ecology, physiology, and biochemistry 1502 of blue-green algae

8/9/2019 ECOLOGY, PHYSIOLOGY, AND BIOCHEMISTRY 1502 OF BLUE-GREEN ALGAE

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Al

ECOLOGY, PHYSIOOG AND BIOCEMSTR OF UE-GREE LGE12

D Institute of Main Resorces, Univerty of Califonia San Diego

ola aliorn

CONENS

INRODUON HOOSYNHESS . ROEN SSMLAON ESPRAON

OOOOCC C ..................................... CHEMAL COMOSON NERAL URON VAMNS AN GROWH USANES YROEN ION EQUREMENS CELLULAR FERENAON CELL OEMENS OX EABOL AORS OLERANE O H AN w EMERAURES AN O ESAON USE O VRUSES

LUEGREEN LAE N MO ELAONSHS

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n the ontnuum o ie om te vis patiles though the uneuamooganisms and nal to highe pants and anals the Canophtauegeen agae oup a unique poton As was desed n Lag8) the bluegen agae hae a poaoti ellua oganiaton whihs sla to that found n batea. he de fo atea howeve nthat the ae geeal obigate otoautotohs obtag the abon andeneg photosntet meanss w ae sla to tose found n

ghe ants. In euaot ogansms vaous sees of metao eatonsma e loaed wtn the e n speaed oganees Studes wth tohonda 1 2 and oopasts ave demonstated an odeed aao enes n se subunts on the onsttuent ebanes o these oganeles Although the membanous stutues and ganues n els o bluegeen agae an oen e eated to spe etao untons the ae notsepaate o the suoundng topam a ontnuou memae n

1 The survey f te literature peraining t ts eview was concuded n Jnua6

Supported by Atmic Energ 'Cmmission Ctract N. A(-)Ge

Annu.Rev.Microbiol.1968.22:4

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pe o h ack o compamenazao he buegeen aga ce apale o all eaoc eacon demanded o a phooauooph oganmand gow and pe a ae opaable o hoe obeed n eucayoagae One o he a eeah pobe aoaed wh poayo e heeoe he ocalaon o enye packe whn he cel and he elula mehan ha enue een unonng o meabol pahwaywhch may be n cloe pomy Thu he enzyme concened wh phooynhe epaon and nogen aon n blue-geen agae ae al locaed on he aeae hyaod bu a ye hae no een epaaed byany hya o hemca mean. Recen edence om eeeech ocopy doe dcae he eence o ubun n he ameae o ue-geencell 5

Bue-geen agae ae wdey dbued n paay a haba whh

w uo e ncudng ho in dee ane ae he Ancand o d up pond hey ae o ondeabe ecooga mpo

ance n egd o nceang he ey o eeal and aqua enonmen and o he acy a olonze o newy epoed land uaceoe o he ehwae pee howee gow n uch pouon ha heyeu n eou loe o economc and aehe eouce Many o heecoogcay poan aec o bue-geen agae ae dued n he oowng eon ogeh wh phyoogca and bohea ondeaonInomaon on ohe apec whh hae been omed fo h eew

may be ound eewhee 11 Ahough he bue-geen agae compe awde aebage o ee-ng and ymboc om he buk o aboaoynegaon hae been done wh ony a ew dozen pece A he peechoen o aoaoy ude ae eneay hoe whch a eay o u

ae pole ha many deaon o he genealaon peenedn h eew may mege om uue ude on ome o he eenowngenea

Thee a lage aebage o non phooynhe ogan whh aybe ondeed "ooe buegeen agae and ncluded n e CyanophyaSuch apohooc yanophya gh ncude he Febaea chemoyn

he o uch a B and hhz nd heeooph om ucha Oc. The phylogene ane o hee aed goup hae ny been dued by Soano & Lewn 1 4 ) o o hee apochoocom ae groued wh he phooynhec buegeen aae pedomnaeyon ophologcal lae u ae que dn on phyoogal eaTh eew onened only wth the phooynthec yanophya.

SSS

The phooynhe mehanm n bue-geen agae baaly ma o

ha n hghe pan phooynhe n ha wae he umae hydogendono and oecua oygen beaed. A ew an hae been hown oe abe o educe wh unde anaeo condon n he gh 5The a phoochemcal and chemal eacon a deemned o een

Annu.

Rev.Microbiol.1968.22:4770.

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BLU-GRN AGA

oosss ) so occu bug g 18 ) usr s covicig vic for occurrc of bo ooss og wvg) sor wvg) d for bo ccic d occic ooosorio ) bcri oosic rus squ d vig ir oosic gms vig c o o oorco (20).

ss of bsoro scr d svr mocufoms of coo x bugr g c 21) s fomr biv o b o r fuco oo sm I ccorig o cur o mi fucio of ccssor is obiirois ) s o bob g qu d rsfr rg o cv coro ocus vovd oo ssm II rsuig bo of d rsg o o cro so c s

rd v co sor c o oosm w cocomformo of r s vidc for ricio of soquo oc cocoe eeco eo s ss of cos bw oossms I d II. ccssoigms r r o ss for Hi rcio cvi ) orfo ooosorio 8 )

gms vovd oo ssm r coro d morcoro como dsg P-70 ) oocos of ss vov cc ooosoro wc sus formo of d occic ooosoio wic sus i ormo obo P sog duc s duc wic is biv o bfrdoxi c uc NP rsc of m frrox-NP-rucs 2, ) ducio of NP b rogs Fj Ms nn clndc ) s osoubo of ss omos c-fr rros o bu-gcs rsus ucoug of bo occc d ccc ooosoo ocic ooosorio c b rsord b ioof kow comouds o wsd m bu cc ooosoo ss o qui som soubiid co wic dos o

o b ow cood i co sor ssm o coros) iggs s ow buk of coco C54 is si wsdo m bu sm mo of i s rm mbs is o si rmov uc ig bou fos of cro rso ors bvd o b of g fucio sgcci oorcios of m ) O cro crirs v b foud i bu-gr g d m ro i oosc mcsm r ooroquo d vi 4) sccocrom -rucig subsc ) vrous ii 7 )

s bsoio xi of ccssor oosic igms rri r ro os of coro- bugr g r goo xm orgiss for sud of comc do wrb

