heritability analysis and phenotypic characterization of ...oar.icrisat.org/10826/1/heritability...

Research ArticleHeritability Analysis and Phenotypic Characterization ofSpider Plant (Cleome gynandra L) for Yield

Ann Kangai Munene 1 Felister Nzuve1 Jane Ambuko 1 and Damaris Odeny2

1Department of Crop Science and Crop Protection University of Nairobi Kenya2The International Crops Research Institute for the Semi-Arid Tropics (ICRISAT) Nairobi Kenya

Correspondence should be addressed to Ann Kangai Munene wakaireanngmailcom

Received 30 November 2017 Revised 30 March 2018 Accepted 12 July 2018 Published 31 July 2018

Academic Editor Mumtaz Cheema

Copyright copy 2018 Ann Kangai Munene et al This is an open access article distributed under the Creative Commons AttributionLicense which permits unrestricted use distribution and reproduction in any medium provided the original work is properlycited

Knowledge on phenotypic diversity among existing spider plant accessions is amilestone in the improvement of spider plant whichis a highly nutritious indigenous vegetable in Kenya A study involving agronomic andmorphological characterization of 49 spiderplant accessions assembled from East and South Africa was carried out at the University of Nairobi Field Station for two seasonsin a randomized complete block design with three replications Phenotypic data was collected on growth habit flower petiole leafand stem colour petiole leaf and stem hairiness number of leaves per plant plant height number of primary branches leaf lengthand width single leaf area and chlorophyll content according to FAO descriptors with modifications Data was analyzed usingboth DARwin software V6 and Genstat Version 14 We observed significant differences among the traits implying great geneticvariability among the evaluated spider plant accessionsThe high genetic variation was further validated using the Unweighted PairGroup Method with Arithmetic mean (UPGMA) clusteringmethod with stem and flower colour as key traits The 49-spider plantaccessionswere clustered into 2major groups each consisting of Kenyan and SouthAfrican accessions Stepwise regression revealedthat plant height had themost influence on yield in terms of number of leaves per plantWe also observed high heritability for severaltraits including days to flowering (91) number of leaves per plant (99) plant height (99) number of primary branches (94)chlorophyll content (94) and single leaf area (87) Our results reveal the high genetic variation between different spider plantaccessions especially from different regions of Africa that could be further exploited to improve productivity in the plantThe highheritability of most of the yield related traits is promising for improving yield in the crop through direct selection

1 Introduction

Cleome gynandra also known as ldquoAfrican spider plantrdquo isamong themost important traditional leafy vegetables widelyused in Africa [1] It belongs to the family of CapparaceaeIt is also an erect herbaceous annual herb that is mainlyself-pollinated [2] The plant is highly nutritive and con-tains health promoting bioactive compounds important incombating malnutrition and reducing human degenerativediseases Spider plant is native to the Southern AfricaWestern Africa Central Africa Eastern Africa and SouthEast Asia [3] In South Africa spider plant has been foundto grow in the wild in KwaZulu-Natal Free State NorthernCape Limpopo and North West provinces [3] In Kenya theplants are mainly found in Western Rift Valley Eastern and

Coastal regions The key counties producing the crop includeKisii Nyamira Kericho Migori and Siaya [4] Despite thiswide adaptation and continued increase in production andconsumption there have been limited efforts towards itsimprovement There is lack of critical information on theextent and structure of phenotypic variation crucial for thebreeding and conservation of spider plant [2 5] Geneticdiversity is particularly useful in characterizing individualaccessions and cultivars in detecting genetic materials withnovel genes and thereby rescuing them from genetic erosionand as a general guide in selecting appropriate parents inbreeding programs Most of the genetic diversity observed inspider plant in Kenya and South Africa has traditionally beenmaintained by farmers in situ This poses the risk of speciesextinction due to loss of natural habitat as humans continue

HindawiAdvances in AgricultureVolume 2018 Article ID 8568424 11 pageshttpsdoiorg10115520188568424

2 Advances in Agriculture

to exploit and develop land divert water flow and changethe environment Secondly as human population continuesto increase there is pressure on natural land being clearedby human activity The need for cultivation conservationand characterization of spider plant remains imperative inmaintaining the integrity of the genetic information anddiversity

