetic characterizationof the extracellular xylanases ofThermomonospora sprdquo Biotech-nology and Bioengineering vol 27 no 6 pp 832ndash836 1985

[105] J-H Shin J-H Choi O-S Lee et al ldquoThermostable xylanasefrom Streptomyces thermocyaneoviolaceus for optimal produc-tion of xylooligosaccharidesrdquo Biotechnology and BioprocessEngineering vol 14 no 4 pp 391ndash399 2009

[106] D Kluepfel F Shareck F Mondou and R Morosoli ldquoChar-acterization of cellulase and xylanase activities of Streptomyceslividansrdquo Applied Microbiology and Biotechnology vol 24 no 3pp 230ndash234 1986

[107] R Morosoli J-L Bertrand F Mondou F Shareck and DKluepfel ldquoPurification and properties of a xylanase from Strep-tomyces lividansrdquo Biochemical Journal vol 239 no 3 pp 587ndash592 1986

[108] M J Hernandez-Coronado M Hernandez F Centenera M IPerez-Leblic A S Ball and M E Arias ldquoChemical characteri-zation and spectroscopic analysis of the solubilization productsfrom wheat straw produced by Streptomyces strains grown insolid-state fermentationrdquoMicrobiology vol 143 no 4 pp 1359ndash1367 1997

[109] L S H Jeffrey and M R Azrizal ldquoScreening for cellulaseactivities in actinomycetes isolated from different locations ofPeninsular Malaysiardquo Journal of Tropical Agriculture and FoodScience vol 35 no 1 pp 153ndash157 2007

[110] W H van Zyl ldquoA study of the cellulases produced by threemesophilic actinomycetes grown on bagasse as substraterdquo Bio-technology and Bioengineering vol 27 no 9 pp 1367ndash1373 1985

[111] T A R Fernandes W B da Silveira F M L Passos and TD Zucchi ldquoOligonucleotide primers for specific detection ofactinobacterial laccases from superfamilies I and Krdquo AntonieVan Leeuwenhoek vol 106 no 2 pp 391ndash398 2014

[112] M E Arias M Arenas J Rodrıguez J Soliveri A S Ball andM Hernandez ldquoKraft pulp biobleaching and mediated oxida-tion of a nonphenolic substrate by laccase from Streptomycescyaneus CECT 3335rdquo Applied and Environmental Microbiologyvol 69 no 4 pp 1953ndash1958 2003

[113] C Ruttimann D Seelenfreund and R Vicuna ldquoMetabolismof low molecular weight lignin-related compounds by Strepto-myces viridosporusT7ArdquoEnzyme andMicrobial Technology vol9 no 9 pp 526ndash530 1987

[114] M Ramachandra D L Crawford andGHertel ldquoCharacteriza-tion of an extracellular lignin peroxidase of the lignocellulolyticactinomycete Streptomyces viridosporusrdquo Applied and Environ-mental Microbiology vol 54 no 12 pp 3057ndash3063 1988

[115] A S Ball W B Betts and A J McCarthy ldquoDegradation oflignin-related compounds by actinomycetesrdquoApplied and Envi-ronmental Microbiology vol 55 no 6 pp 1642ndash1644 1989

[116] B Godden A S Ball P Helvenstein A J McCarthy and MJ Penninckx ldquoTowards elucidation of the lignin degradationpathway in actinomycetesrdquo Journal of GeneralMicrobiology vol138 no 11 pp 2441ndash2448 1992

[117] M B Pasti A L Pometto III M P Nuti and D L CrawfordldquoLignin-solubilizing ability of actinomycetes isolated from ter-mite (Termitidae) gutrdquo Applied and Environmental Microbiol-ogy vol 56 no 7 pp 2213ndash2218 1990

[118] M L Roes-Hill J Rohland and S Burton ldquoActinobacteriaisolated from termite guts as a source of novel oxidativeenzymesrdquo Antonie van Leeuwenhoek vol 100 no 4 pp 589ndash605 2011

[119] D Jing and J Wang ldquoControlling the simultaneous productionof laccase and lignin peroxidase from Streptomyces cinnamo-mensis by medium formulationrdquo Biotechnology for Biofuels vol5 article 15 2012