Annu

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5 HA

cncentrtin iments is rtiy cntre by te wvent n intensiy f e bsbe it (21, 442), n s by temerture (8)Gs Gvnjee, n stues n te ecency f enery rnsfer fryccynin clry cels wit wiey vryin rs fyccynn/lry, e sue t e cry- n syse is ily urescen wile n syse is wely urescen (43). Jne Myers (21) ve swn rw f n re li resus in rtc ecese in e cryl cntenwile nt ecin te cncentrtin f yccynin Tey inerree isien terin s cntr mecnism t bnce te tw i rectins syntesis e cncentrtin f yccynin ws nt wever ece by rwin te ces n mncrtic i f 620 ; tistey secue mi be ue t enery trnsfer frm yccynin irectly

t syste in iin s usu uncinin in syste Psynec incrrn f CO n uereen le s een swn

t ccur remintey vi te Cvin cycle invlvin crxytin f ribuse ise (5) e CO xin y s ccur vi suc recins s e crbyl e synese recin (46), te reucive crbxyic c cyce s escribe fr se ceri (47 n y crbxytin senyruvte (48) ere is n evience wever tt tecrbxyin recins in tis lter ru re invve in te i-inucessimitin f by ces bue-reen e. me tese recins

re invve in rkxtin retis weer, s te cmuns wicincrre rcrn frm CO rn rn re reinny crbxylic cs n sscite in cis, ncluin ctrulline 4950 ere re rers in e ierure t uereen le cn live nerclly v cesynetc ecnis uner nurl cniins (5eiter te iiy t rw nerbicly nr te ity ve cesyneicly s ever been exerienly ensre wi ny

blue-reen l

Buereen e cn ssimite nitren rm nitrte nitrite (52), mni (3) yrymine () ure ( ) csein () min cs(57), uric ci (58) n N (59). ere is cnsierbe vriin mnsecies s t wic frms f niren cn be ssime, but ue-reene renty cn rw wit nitrte s e sle ntren surce Te reucin f nitrte rcees ru nitrite t mmni Nitrte reucse,wic s Mcninin rtein is un in e ricute frcn cllfree rerins f b dr, weres nitrie reucse isrecvere in te sube fctin (60) Ferrexin n re requirer nre reucin we ferren n NADP re reuire r nri reucin 61). As tese reucns re ville ru e recins system I tsyntesis e reuctn f nitrte in inc cells isi-stimute. Sme uereens s ssess yrense wic ctyes

Annu.Rev.Microbiol.1968.22:4770.

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BLUE-GREEN ALGAE

e reio o ire o mmi i i w s e dre d6 e roese me is oed e melle b is esi sbiie eoe reme

e ili o ree molelr iroe is o i m lmesrms 63, b s o bee reor or ielr seies o rder Croooles 5 e imre o iroe-xi e ormiee sil wer erii s eel ee smmie bewr e xi N is rir imr i e lii ree exse sres d i mli reservir reded r ire mds wi is esseil r e eslsme mes eri le d ier ls 4 5 S lii is imor res dee ie b vi ivi r exessive rdiio s we s i eser res sd es d d exsed

b reei liers 6 e siie N xi b mrie bleree le s o bee deel ssessed l ere re m sec h bhc f h c whch c 8 e ee rms ssoied wi N io re eer seieso mm r 3 For ms rii sdies N xie sbe isoo N s bee mlo b re i ws ie xio e olowe i s mor b deemiie mo redio elee eee 9 s e meold eie reire or e er meod re m simler se

reired r N sis is eie sd id osiderb i evli e ribi be-ree le e erili erresri i eviomes

ies wi i ells reveled e meism N xi ible-ee le d m similriies o d i sme ree-liviberi d i dles lemis rs 0 e ees vrisiibirs d e msii e ses se i exerimes wielree reris 1 ve ls demsred e bsi imilri ie meim o N i i ll mirorisms. e emes ivlvei i molelr iroe re led o e lmele 3 ; e

s e ve o bee sere rom e emes ivolve resiri osesis e rs se iermeie w s bee emosre i ese exerims is mmoi e iermeies bewee N N e o ee eee ( & o e sese ooemill roe roe oors re iie i N xi is s reel bee esioe Cx lo ere seems be ose il relisi ewee sesis re xi i els s wiesse b m ier re N xi i e is mred e rk ere des seem be bie em

l rei r e xi melr iroe e evidee r is is m be-ree e x N i e drk i slie wi eriri ssres l e re xi is ver w x8 s sow e re xi i e rk is simlr

Annu.





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52 HLMHNSEN

i he ight if CO s excluded Ths ow rate of xaton of N2 te lt he bsence of CO2 indictes the possble deendence of ntoe xationfor crbo compouds suppled by te phoosytc corporto of CO,Snce N xo in ceee pepatons is cceleaed by pyuve and isaccompaied by libraton f C2 (78), nd snce decaroxyao enzymeshave pevously been shown to be assocted wth he amelle ( he eegy fo N2 eductio my be dered from he phosphoroclastc cevgeof pyruv, s as een show fo h ateal system (80). Alhough hxato of N2 ca hs be dve by ght-depede elecron trspotsystems on sould no eclude te possbly n culturs expose ogh much r o he rqure reucg powr AP s urshe drectly b he phoosyhec pprus Vrous mnerl elmts such sMo M Co Fe nd Na ave been shown to have mked eects on te

ntoen xato process (59 71 81-3)

Vry ow of he rsprory pahys i uegre lae.The EmbdenMyehof ycolytc pathway appaently fucios n heseg s dos he enose hospe cyc (84 8 n h ss of noroaton attens om C-aee oanc states o on enymac analyses seveal nvestiatos hae suested that e carboyc acd cyceTA is vy suggsh or s no fuy opertive (86) The minevidece fom t big eperimes s rdocrbo from-ce is fo i ps four mno ids (guamc prine arnne d lece bu n n spartc cid hs dctes athe TCA cyce may be operatve up to he omaon of -keogutaric cid,whch s hen convted o i d he oth abeled io id cane obtaed from glutamc cd by h usual metabolic phways. uppofo hs hypothess s aorded by he fae o detec keoglutaate dehydogenase and ccnyCoA in beee cells (86 88) AceyCoA synhease cirate syhse conitase, iso-crate NAD dehydogease succnc deydoease and malc deydoenase hae l been demonsred

i -green gae (8 The ruent at e TA cye s no fncong i these ae mus e consideed eatve to he fact that he propres of some eyms h buegree ge r suciently derntfrom he correspong eymes higher pls th he ssy mosmay o be hs althoh cean eaos wee uable dmonste aldose n any o the blueee algae tested (89), t s nowknw th dos (ype II is pesen hese alge bt was no deected ealer becuse of usatsfactory assy methods 9-93

he enymes socate lyase and malae synthase, whch ae necessryfo the opeaton o he yoylc cycle, have been demonstraed n cells of

bue-geen alae houh they do no show the adpive beavo wch hecorespong zymes show n prepraons fom ge plants (4, ).The ec of he glyoxylc cycl s o synhes on suca moleule