Mendelian analysis of discrete morphological traits canbe used to estimate genetic diversity in plants [6] and hasbeen successfully used in spider plant [5] Some of the keytraits that have been used as a guide in selection for goodgenotypes in previous studies included high heritability traitssuch as days to flowering plant height and number of leavesper plant [7] Omondi [7] observed that higher leaf yieldplant uniformity longer vegetative phase late flowering anddrought tolerance could form the best criterion in selectionof good performing spider plant accessions However foran efficient crop improvement program information onestimates of heritability for these desirable traits must beestablished [8] Due to limited knowledge on the geneticvariability more research remains of the essence to elucidatethe genetic and phenotypic diversity of existing spider plantaccessions Thus the main thrust of this study was tounderstand the extent of phenotypic diversity and heritabilityof qualitative traits among 49 spider plant accessions assem-bled from Kenya and South Africa Promising spider plantaccessions can be utilized in various breeding programs andhave the potential of enhancing its utilization while aiding tofight hidden hunger in Kenya

2 Materials and Methods

21 Plant Materials The study used 49 spider plant acces-sions mainly local landraces assembled from 3 sources Genebank of Kenya (25) Gene bank of South Africa (9) andKenyan farmersrsquo landraces (15) (Table 1)

22 Experimental Design and Study Site The experimentswere carried out at the University of Nairobirsquos Kabete Fieldstation (Nairobi Kenya) for two seasons fromMarch 2014 toMay 2014 and October 2014 to January 2015The experimentswere laid out in a randomized complete block design withthree replications Kabete Field station lies at 36∘411015840E and01∘15rsquoSwith an altitude of 1737m above sea level It receives anaverage temperature of 23∘C with a bimodal rainfall patternand an annual precipitation of 600mm to 1800mm The soiltype is well drained very dark reddish brown to dark redfriable clay locally known as Kikuyu red clay loam with anaverage pH of 62 [9]

23 Crop Husbandry Pregermination for each accession wasdone for 72 hours under treatment with 02 gibberellicacid to break seed dormancy and enhance germination [10]Each individual accession was planted by hand in two rowscomprising ten seeding holes per row (20 plants in a plot)Row plots were 3m in length with inter-row spacing of30 cm and intra-row spacing of 30 cm Farmyard manurewas applied to rows at the rate of 105 gaccession and mixedwith soil at planting Hand weeding was done throughout the

Table 1 List of Kenyan and South African spider plant accessionsevaluated in the study

Entry Accession no Country of origin Region1 1ke Kenya Siaya2 2ke Kenya Bungoma3 3ke Kenya Kakamega4 4ke Kenya Kitale5 5ke Kenya Mbale6 6ke Kenya Bomet7 7ke Kenya Busia8 9ke Kenya Marakwet9 10ke Kenya Kisumu10 11ke Kenya Homabay11 12ke Kenya Nandi12 13ke Kenya Kakamega13 14ke Kenya Kisii14 15ke Kenya Mbale15 16ke Kenya Meru16 1959sa South Africa Mpumalanga17 1988sa South Africa Mpumalanga18 2000sa South Africa Mpumalanga19 2232sa South Africa Northern province20 2241sa South Africa Northern province21 2249sa South Africa Northern province22 2279sa South Africa Northern province23 2289sa South Africa Mpumalanga24 2299sa South Africa Mpumalanga25 30316ke Kenya Western26 31990ke Kenya Western27 31992ke Kenya Western28 45426ke Kenya Western29 45446ke Kenya Central30 45451ke Kenya Central31 50259ke Kenya Kisii32 50264ke Kenya Nyamira33 50265ke Kenya Nyamira34 50273ke Kenya Nyamira35 50290ke Kenya Nyamira36 50296ke Kenya Nyamira37 50298ke Kenya Nyamira38 50299ke Kenya Nyamira39 50307ke Kenya Kisii40 50319ke Kenya Nyamira41 50325ke Kenya Kisii42 50326ke Kenya Nyamira43 50328ke Kenya Nyamira44 50330ke Kenya Nyamira45 50332ke Kenya Kisii46 50339ke Kenya Nyamira47 50353ke Kenya Nyamira48 50584ke Kenya Nyamira49 50600ke Kenya Kisiike=originated from Kenya sa=originated from South Africa

Advances in Agriculture 3

Table 2 Character descriptor and codes used for characterization of qualitative traits in spider plant accessions