[120] E L R Escudero O D S Daza and J H Torrs ldquoCharacteri-zation of lignocellulose degrading rare actinobacteria demon-stration of laccase activity in two isolates ofTsukamurella sp andCellulosimicrobium sprdquo Revista Colombiana de Biotecnologıavol 14 no 2 pp 70ndash80 2012

[121] M G Mahmoud H M Rifaat O H El Sayed F M El-Beihand M S Selim ldquoEffect of inducers and process parameterson laccase production by locally isolated marine Streptomyceslydicus from Red Sea Egyptrdquo International Journal of ChemTechResearch vol 5 no 1 pp 15ndash23 2013

[122] K N Niladevi P S Sheejadevi and P Prema ldquoStrategies forenhancing laccase yield from Streptomyces psammoticus andits role in mediator-based decolorization of azo dyesrdquo AppliedBiochemistry and Biotechnology vol 151 no 1 pp 9ndash19 2008

[123] K N Niladevi and P Prema ldquoMangrove Actinomycetes as thesource of ligninolytic enzymesrdquo Actinomycetologica vol 19 no2 pp 40ndash47 2005

[124] M Santo R Weitsman and A Sivan ldquoThe role of the copper-binding enzymemdashlaccasemdashin the biodegradation of polyethy-lene by the actinomycete Rhodococcus ruberrdquo International Bio-deterioration amp Biodegradation vol 84 pp 204ndash210 2013

[125] A Aoyama K Yamada Y Suzuki Y Kato K Nagai and RKurane ldquoNewly-isolated laccase high productivity Streptomycessp grown in cedar powder as the sole carbon sourcerdquo Interna-tional Journal of Waste Resources vol 4 no 2 pp 1ndash5 2014

[126] C Strachan D VanInsberghe and D Williams ldquoLigninaseactivity is not consistently predicted by the presence of man-ganese coordinating residues in dyp-like proteinsrdquo Journal ofExperimental Microbiology and Immunology vol 16 pp 66ndash722012

[127] J B Sutherland R A Blanchette D L Crawford and A LPometto III ldquoBreakdown of Douglas-fir phloem by a ligno-cellulose-degrading Streptomycesrdquo Current Microbiology vol 2no 2 pp 123ndash126 1979

[128] A L Pometto and D L Crawford ldquoCatabolic fate of Strep-tomyces viridosporus T7Amdashproduced acid-precipitable poly-meric lignin upon incubation with ligninolytic Streptomycesspecies and Phanerochaete chrysosporiumrdquo Applied and Envi-ronmental Microbiology vol 51 no 1 pp 171ndash179 1986

[129] M Iqbal D K Mercer P G G Miller and A J McCarthyldquoThermostable extracellular peroxidases from Streptomycesthermoviolaceusrdquo Microbiology vol 140 no 6 pp 1457ndash14651994

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

14 Enzyme Research

[130] R M Gulve and A M Deshmukh ldquoEnzymatic activity of acti-nomycetes isolated from marine sedimentsrdquo Recent Research inScience and Technology vol 3 no 5 pp 80ndash83 2011

[131] P Chellapandi and H M Jani ldquoProduction of endoglucanaseby the native strains of Streptomyces isolates in submerged fer-mentationrdquo Brazilian Journal of Microbiology vol 39 no 1 pp122ndash127 2008

[132] M F Eida T Nagaoka J Wasaki and K Kouno ldquoIsolation andcharacterization of cellulose-decomposing bacteria inhabitingsawdust and coffee residue compostsrdquo Microbes and Environ-ments vol 27 no 3 pp 226ndash233 2012

[133] A M Al-Tai S A Rzzak S S Al-Attiyah and B A Abdul-Nour ldquoCellulase production from actinomycetes isolated fromIraq soils II cell growth and cellulase activity of Streptomycessp strain AT7 at different temperaturesrdquo Journal of IslamicAcademy of Sciences vol 2 no 3 pp 185ndash188 1989

[134] L T A S Semedo R C Gomes E P S Bon R M A SoaresL F Linhares and R R R Coelho ldquoEndocellulase and exocel-lulase activities of two Streptomyces strains isolated from a forestsoilrdquo Applied Biochemistry and Biotechnology vol 84ndash86 no 1pp 267ndash276 2000

[135] A Nurkanto ldquoCellulolytic activities of actinomycetes isolatedfrom soil rhizosphere of Waigeo Raja Ampat West PapuardquoJournal of Tanah Tropicals vol 14 no 3 pp 239ndash244 2009