Annu

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m w mu. bun n he enzm ue n ee un n n le d w n be ne y nmnn hue e nbl mne lbeled n blue-ne we ae w - b ne f leea e e paways y w aeae s assae aa e ase ay paa eye syse

Te o apo a oe espao aso appeas o eassoae w e aa We ow e ao eo eaos,e ose eys a e ea oase ese ogass Sea esgaos ae sggese a egee aga ae epee popoopospoyao o sppy os o a e A eqe y e eese speaos wee ase o e ae o AD oase (6

a o e asspo a s e a o o sese aeo o aeae (96 S a eee ew oeg e a oese agae o syese A T e a sees waae ew oe oowg pos Fs a ew ege s ae e sow o eapa o sow eoop gow op ass (6 9 soo a e ae a -geeag ea oe ah of popn end een hoe bueeen e whhe n b f eh owh e ble of un n omee dkn f mn mnh dun whh me e eumbl

enen ATP fm ellul mnen T e AT nen of s bh n n n dkn bu n of e ywh (HmHn unublh wh mb a fo oe agae a ae ee esgae 9 99

Aog S oo Sae (6 wee ae o ADHoase ns ndulns, hls pns o Glps lpl Wse Ha (00) osae e pese o s eye spes o apooo -ge agae ( aos ee oese as e-gee agae ey ae Feaeas e ADH oase ay assoae w e paae fon o ese ogass

sowe on o nhbo een fom a of gee agae age pas

oo Myes ae ese ee ee -pe ooes (549 a 2) ns ndulns (2 . e a oase eeeo aspo syse as o ee eoay eosae ses w os ae ae e og o yooe oase( oypeooase ay as o e eee (

aos ses a po o eao o e pooy apaoy a egee agae a away (13 sow

elle n f b vbs due end-,dhlenl n he l un dn fm he l f wewee n he d endenu n ube e he hoenono n he b f nhb ue h nue a e e me

Annu.





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4 HOLM-HNSN

electron trsport chins functon in both photosnthesis n resprton, rth htthet eet tt ee the tvt f the et ee tt te. Othe et hw tht the et

tke o oxen is eprese t ow it levels but be tted bright light (104 S) tiultion t high lght level compied by edg ee CO dt, ght e f hedt f whh the xded v the the te t edATP 105. This upte f O2 wthout concomitnt libcrton of CO

iht et f ft f ede v ecc hthht.

Hoo rr 84) hve shown that the lcerlehe3-phosphte ehoee n be-ee le ue eer NAD AD s ezeThe tte e tve te the ee tht t ght ftethe the et hhte e in the gt thw deeding on whc ee g the tive site

BG URYMt blue-reen le whch hve ben investigt re oblite h

tth, n tht the equie ih f ted gwth d etcn not grow hetrotrophicll A few e 06 107 of heterotrophic rowth n certn of these le, notl speces of osto hveben ted fte ht fttve heteth n tted t

he Cht t ex the ede f begee ge wte h dved g te The e et f hetet e f egee e hweve hw rowth rte in the dk de ew tht de tth dt d de edeteth oteti i n

Recentl there hve been severl new reports o the blt of ew spee f buegree t row ver slowl ete rkness f supplewith the proper crbon source A strn of Chorogo frtsh hs bnshown to grow contnuousl n the rk 76, though onl fter lon ptton pero In te eperments of F there ws no rowth of ths ldrng the rst onth n the r, n onl slow rowth urng the seconn thr onths. Toohrx teus hs lso been shown to grow in ther f ed wth ge d d 97) but not f te the e tge e The th et tht the heteth gwtw ve w ed tht tth dt d tht heth w t theed the dk. t t e if th g e o ntno gowth n the drk, ghtgown te we ed t ss which wee plce n the dk f jt 10 d efegwth w eed. Othe vetgt hve ted et rowth

blue-reen e when rst plce n rkness but tht subsequnt trnsersfed to show n heterotrophic rowth 108

he k f heteth gwth t ee f egee ge nth lck stiultion of ther respirtion upon tion o ornic sub

Annu.





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AA

a 9 a o b oad o b d o a o ogac b o a aa ba o cl 110. o do o aa o b d ow a cl- aao o

ay bl cll ow ll o o al o addd oga ba alo d w Clbld cood doa o c ca ala addd ba w ow l T Ca ac ow a w cll o . vb o -dln a ow w U--co o -aca o 3 c o clla d wg dd o labld oc ba T ooalao o lald aca l docabobg dbd o ld o o o ao acd gac acdag ol d lc ad o cabolc acd 96 T n alo ow ooalao o co da lco wad wa wa od o olaccd

boccal ba o obla ooaoo o ow oo a cly b ad a ol col coo CA cc o a abl o a A da adc o a col CA ccl abc o doabl olaa ddoa ac ad ac o adocabo aacacd cbao w Cac 8688 o dc dc o o clao 86 96 a bl- ala cao aAP da w o dod 86 co o ac

o acl-CoA ad c o -olac acd d lad coodad coao o ATP cll l ad da co o ao d a bl- aa a caabo a a o AP da

obl a l ala a dd o oooolao o b o T b a ca z o dak b ba oolao aco T o (86), wc bad o abl o doa AD oda ac ala acco o ow alao o oac cood ad al o co A w odao T a clao b ald awa o d aoo o awa alga

I I

doboclc acd o b aa o locazd a bad cl o do cod o coloca doabl cooo a d o caoc cl I aao cola & 114) a ow c o cob c o a DA bl a a c dcbd

occc o o DA bl vbs a o o da o aao o 10 o 5 da wc DA dlcad ad ddd bw dag cll a o b dd w blg lga I l ow

Annu.