SNo Character Descriptor and code1 Growth habit Erect (2) semi-erect (4) and prostrate (6)2 Flower colour White (1) purple (2) and pink (3)3 Stem colour Green (1) pink (2) violet (3) and purple (4)4 Stem hairiness Glabrous (1) weaksparse (3) medium (5) and profuse (7)5 Petiole colour Green (1) pink (2) violet (3) and purple (4)6 Petiole hairiness Glabrous (1) weaksparse (3) medium (5) and profuse (7)7 Leaf colour Dark green (1) and light green (2)8 Leaf hairiness Glabrous (1) weaksparse (3) medium (5) and profuse (7)Source Food and Agriculture Organization of the United Nations (FAO 1995) numbers in brackets on the right-hand side are the corresponding descriptorcodes listed in the FAO publication with modifications during the development of the list

experimental period The experiment was conducted underrain-fed conditions with supplemental overhead irrigationwhen required

24 Data Collection and Analysis There were two sets of datacollected in the study namely qualitative (morphological)and quantitative (agronomic)

25 Qualitative Traits Spider plant traits that were consid-ered qualitative included growth habit flower colour stemcolour stem hairiness petiole colour petiole hairiness leafcolour and leaf pubescence based on the list of modified spi-der plant descriptors [11] (Table 2) Three randomly selectedplants were tagged per accession per replicate during cropgrowth before flowering The data was subjected to DARwin50 software as described by Perrier and Jacquemoud-Collet[12] Euclidean distance matrix and hierarchical clusteringanalyses of Unweighted Pair Group Method of Arithmeticaveraging were used to estimate dissimilarities among theaccessions and results displayed in a dendrogram Thiswas followed by the identification of the most significantdescriptors contributing tomost phenotypic variation amongthe spider plant accessions through a stepwise regressionanalysis

26 Quantitative Traits All the yield and yield related traitswere considered quantitative including days to 50 flower-ing SPAD values plant height number of primary branchesleaf length leaf width single leaf area and number of leavesper plant Using Genstat version 14 software as describedby [13] the data was subjected to Analysis of Variance toestablish any significance differences among the traits and toobtain genotype means which were then separated using theFishers protected least significant differences (LSD) at P lt005

To establish the relationship among the traits collected atwo-tail correlation analysis was performed to estimate quan-titative relationships among the traits and also to identifythose traits that could be of great significance in a spider plantbreeding program

Heritability in the broad sense was estimated as a ratio ofgenotypic variance to the phenotypic variance and expressedin percentage [14] as per the following equation

Heritability (1198672) = (119881119892119881119901 times 100) (1)

where Vg is the genotypic variance and Vp is the phenotypicvariance

Genotypic variance (1205902119892) was derived by subtractingerror mean sum of squares (EMS) from the genotypic meansum of squares (GMS) and divided by the number ofreplications as given by the following equation

1205902119892 = 119866119872119878 minus 119864119872119878119903 (2)

where

GMS is the genotype mean sum of squares EMS isthe error mean sum of squares and r is the number ofreplications

Phenotypic variance (1205902119901) was derived by adding genotypicvariance with error variance as per the following equation

1205902119901 = 1205902119890 + 1205902119892 (3)

3 Results and Discussions

31 Qualitative Traits There were three distinct flowercolours displayed by the spider plant accessions purplepink and white Among these the purple flower colour wasthe most dominant among the Kenyan accessions at 49(Table 3) while the most dominant flower colour amongthe South African accessions was white Most of the SouthAfrican accessions displayed a green stemwith a green petioleas opposed to the Kenyans accessions which displayedpurple stems and pubescence Previous study byMasuka andMazarura [15] reported that purple-stemmed plants tendedto bemore hairy (trichomes) than the green-stemmed plantsAnthocyanins have been implicated as responsible for the