[136] H-D Jang and K-S Chang ldquoThermostable cellulases fromStreptomyces sp scale-up production in a 50-l fermenterrdquoBiotechnology Letters vol 27 no 4 pp 239ndash242 2005

[137] R K Rathnan and M Ambili ldquoCellulase enzyme productionby Streptomyces Sp using fruit waste as substraterdquo AustralianJournal of Basic and Applied Sciences vol 5 no 12 pp 1114ndash11182011

[138] A J McCarthy ldquoLignocellulose-degrading actinomycetesrdquoFEMS Microbiology Letters vol 46 no 2 pp 145ndash163 1987

[139] R K Harchand and S Singh ldquoCatabolite repression of cellu-lase biosynthesis in Streptomyces albaduncusrdquo Journal of BasicMicrobiology vol 34 no 6 pp 371ndash378 1994

[140] R K Harchand and S Singh ldquoCharacterization of cellulasecomplex of Streptomyces albaduncusrdquo Journal of Basic Microbi-ology vol 37 no 2 pp 93ndash103 1997

[141] X Li ldquoStreptomyces cellulolyticus sp nov a new cellulolyticmember of the genus Streptomycesrdquo International Journal ofSystematic Bacteriology vol 47 no 2 pp 443ndash445 1997

[142] A L G de Lima R P d Nascimento E P da Silva Bon and RR R Coelho ldquoStreptomyces drozdowiczii cellulase productionusing agro-industrial by-products and its potential use inthe detergent and textile industriesrdquo Enzyme and MicrobialTechnology vol 37 no 2 pp 272ndash277 2005

[143] N A El-Sersy H Abd-Elnaby G M Abou-Elela H A HIbrahim and N M K El-Toukhy ldquoOptimization economiza-tion and characterization of cellulase produced by marineStreptomyces ruberrdquo African Journal of Biotechnology vol 9 no38 pp 6355ndash6364 2010

[144] M Murugan M Srinivasan K Sivakumar M K Sahu and LKannan ldquoCharacterization of an actinomycete isolated from theestuarine finfish Mugil cephalus Lin (1758) and its optimiza-tion for cellulase productionrdquo Journal of Scientific IndustrialResearch vol 66 no 5 pp 388ndash393 2007

[145] F N M Da Vinha M P Gravina-Oliveira M N Franco et alldquoCellulase production by Streptomyces viridobrunneus SCPE-09 using lignocellulosic biomass as inducer substraterdquo AppliedBiochemistry amp Biotechnology vol 164 no 3 pp 256ndash267 2011

[146] N E-A El-Naggar A A Sherief and S S Hamza ldquoCellulolyticStreptomyces viridiochromogenes under solid-state fermenta-tion conditions for bioethanol productionrdquo African Journal ofBiotechnology vol 10 no 56 pp 11998ndash12011 2011

[147] P Prasad S Bedi and T Singh ldquoIn vitro cellulose rich organicmaterial degradation by cellulolytic Streptomyces albospinus(MTCC 8768)rdquo Malaysian Journal of Microbiology vol 8 no3 pp 164ndash169 2012

[148] P Prasad T Singh and S Bedi ldquoCharacterization of the cel-lulolytic enzymeproduced by Streptomyces griseorubens (Acces-sion No AB184139) isolated from Indian soilrdquo Journal of KingSaud UniversitymdashScience vol 25 no 3 pp 245ndash250 2013

[149] P Prasad Tanuja and S Bedi ldquoCharacterization of a novelthermophilic cellulase producing strain Streptomyces matensisstrain St-5rdquo International Journal of Current Microbiology andApplied Sciences vol 3 no 3 pp 74ndash88 2014

[150] MAMYassien A AM Jiman-Fatani andH Z Asfour ldquoPro-duction purification and characterization of cellulase fromStreptomyces sprdquo African Journal of Microbiology Research vol8 no 4 pp 348ndash354 2014

[151] M L Langsford N R Gilkes WWWakarchuk D G KilburnR C Miller Jr and R A J Warren ldquoThe cellulase system ofCellulomonas fimirdquo Journal of GeneralMicrobiology vol 130 no6 pp 1367ndash1376 1984

[152] A K Srivastava and P Agrawal ldquoCellulose hydrolysis by Cellu-lomonas fimi and ethanol production by Zymomonas mobilisrdquoJournal of Atoms and Molecules vol 2 no 2 pp 214ndash222 2012