10/24

HMHA

v h h ognon o h DN b ht ond n ob n Eshh o h b hooo no on o ng oo w-sranded D 116 17). A t dt to obtn

n dg o nhono dvon n bgn g no knownwhh th nh o DN o dng ho od ding he grwh o o during ot f he geneion ime as in obacteria 7) Th DA in ot is oxd wth htonwhh bv to b nvov n tng th tvt o h notnt o th DN o No hon hv bn ontt, howvn b o n bgn g 1 8 1 1)

n ohnv td o 29 n o bgn g dn ndowok ot tht bo n tt th N gdn howd nod dbon 120) Th w no D ie bands alhugh here was a smal saeie band pssibly aribuabe cnaminan polsacchads This is in cnras eucayoic ces inwhich h DN en how bod o od n o DNnt o dn gn ngton tng vng h ooon o h DN on n h n hoond tohond Th no bgn g xn hd kb b ooon o h DN bo n gnton) td om bont nt o . /m hb ooon o th DN n n b o h hoo

o) w h o vb ngng o 35 n o 71 nt G Oth invsgos hv ot fo Pom born tt DA band which had a ow G onnt han he main band DN 12 1 ) nd whh th d gh n DN od nh hoonh .

Th ognton o th geneic t n bgn g non gt n o on b dton o tn btn whh n ot om t wth von 221) h ot th n o bgn g d nont g xo o 2 d ov oho odo o 124) Mo bgn o b o vvg do o tvot don th on o vo h bntd to obtn b t o boto nvtgon 12512) K h hown ht ohn h no on h gowh o thagae ) ; his is undubed relaed he ac ha clchicine which inereres wih prper uncining he spinde in eucaryic ces has nocrrespnding t o ton n o Kumar has a caimedtton hn n bgn g t xo to ton ndnn b no whh h on o hoog

on 128) . n n h od obtnng on nt oyss dus b nt wth ntbo b gv no d on thhtt nd tbt o h tnt tn 12) .

h hv bn v w t o RN ooon nton n

Annu.





11/24

LU-GRN ALGA

bgr g. & Rchr hv rord h idu oof h olclod hohorl cv ocd wh h rboofrco 3 d h R olr ocd wh h D frc

o 31. oro&

oh hv dcrbd R fro . du- whch cv g 2O-h R 132 Th hv o rord cd R whch o b locd hr o or rh c wl of 133

CHEMAL OMPOTON

Th gro chc cooo of b-gr cl bcl r oh of croc lg Thogh vrol cor c grl h cooo of c c 134, 135 of r d

fr-wr c how h folowg cooo, bd o dr wgh:crbhdr, 30 o 55 r c; ro, 2 o 45 r c; ld, 1 rc; R, 2 r c; D, 04 o 0 r c; o g, .5 rc

Th o cd of h ro fro blgr lg do o drrcbl fro ho of o ohr croorg 13 13 Orh co o cr cclc old br 13 hb rord o h wrob rco o cr b-grlg 1 h r o h cood whch orh occr w o

frhr vgdTh bl o h co f cd blgr c co w 10 d 1 crbo o d r rd or hv o o dobbod M o h lo or o co d d rord f cd 043. cl cog -olc cd o o hcoo o b ocd h l 12 ch Wood1 hv rord h Su co rg q of"-oc cd ,,12-ocdcroc cd Thr o o gc o of brchd or f cd, ch r od bcr , . Th ohr coo ld of bgr g r

b ho of bcr ckg rol, ch, hohdlho,d hohdl-oo d rb ho of hghr l corgwo glco dgcrd, hoovo dglcrd, d hohdl-lcrol 14. rr 15 h rord h h hloogo fschi grow wh c h gh bo 10 r c of h cllr drwgh c b rcovrd o--hdrobr.

Thr codrb vro h occhrd hzd bblgr g d h coo oocchrd . h o-cld d gr, whch b org odc, r bvd o co

of olcchrd rld o glcog 14. bo 25 r c of hcrbohdr of Toohx u co of ordcg gcofrcwhch was l dd a fucofurnosl (,-fuctofurnol-2-glcorod 4. h rclllr olcchrd of

Annu.

Rev.Microbiol.1968.22:4770.Downloadedfromwww.annualreviews.org



12/24

HOLMHANSN

Anabena fosquae ml glu a yl u hl amu gluu a a b 148

Th ll all h h u la lg b h lama mmba a h hah a h hah bu h ha

n a m hh mb h variousbaa Th ll al f A. nidulan a 2 ua a a D & llz 19 Th ma m h abhya fa malu gala a u u ll all f Phrmidium uncnatuma gh u am a h u a ll a26amml a a muam a 0) Th la a gh h guam m b haa blug all 1 1 aly ma ll all m ha b yng

bya a osoc Th mma h l! al f glbula a muym mlul Th mu ayhh b b a b muh h g ll all a b u by h a lyzm a ll Th lau hah ha b b a f llu bl a au ma huh h m qu h val f h llul a h h hah matal hlzh ma maha u a gay glu a l h amu aab u a gluu a 148 1

a h uual gu ba ul a ua y a hym a ual h mhla bamhyamu a mhyly ha al b Pconma byanum )

Th ma hh m blu-g ala a hhll-a h hbl ha allha a hyh Th h hyb m ha b yal by Ha Fja IS; h hmh f hyya a allhya aal a ha h n vvo al a u h m gm h a ab ba a

70 m ha al b a blug alga bu ha b 1 18 Th a m mml u a a zaxah h mxahhy llaxah aaaha ahazhll yah a hyxya ka (9-6)

NERAL URION

a h ma ma lm Mg S, a K 1h llg lm a al b a a la ma ha all bl-g alga alum (3 63b 6 mlbum 6 6) maga 8 um 6 aba 8 1 1 qum a ha b ma h alga bu bau h ma u a

Annu

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13/24

BLUEREEN LAE

enzyme co-facors, i may be assumed that they are aso essentia microeements in these algae The functions of these microelemens ae basicalysimilar t se descried fm higher plant and anima studes (171 17ith the impliain f M, Mn Fe and C as in he prcess of nitrogenxatn anadium ha een ited t pay a role in nitrgen xation (73),ut the enea eet hh re tained by the addtn vanadium mght have een due t mydenum ntamnatng h vanadiumsat HlmHansen (6) ud nt nd any evidene fr eentaity fvanadium fr thr r petn There haeen n ent demnstatn V essentaly r -green agae.