Table3Morph

ologicaldescrip

torsrecorded

forthe

49field

grow

nspider

plantaccessio

nsforthe

combinedseason

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leaf

hairiness

Growth

habit

11

Kenya

Purple

Purple

Green

Profuse

Medium

dark

green

Medium

Erect

22

Kenya

White

Purple

Pink

Profuse

Medium

light

green

Sparse

Erect

33

Kenya

Pink

Purple

Pink

Profuse

Medium

dark

green

Sparse

Erect

44

Kenya

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

55

Kenya

White

Purple

Purple

Profuse

Profuse

light

green

Profuse

Erect

66

Kenya

Pink

Purple

Pink

Profuse

Profuse

dark

green

Medium

Erect

77

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

89

Kenya

White

Purple

Purple

Profuse

Profuse

dark

green

Sparse

Erect

910

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1011

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Medium

Erect

1112

Kenya

Pink

Purple

Purple

Profuse

Medium

light

green

Sparse

Erect

1213

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1314

Kenya

Purple

Green

Purple

Profuse

Profuse

dark

green

Medium

Erect

1415

Kenya

Pink

Purple

Purple

Medium

Sparse

dark

green

Sparse

Erect

1516

Kenya

Pink

Purple

Pink

Medium

Sparse

light

green

Sparse

Erect

161959

SAfrica

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

171988

SAfrica

White

Green

Green

Sparse

Sparse

light

green

Sparse

Semierect

182000

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

ssemierect

192232

SAfrica

White

Green

Green

Glabrou

sGlabrou

sdark

green

Glabrou

sErect

202241

SAfrica

White

Green

Pink

Sparse

Sparse

light

green

Sparse

Erect

212249

SAfrica

White

Green

Pink

Glabrou

sGlabrou

slig

htgreen

Glabrou

sErect

222279

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

sSemierect

232289

SAfrica

White

Green

Pink

Medium

Sparse

dark

green

Sparse

Erect

242299

SAfrica

White

Green

Green

Glabrou

sSparse

light

green

Glabrou

sErect

2530316

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect


Table3Con

tinued

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leafha

iriness

Growth

habit

2631990

Kenya

Purple

green

Green

Medium

Sparse

light

green

Sparse

Erect

2731992

Kenya

Pink

purple

Green

Profuse

Medium

dark

green

Medium

Erect

2845426

Kenya

Purple

Purple

Green

Profuse

Sparse

dark

green

Sparse

Erect

294544

6Ke

nya

White

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3045451

Kenya

Pink

Purple

Purple

Profuse

Profuse

dark

green

Medium

Erect

3150259

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3250264

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3350265

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3450273

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3550290

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Semierect

3650296

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3750298

Kenya

Purple

Purple

Purple

Medium

Sparse

light

green

Sparse

Semierect

3850299

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3950307

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

Erect

4050319

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4150325

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

4250326

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4350328

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4450330

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4550332

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4650339

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4750353

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4850584

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

495060

0Ke

nya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect


stem pigmentation in most herbaceous plants [16] and havealso been widely studied for their potential medicinal value[17] Although spider plants have been traditionally usedfor medicinal purposes [18] the obvious contrast betweenSouth African and Kenyan accessions in their anthocyanincontent calls formore studies in order to elucidate the benefitsof the variations observed Presence of trichomes on theother hand has been associated with insect resistance inseveral studies including soybean [19] pigeonpea [20] andBrassicaceae [21] Trichomes are considered a domesticationtrait and are often more abundant in unadapted landracesthan in improved germplasm The absence of trichomesin South African accessions may suggest that they haveundergone much more intense selection cycles than theKenyan accessions although more studies would need to bedone to confirm this fact

Erect growth habit was more dominant and was observedin 90 of the accessions as opposed to the semi-erect growthhabit observed among 10 of the accessionsThis observationagrees with earlier findings [22] which reported over 80erect growth in the studied spider plant accessions Otherpast research has also shown that majority of the spider plantmorphotypes present an erect type of growth [5] Growthhabit is crucial in vegetable breeding as reported in otherindigenous vegetables [23] Bushy growth habit results inmany small leaves arising from numerous shoots that are nota preference trait to producers while the erect growth habitwith only primary and secondary branches maximizes theleaf area This implies that yield improvement in spider plantcould be exploited through selection of genotypes exhibitingerect growth habit and therefore large leaf size Furtherreports have suggested that inmixed cropping farmers couldadopt the semi-erect type whereas the erect types are ideal forintercrop adaptability [14]