[153] S B R Ali R Muthuvelayudham and T ViruthagirildquoEnhanced production of cellulase from agro-industrial res-idues by optimization of medium components using centralcomposite designrdquoAsian Journal of Food andAgro-Industry vol6 no 3 pp 113ndash131 2013

[154] D W Thayer S V Lowther and J G Phillips ldquoCellulolyticactivities of strains of the genus Cellulomonasrdquo InternationalJournal of Systematic Bacteriology vol 34 no 4 pp 432ndash4381984

[155] D Lednicka J Mergaert M C Cnockaert and J SwingsldquoIsolation and identification of cellulolytic bacteria involvedin the degradation of natural cellulosic fibresrdquo Systematic andApplied Microbiology vol 23 no 2 pp 292ndash299 2000

[156] M I Rajoka and K A Malik ldquoCellulase production by Cellu-lomonas biazotea cultured in media containing different cellu-losic substratesrdquo Bioresource Technology vol 59 no 1 pp 21ndash271997

[157] G D Saratale R G Saratale Y-C Lo and J-S Chang ldquoMul-ticomponent cellulase production by Cellulomonas biazoteaNCIM-2550 and its applications for cellulosic biohydrogenproductionrdquo Biotechnology Progress vol 26 no 2 pp 406ndash4162010

[158] R Agarwal B Mahanty and V Venkata Dasu ldquoModellinggrowth of Cellulomonas cellulans NRRL B 4567 under sub-strate inhibition during Cellulase productionrdquo Chemical andBiochemical Engineering Quarterly vol 22 no 2 pp 213ndash2182009

[159] K R Sugumaran S P Chakravarthi and V Ponnusami ldquoCas-sava bagasse a potential and low cost substrate for cellulaseproduction in an economical fermentationrdquo Research Journal ofPharmaceutical Biological and Chemical Sciences vol 4 no 2pp 1168ndash1175 2013

[160] J Premkumar T Sudhakar and K Srikiran ldquoIsolation andidentification of Cellulomonas cellulans from silver fish and

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009

Enzyme Research 15

characterization of cellulase enzymerdquo Journal of Chemical andPharmaceutical Research vol 7 no 1 pp 346ndash349 2015

[161] MMalfait B Godden andM J Penninckx ldquoGrowth and cellu-lase production ofMicromonospora chalcae and Pseudonocardiathermophilardquo Annals of Microbiology vol 135 no 1 pp 79ndash891984

[162] J Gallagher A Winters N Barron L McHale and A PMcHale ldquoProduction of cellulase and 120573-glucosidase activityduring growth of the actinomyceteMicromonospora chalcae oncellulose-containingmediardquo Biotechnology Letters vol 18 no 5pp 537ndash540 1996

[163] C R Waldron Jr C A Becker-vallone and D E EveleighldquoIsolation and characterization of a cellulolytic actinomyceteMicrobispora bisporardquo Applied Microbiology amp Biotechnologyvol 24 no 6 pp 477ndash486 1986

[164] S P George A Ahmad and M B Rao ldquoStudies on carbox-ymethyl cellulase produced by an alkalothermophilic actino-myceterdquo Bioresource Technology vol 77 no 2 pp 171ndash175 2001

[165] F J Stutzenberger ldquoCellulolytic activity of Thermomonosporacurvata optimal assay conditions partial purification andproduct of the cellulaserdquoAppliedMicrobiology vol 24 no 1 pp83ndash90 1972

[166] K G Johnson P H Lanthier and M B Gochnauer ldquoStud-ies of two strains of Actinopolyspora halophila an extremelyhalophilic actinomyceterdquo Archives of Microbiology vol 143 no4 pp 370ndash378 1986

[167] B G R Hagerdal J D Ferchale and E K Pye ldquoCellulolyticenzyme system ofThermoactinomyces sp grown on microcrys-talline celluloserdquo Applied and Environmental Microbiology vol36 no 4 pp 606ndash612 1978

[168] C-C Chang C-C Ng C-Y Wang and Y-T Shyu ldquoActivity ofcellulase from Thermoactinomycetes and Bacillus spp isolatedfrom Brassica waste compostrdquo Scientia Agricola vol 66 no 3pp 304ndash308 2009