VINS AND GROW USANES

here e only two reorts n he leatue o the auho's knolede

n which a amn equemen was demosaed o bueeen aleVan aaen (18) eoted ha eh 1 mane bueee seces heolaed equred he addtion o itamn B or rowh; Pntner & Proasol(174) aso epoed a iamin B requirement b ormm percnumone of the commonly cultued sans of blue-green reques an oanicadditons. he cobalt requiremet can howee be pata o whllsustiued for b the addition of vitamn B (8). The tamin cmpostinf these alae (other than that f B) has not een studied hough thereis n eason t eliee tha it is signicantly dierent from tat of other

algae and hgher lans.The authr is unaare of any repors on the occurence of natura

gowth sustances (auxns, kinetics etc) in pue cultures blue-greenalgae or on the eecs n celula growth elicted by the addiion o suchsusnces t te nutrent meium

HYRE QUIREES

Mst bu-green agae gr in envirnments whih are neutral t alaline th a fe speis uing n haitat wth a p eteen 50 t 0

(75) kaline hot spngs geneally hae aundant goth of these algaewheeas acid hot sprins (H 5 and ower) do no (8). n aboary cutre the p ptimum o groth seems o e eteen 75 to 100 butgrowth stl ccus at aues e 110 (3,4, 163 176) Te we Himit fo aboratory cultures is about 6. to 7.0 (54,57). Thee are no daaon the ntenal pH of blue-geen cels,nor s he author awae of any explanation as to wh these cels geerall seem require such alalne condiions. The p otima f extacted enmes rm them aparentl do not

3 e auxin acivty has eed n amples f ehwate bm

peie: ee Mo J A., 8 14955 (196) Ar pap prs slgh f nde-3e d n h rowh si e-een ae: Amd, M R, inte A. P 78 2619)

Annu.




14/24

6 HOLMHANSEN

i signicnty m ths ucytic cs t is ssi tht th highH s my chcstcs f h c w n mmnn thus cts cuts ith in th in utk y th cs in sst h nutin suin su ssni mits

CELLUL EENON

Vius scii stuctus ccu in t u-n such s htcysts knts, nss, n hmgni Litt is knwn cncningh cn f nn f sucus intn fm vtv ckns n nss n ucv sucus nsuviv th nism uin is unv wn cnitinsut th unin t hcyst is mu m scu t hs n sust th tcysts civ s stg vsics s uctiv cs,

in nin in in cnin intitin th s my vstigi stuctus (77 7) Wk (7 ) hs t gmintn f htcys, ut thi h yt t cm y h invstitsFy & sy () hv t h htcyst ns ntshw ny n ty u tht thy sw t sy hn th th cs h mn hs cnty n shwn n ) ht hcysts nhit th vmnt ny csint htcyss s ss cmmny un cn t htcysts itsms ht th intin cs is cnt sm tn y h

ctivity th cs in h mnhuh sm uths hv t sm g snchniwh y mniutin igt n tmtu 8) syncnus gwt ugn g hs nt n chiv t t in with ucytic L n c-wks hv utii ih wh sn Nostoc muscorum tin cutus whchn fm snus (s) s hcysc st(3 ) n s wn hthcy n h k cu vmn s ck h si s futh vmnt n h mnus s s inuc y iht in th 7 m scin f scum

Such hinuc mhic intiin cn vs y ihthvin wvngth twn m[ Lz hs sust thhyccynin is th c ign th htchmic civincss n h n m ms hycyhin nvv in hvs f hctivin (5) huh L & ishnic nt hi us s shwin htchmic uctin sm wthsusnc cssy mhic vmn ( ) this hs cntyn v y th vns f (5) inctin h hs n nnu nuctn f mhc vmn n h sn f

ny hchmc stmuus Nsto musorum cn cmt ts f cyc nth snc f iht u th is s vy sw css n ws nt ny t fw mnhs n k

Fy Kum & F 8) hv s sc sm f cyc

Annu.




15/24

G G 61

Chlorogloea fitscii involvng n nopor g nd a st nousta s mologcal stas a als ct y chng n tma compostn of t s 34) and n appaan an stutn t lama an vaus ytpasm anus (

un a a ny s as avn n sua stasaza & Vsna wv cam t av sn fusn wo srom in n N muscrum t fm n a (184).s svatns wc wu cnsttut t st vnc suaty nlu-n ala av nt n v y t nvstats ut vdnc o exuiy w cmd y uma n ass stus n cuir un aa t anoics (188) tanstans A ndu wc w sstant t t pnln stptmycn bu o t t anttcs upn mn t tw stans an tn

tm w two ks tan stans w w stant t tantts s appant ants u wt a uny aut n cs uma's wk cou n at wv yPik (189), wo coud d no vn f coinon n A uas

CE MOVEENTS

o grn aa ow gidn na, taty mvmntou y do o po a at an sa mcansmcausn ts mvmnt s nt unst tu vaus yptss avn ug wc nvlv xcron o mulanus mata ppaatn tm was f cntactn n t c mcansms asscatwt smss sua tnsn 51 Many t lantus sw sw mst ap vnts as sw t psn pstu t c wa ( tu wc stans mucnus sustancs mt r In so utus f amntus ms t mvmnts sut n aatn f f lamnts t m tan tufts ays u on an inrwning n sust a su

a u o ropic or in -owin stams as t pvnts amnts m n was away (1 ) ln mvmnts may

ct t twa away m a su t t tn pnn n t wavnt an ntnsty (12). tn spcta tvaus typs f ll mvmnts av n pt y sva nstats(3 ) t pmnts spnsl ts t-nun mvntsas t n t sam spcta ns as t ooic imnts

XIC EABLIC F

n o oo-roucing u-grn g oduc oxi cow a u mtaty m aima (15 tm Mcrysis rugis a yi poppd onsising f tn amna sus (1 ) . In tsts wt vus anmas ts ala put wasfa t tan t amn atan an stl ss tctan t psns tact m t musm Ami phid (

Annu.