32 Cluster Analysis Sufficient phenotypic variation wasobserved among the accessions as revealed by the clusteranalysis (Figure 1) Two major clusters namely clusters 1and 2 were distinguished using the eight morphologicaldescriptors Cluster 1 included 9 accessions of South Africanorigin with an exception of one Kenyan accession 1959 whilecluster 2 is comprised of 40 Kenyan accessions inclusiveof one South African origin Stem and petiole colour werethe major traits that contributed to the Kenyan accession31990 grouping together with the South African accessionsthat were mainly green stemmed with green petiole Theexceptional South African accession grouped together withthe Kenyan accessions due to the purple stem colour and pro-fuse pubescence that were predominant among the Kenyanaccessions This clearly revealed the differences in the geneticmakeup of the accessions from the two regions Howeverthe South African accessions 2279 and 2000 which werecollected from the Northern Province showed similarityin their phenotypic traits Most of the Kenyan gene bankaccessions namely 50339 50330 50328 50326 50299 and50273 from Nyamira region clustered closely together withaccession 30316 from Western region despite being collectedfrom different regions Additionally Kenyan accession 45451from central region and Kenyan accession 14 collected from

Kisii region grouped together suggesting some degree ofsimilarity in their morphological traits Most of the Kenyanfarmersrsquo landraces clustered together based on their originThere were major overlaps with the accessions assembledfrom the gene bank implying that they could have samegenetic makeup A study by Krsquoopondo [5] has demonstrateda close relationship among spider plant genotypes followingthe evaluation of the variability in seed proteins among themIn addition this uniformity could also arise from the self-pollination status of spider plant However more character-ization needs to be done to validate such findings The clusteranalysis which clearly grouped the accessions according totheir geographical origin suggests that crop improvementof spider plant could be achieved through exploiting thevariation revealed However the current cluster analysis wasdone using morphological traits which can be influencedby several environmental factors There will be need toundertake a more detailed genetic analysis using molecularmarkers to confirm the existence of genetic variation acrossthe different geographical regions There is need for specificregional breeding efforts to target preferred traits [24 25]

33 Quantitative Traits

331 Analysis of Variance Spider plant accessions showedsignificant differences (Plt005) with no seasonal effect forall the traits namely days to 50 flowering single leafarea leaf length and width chlorophyll content number ofleaves per plant number of primary branches and plantheight (Table 4) implying the existence of variability for therespective traits among spider plant accessions Accessionsthat exhibited longer days to 50flowering also yielded moreleaf count Late flowering enables a plant to have a longervegetative phase during growth period [7] Past researchhas associated late flowering with increased leaf yield andconsequently early flowering as a limit to leaf yield in otherindigenous vegetables [23] This suggests that late floweringwould be a good selection criterion for yield improvement inspider plant

Other traits that contributed to increased leaf countwere plant height and number of primary branches Kenyanaccessions were taller than the South African accessionswith plant height varying from 21 cm for accession 2249 to113 cm for accession 50296TheKenyan accessions performedbetter than South African genotypes for the number of leavesper plant number of primary branches leaf length leafwidth plant height single leaf area and chlorophyll contentconforming to past research by Wasonga [22] The best 5outstanding accessions with regard to yield related traits like50flowering single leaf area and number of leaves per plantincluded Kenyan accessions 3 7 45451 50296 and SouthAfrican accession 2241 (Table 5)

332 Correlation among the Traits Knowledge of the cor-relations among yield and the yield related traits is ofconsiderable importance in crop improvement because it aidsin indirect selection [26] There was positive and significantcorrelation between leaf length leaf width and leaf areawith number of days to 50 flowering Table 6 This agrees


Cluster 2

Cluster 1

Kenyan gene bank accession Kenyan farmerrsquos landraces South African gene bank accessions

50353

50332

50319

50296

50265

7

50290

11

50600

50339

50330

50328

50326

50299

50273

50264

30316

50259

45446

12

50584

50307

13

10

50325

16

15

50298

45451

14

6

31992

1

1959

4

45426

9

5

3

2

31990

10

2289

2241

1988

2249

2232

2279

2000

2299

Figure 1 Phenogram showing relationship among accessions characterized using morphological traits

with the findings of Kiebre et al [27] who also reporteda positive significant correlation between number of daysto 50 flowering with leaf length and width This furthersuggests that the late flowering genotypes could be selectedfor their big size which is a crucial trait to the producers whoregard leaf biomass as key in leafy vegetables production