16/24

62 OLMANN

Si (9) as rny rwd g n, inudng rmgrn aga, and srssd r pn appan n pygi a an as trpu gn

OLERANCE TO HIG AND w EMPEAUES AN TOIATN

Aug r ar m rpr f bugrn g grwing in prings a mprr v 9 C, rn a n a C may pp mpra mi f gw f n m mpii spi (, ). imia ai f t abiity gw a ua mpras rmains an igma. a trma ain aigy spiiz survi n ndns f g mpurs s ndid y wr tmprau ii f grwt in try t

rm srin wi t grw i brtry bw abu , wii mprur pm r grw i bwn 4 C (99 aiity s rma rains gw a u ig pratur indt tt uur an guain f ti pi mu nfstaiy f ts mrmu d mpu n nas pn n m nnma rganim wi ndg nauain w t . prty tr v b ui t in sua ppris f prin m mpii -gn aga an a fw ingan n t a saiy f ia nzym()

-g g r rmy y i rgr t urvivng gnray rius w tpratrs 1, ). ugrn s urs n pd n sur nd in r war in nrti, wi prt vr ii iti () t ny mprars rp s w a bu p vr iatinntns mps y w msp mdy y a as u tig ig nnsiis in summr mns, fwd by many mns an g inny in u, upd w wna mpau u n undn pa y awn and

fzing, n ring i ptimum gwing pi abay iwit bu-gr id rm rti m ti iiy urv qu n sw frzing as w ptd ys f frzing awing, a prss wi is vry dsruiv t mt uaryti s () ai f ti m biiy t uviv t f w mprm iy ri i bi rutur rgniztin uprins Lvit (4) as psua a rs rinss in pns td inr sabiit in ti prtn brug au b rsistan tw rin f rur iupi n n wr,sd prryi s in i insigains i w ining sy

irun f ufydry nd dsupid gp in t prins f ugrn aga if vitt' ty migt rat wit t u ran t ag t b w nd g mpatu I ud b pint

Annu.Rev.Microbiol.1968.22:4770.

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UE-GREE ALGA

ot tat not a -gn aga sow t aiity to svv sc tp cotos. any isoat ro t Unit Stats a t v g bg b vv v w ()

b -g g vv g p l k. b p av b to viae 87 o toag 05) aga wic svivng an tawng can aso srviv yoiiation 0) Eton croga o yoii grn cs o not rva any oroogicacangs as coar to conto scins wic wr not yoii0) s ay atia to t ack o any ao vacos nts aga So stains o s av own no cn n aty g v y g t yoii o in t aiity to sviv

o 0 to o. is i sa contrast to t station wt gn aga wic a mk vbl o cts an wos csc vv atn at 00 evn 0 ints 208 )

O ag vb ugaga as at to nvronnta acto as fzng an yng n t ic qattg t n of via cs n any ssn. l nt ca o i ti ga tvn t nica os can not cont in aga p b v. V as l tat cnt cincy lating o ing c vaio cocco -gnsci y incooatng cataas o FA t ntnt aga tact t tios ct o rois 0, 0 ts vay gat a o ci o ci owv an t gwt of a coony sng c is ao on trat an ot actos

EIIL O VUE

Snc aa & ors ) rst sci a virs cang y oa grn aga otr viss cyanoags ) av n soat wc

cuse lyi n flmentou bue-green lgae ( 197, 213 1 4) . The lytc i pg g g c p b g & g 1) v b o p b vl l k b v t ag l.W, L & W ( v vb b N v ema baum taotnactat is was t agap consia action oto actvat y ow gt conc at t oto activation rocss occr att ntranc o t vis into t aga c, a tat t vaai canis wa not ircty assocat wit otosyntss

coogca signcanc o cyanoags s no nown, tog ty

Annu.





18/24

-

may pay a oe in t uen eat an ecay o uegren cs constutng a war oo Suc a rap nraon o a arge popuon o cey o e cue y prsitic or aprphytc aceria. io ) a recny oae a nur o aceria wc cau ecopoio o vgtav u-grn cs oug roys wr no ac Tr apparny s o spccy nvov wn actra an aga ces ae ame bacrial iolate did no cue y of Chlorella ce

BE-GREEN AE N YO EATONH

Bugreen aga occur n mny yotc aonip uc as n cens 6) n t roos o cycads 17) and w cran bryopy anpteriope (59 In om o te reationip u-green aga soug o conru os or a o e euce caron an nirogen co

pouns requre or growt y o c. Drew & it 218 21 ) vrport at t Noo n a icn u ecrt up o pr cn ot nwy assiae caron n e or o guco wic i en convreno anno y e ungus Ts s n sarp conras o rvng ugrn aga wc cre ony sa ouns o ino acs ppi ororganc cs no nurn u 220) T Noo c n cn au ack hck geanou et a te evng or noray posss as en own eprinay a Noo cs, soryaer soon ro t cn au, connu o ecree c o ter

poosync rouc no te eium ut wtin 4 hours ty eveo ac sea an con o organc caron rese e ecanmswry t ungu semingy aters te strucure an inunces t abolc recon o te uegreen smbon i competey unknown. I s onrt n is rgar tat t ue-gren ymion (Anaban p) n twa rn U as no en cuture ous os su 177) wc ay rc so srucura o occa araons o apparny croscoc enon cen ungi comony requir uy oeognous viin B an ioin 221 ) i s iey ag syiontin cens ao srve a t source o viains an osiy oer growt

actors or e ungusI as bn ugget 222) a cloropass o igr n y

orgnad as ndpndn c wic nvadd an ece tie a intraceuar syonts evenuay ecoing moie to unction a organes o o ce Bcau of any imiarts ween choopaan ue-green agae, auhors who e suh spulaons on consdrt ry enoyon a bng ugren ce er conieraorpoogica 223 224 enyatic (225 an cca vinc 106 n avor o s possy atoug uc an vouonary veopn

o coropa rom ue-green ces wou nt mny eron in srucure an ceica copostion o e enoyion Ter av n anyus o pign pas cyans) in te apocoro gn agaGlauoy noohinam an n t crytomona yanophora paradoxa.

Annu.





19/24

G G

e cyaneles o bo ese oms appea o be sma o be-geen agaen eecon cogaps 223 22) e an sca eence beweene cyanelles an ee-lng be-geen cels s e absene n cyanees oan oe sea an cell wal only a n membane seaaes om sos I ese cyanelles o epesen be-geen algae wc ae los eseas an cel wals ey ae mos lely moe n meabolc nconsas wel o ey ae o a poe o be naabe o nepenen gwan vson wen emove om e os e 22

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21/24

BLUEGRN LGE 67

ako S Mikbly, 4209-1 19

4. Ghsh A K Govindje Bophs 6, 11-19 19

44 Kidel P Gibbs, M, Natue, 200

26061 (1963)45 Norrs, Nors R E lvn M xpt. Botany 6 4-74 19

Hlm-Hanen Brw, G \T JrPlant e Psiol (Tkyo) 4993 9

47 ns, M W. Buch B. B,An 1 Proc. Nt Si. 55 98-4 9)

48 lke, D. A, Eneaur 2 (19)

Hlm-Hnsn 0. Bass-m J A. Cv M J xpBoa, 8 47 97

O M V ann 0 lvnM erl 77 70-78 9 59

1 Ehln Sc Am, 4 748(9

Htti Pant Ce h(Toko) -9 19)

53 Glof G c. Ftzgl, G Sg A ! otny 9 2 9

4 Katz W A., Myes, J A. Bo 42 8-87 (955)

Cbb . D J. My J. m !.Boan 7- 94 len, M B Ar Mikool, ]

3-3 7 McLch J Gh an

Mrobl, - 9)8 V c. Mer J E,

Gen. Mrol., 47(96

5 twt, W D P. Ntrogen Fxa Pa T Alo U 18 pp.,)

o A. Ms J Pa CePo , 8 7-7(97

1 tt A. Myr, J PnPl 41 101- 19

Fj Y O o, APan el Phso ( 5,3054 (9)

Stewt W D Siene, 1584 197

4 hields L M, u WBa e 0 92- 8 9)

w W D Nu

6 (96) olmHn 0 Sece 19

090 197 Dugdl, R c Gg J

R . Lm Oceag9 071 194

Sw W D P Ba(Lnd 1 8-48 97

9 wo M J Bohm oAa 1 894 19

70 Bi, R Wilo, P W Da

z 2 )7 1 ur R A laP:'s, 1 -84 (19

7 ox R M Fy F G. E.Bohm Bo ta 88 080 194

7 Fy, ox, M., cm. Bhs Aa 1 -9 97

74 y P Fogg G E A Mbo., 4 (9

7 o R M, rch Mro 6190 (197

7 F P ! e Mobol 9 0 (19

77 Wanb A a Y aure, 214 738 97)

78 o R M A Mikrobiol., 53-7 9

79. Fy, P Cx R M Bh Bop Aa, 1 4-4 9

80 C J E Moteon EMowe F Csf J E. Bhm Bo Aa 44 -19

8 owe P Nos D D

Pla Pl 11 18 HolmHan , Geof G Co F Pl Paa 77 94

3 B Lanwrt h:90 7 94 )

8. ood W C, N G BomBo At 14 09- (197

8 hu W Gbb M PPl. 41 337 9

8 m A oon aR ! Ba 9 7-897

87 o D o S. MooeR B ! G Mobl, 9170 (97

88 a N G BI 10 45 P 7)

89 Fwo A. Idh M GbbM Pa P 40-62)

9 t N Pol. 8-897

91 Wl J M Schulmn M bb Ntu 20 9 9

9 Vn Blen C Na 206 9-9993 u W a- P

17 94 Kornb Kbs

r 179 98-9 97

Annu.R

ev.Microbiol.1968.22:4770.D

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22/24

HOLMHANSE

95. Per J ' N. , . Genirobo 49 -1 ( 1967

96. Hr, D S, Moore R Bhm . Bohys Ata, 109 6( 1 9 6 5 )

Kyohaa T. Fujta, Y HattorA aaa Ge. App.Mcbiol 76-82 960)

98 HoHanse, 0 Boot, C RLmnl Oceag II 5 10-1 (196)

99 Coob, Halc, P , HmHase 0, Vai B E, Erp/.Cl! R. 47 32-28 (1-67

00 Websr D. A. Hcket, D P.Plnt Phyiol 4 -( 1 966 )

101. Holton R W. s cience

4-5 1910. Wbt, G c Frl, A. W,

la Physiol. 2 -9 ( 95 3)03 arr N. G Hallwy M. J Ge

Mrool 3 3344 (19610 Brw A. H., Wbr, C A.

Boty 40 75358 5310 oh, G., Ows 0 Kk, . A rh.

op 171-0( 1 9 6 3 )

rdr Z oa 9 152( 1 9 1 )

07 Ais F Hvr, S R MiH J. Bot. G: 98 ( 1 9 3 7 )

108. Va Baal c Bt Mri 4299 (92

1 C G Exe G y Hlw M, ld, S, chBiochem. Biopys. 120 5 03-71 9 6 7)

1 10 Ktz, W. A M Phsil. 0 5-80 1 9 5 5 )

1 1 ar N Peae J. B 28 19)

1 Allson R K Skppr, , RidM hot W A Hg .C Bi Che. 20 197-05( 1 9 5 )

1 Kioaa . uj ia Y., Hattoi A Waanabe A J e. Al.Micl. 16568 (16)

1 1 R H Sngh, N, Bioph.Boc/ze. ol" 9 6-0 196 1

1 1 5. Lea, ! Ursuc. Res20 19020 (967)

1 1 b See 5 1707 66)

r J, Mo BoI 6 208-3

963)1 1 Zua G., \t M R J Bio

phs. ich y 5 1-5 1 )

1 9 . B T o Boo

of Devee 17 (foU Prs, N.Y, 165

10 Edelman M, Swi, D Sif, A" pte H Zl BBail Rv., 1 ( 1 9 6 7)

1 1 Kay A . aomon Fil Mo o. 2 7983 ( 1 9 6 7)

1 Shield L Dot Botany 49, 47-4 (16)

23 rtb:y, T, Vis, " Bio. Acd S Ing 1 1 -60 967

u M. P N' , 3( 1 9 6 6 )

15 Grf, G C. izd G k, F, Am 37,2168 ( 1 5 0

1 :6. Kumar, H. D An Boa, 3-33 ( 9 63 )

7 H n 313-2 ( 19)

umar, H , Exptl. oy, 530 16

9 V Bale c Sciece 9 0( 1 9 6 5 )

10. C Rh, G Ntforc 22 0-15 197)

1 1 Cu Rch, , Ntr-fosch. 22 876-8 ( 1 97)

Nr, , Rh, J oChe" 24 029-3 ( 9

33 t J J. S, omp Biochem Ph'i 36- 67

1 4 Holh P D M R B ,Bateri. O Abs 1196

1 3 . Par, T Sph K Srick-an, J. D H J Fisheie Re.Bo C. 18 1 0 0 1 1 ( 1 9 6

6 Willi, A. E i R H., . B, 9 30- 15

. Rho, G G V

Mkroolya 34 282 1965 Mller M W i Pzer of Mcol Metolite 33-97(GwHi! Book o, N. Y6

3 ma, 0 Lwi R A,h u 8 41823

1 0 Prk, P V Bl, aurer L, Siece 15 77(1967

41 . H, R W Be H. Hr, hytchemisry,

595-602 964)142. vi, nnz W J. oc

K Biochi Bioph. ct 877 964

43 Nh B. W Hais R as,

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LU-N ALG

T, Boem. Bopys Rn 20 252 19

144 Nichols, B. W Wd B J. BLp 3 - ( 19 8

1. Car, N G Bocm Bopy. ca

20 3081 (19)1. Gesy, R. M A J Bty 51

3889 (97 Tuu, Y Ymkw B

cm. (Tyo) 58 879 (98 oo B G T R. G

J Mrb 1 8-85 ( 96149 Dew G. , Glwtze W, Arc.

l. 5 798 (950 an Lfr M., arti

Bem. Bpys Res Commn 32225 92

1 HlmHen, d, RLn R A Pyc% g 5 1 5

12. Put rrk E Gen Mrobol., 11(9

53 Mo, B c Sn 4 887 (9

6ha, C, Chmy a B-hmy o n igmns 59. , T W, d d Ps NY 83 19

1 Ho A Fuj t, Y., Bom.y 6 33 (1959

1 apma D , le W m H W Bioh J 1059 (196

G, E B Pln yso 37339 (9

58. ovnj n C Rbnwch E 39-92( 1 9 1

15 9 Hezbeg Jense S L Ptmstry 5 -3 19

Hz S, Jn S . o 9

6 Heeg Jen L Pcmtry 6 96 19

2 Gw, T W dBomtry P Pgms4 (Gn, . W Am , Y 83 19 5

Gro G Fza G F m. J Btay 383- (

A M. B Aro, PaPyso 372

1 Th, M . Er Ar krobol -192

1 Natur 0 5 9 5 ) Wf nn.

18, 39-2 (198 HmHn A Sy of Mj

a E t qurmt t N utr/ o Buelge D s Un. fWn, Mn W 19

19 Al M B , D Pys

Platarum 8 5 955 oon, G V x ,

Plat so. 8525 (

Bowe, J. u Y 2 pp

72 m, A, Btley Ba M (d), m 0 pp198

13 Brel Arc. robol 1118 (19

r , ov L J GMrb 1 90 (58

1 , G a R 018- 95

lenn K. A, Bwn Eyse H B4 93 (19

n P 1 (

18 rh Pr Lean ocLndn 1 9 2 1 1 1 9 1

1 79 Wk C P. atur 205 2219

18 Fy Wby A E Ntur 20999 (19

8 Wlk C P, Proc Natl ad ScUS. 121 9

1 Fy . Kumar H D F G EJ G Mcobol 3 1-19

18 Lzo, S, Sc 0 ( 9

18 o V W Gn.cbl. 28, - (9

18 z N Pycl

( )8 L nc, W J r. 3- 9

8 7 Pet , n A . , ArcMrbl 7 8 (9

88 Km H D Nar 16 5 (

189 Pklek P. Ntr 5 (19

1 Pnkz Bowe, Am.J Boay 899 (

11 , R W Nar 21528-8 (9

92. Bndx S, n omarate B-cstry of Potreactve y 0 EAam K p9

Annu.




24/24

70 HMH

193 ult, W, Pnta 58 647-6(96)

194 Haldal Phh Phl 5-60 (67)

1 Fln

New Zealand Ve. I,14 1818 66)

16 Kons, H McKerchr P D Gom R Roerso A.wl, Cn 1 omp MeV 29 -8 96)

97 Sio M Bacteriol Rev, 31 0- 67)

18 ock T D, Sn U 02-9( 67)

1 ry J A Cholz, R atur 20 720- 64)

0 Mrr E Pysigy i

chmtr f 10(Lw R A, d Admice NY 99 pp 96)

20 Won B A. Brt Phol Bull.2 7778 (162)

0 HolmHs 0 Physl. l-am 300 163

203 HolmHn, 0 ycloga 4 3 964

4 v 1 Tre B 3 -9 96

Lpmn, ul BnClb 68 664-66 94)

06 HolmHnn 0 Btany 7- (964)

07 og T D ol V ESclicg H Jr Ph)'l3 16-6 (96)

8 HolmHnn, 0 Cryobilgy 41723 167

20 Vn Bln C 1 Phyc 1 1 96)

Mlr E V Bl fPhyol 0- (96)

V Bl C, Pyc., 3 147 (967)

2 M M E Sn 0 67-8

w R

H Vrly 66 67)

4 Sfmn R Morrs M E 1aerl, 7717 (6)

Sng R N, h K atur2 00- (97)

6 Amdji V Phc% ga 6 76 967)

217 b Kyo T Mralae an otynttaera 186 (Jn SoPln ysol Eds Unv Tokyors okyo 96

8 Drew E A Sm D g 7-88 (67)9 Drw E A Sm D C. New

Phtolois 66 900 967 on B A f n Mirl

40 11 (6)22 1 dn T HolHnn 0

Pla Cll Pyol (Tkyo) 90 94

Thor BI 4 74 (67

Hll W T us G Phyl3 71 1 67)

4 Rs H l, W ll 13-9 96) on D S, 9 1606

166 m S G n Bhmstry

Chlooplt I 99 Goow W E Amic PNY 776 67)

227 Hll T, Cl G Cll B9 6 (96)

El Bi Phol Bll, 3-9 967)

Annu.



ecology, physiology, and biochemistry 1502 of blue-green algae

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