There was a significant positive correlation between plantheight and number of primary branches conforming to theresults of Kiebre et al [27] indicating the taller the plantthe more the number of primary branches This is furthersupported by the observed correlations where yield in termsof number of leaves per plant had a positive and significantcorrelation with plant height (r = 069) and number ofprimary branches (r = 063) implying that the higher the

number of the branches and the taller the plant the higherthe number of leaves Single leaf area correlated positivelywith leaf length width and days to 50 flowering at r =092 r = 088 and r = 021 respectively As expected the leafarea would be determined by its length and width suggestingthe longer and wider the leaf the bigger the leaf area Thissuggests leaf length and width as important traits in selectingfor vegetative yield in spider plant

However there was a nonsignificant negative correlationbetween leaf size and leaf yield indicating the more thenumber of leaves in the plant the smaller the leaves Yield isinfluenced by complex soil plant interactions in many cropsIn this study the chlorophyll contentmeasured in SPADvaluehad a positive significant correlation with number of leaves


Table 4

(a) Analyses of variance showing the mean squares for the agronomic traits in Cleome gynandra season one (April-July 2014)

Source of variation df DTF LL LW NPB NLPP PH SLA SPADRep 2 43 06 17 16 279 65 04 1Genotype 48 347lowast 52lowast 163lowast 147lowast 68956lowast 27340lowast 81lowast 1654lowastResidual 96 1 02 14 03 134 82 04 21Total 146

(b) Analyses of variance showing the mean squares for the agronomic traits in Cleome gynandra season one (October-July 2014)


(c) Analyses of variance showing the mean squares for the agronomic traits in Cleome gynandra for the combined seasons

Source of variation df DTF LL LW NPB NLPP PH SLA SPADRep 2 91 1 12 16 16 47 09 32Genotype 48 646lowast 103lowast 326lowast 261lowast 138402lowast 54511lowast 159lowast 3008lowastSeason 1 1616lowast 10lowast 41lowast 21lowast 54 334lowast 21lowast 48lowastGenotype Season 48 12 01 02 04 167 67 01 41Residual 194 09 02 09 04 155 73 03 32Total 293lowastSignificant at P lt005 DTF days to 50 flowering SLA single leaf area (cm2) LL leaf length (cm) LW leaf width (cm) NLPP number of leaves per plantNPB number of primary branches PH plant height (cm) and SPAD soil plant analysis development

per plant (r = 045) number of primary branches (r = 054)and plant height (r = 059) This contradicts the findings of[28] who reported negative correlations between chlorophyllreadings with yield related traits except for plant height inbeans This positive significance correlation between SPADvalues and yield related traits calls for more studies toelucidate this phenomenon Leaf yields may be improvedthrough selection of accessions that showed high leaf countas well as large single leaf area

34 Stepwise Regression Themost important traits that havea considerable effect on the dependant variable are verifiedthrough a stepwise regression analysis The traits selectedthrough the regression model can then be used as a selec-tion criterion for indirect selection in a breeding program[29] A multiple linear regression analysis was calculated byconsidering the number of leaves as the dependent variableand other characters as the independent variables Results ofregression analysis showed that plant height had a significantinfluence on yield (R2 = 467 P value le 005) (Table 7) Thisimplies that selection based on plant height will influence andincrease vegetative yield in C gynandra This further agreeswith other findings by Nwangburuka et al [30] in vegetableC olitorius where plant height was found to significantlyincrease leaf yield

35 Heritability Estimates for Yield and Yield Related TraitsThe estimates of heritability in broad sense for all the traitsranged from 78 to 99 (Table 8) High percentages of

broad sense heritability were estimated for number of leavesper plant plant height at 99 and SPAD value at 96Leaf width exhibited a moderately lower percentage at 78followed by single leaf area and leaf length at 86 and 89respectively (Table 8) High heritability plays a great role inselection for crop improvement as the traits to be improveddepend immensely on their heritability and variability [27]In this study the genotypic variance of all traits was higherthan the environmental variance implying that much of thephenotypic variation among the accessions was attributed tovariation in genotype as opposed to the environment Thehigh estimates of heritability displayed in the study suggestthat selection for yield improvement in spider plant couldbe based on traits like number of leaves per plant and plantheight

4 Conclusions and Recommendations

This study reported the existence of significant phenotypicvariation in Cleome gynandra as evidenced by the mor-phological characterization which clearly distinguished theaccessions from the two regions The new knowledge gener-ated on the spider plant morphological structure could offera great potential in developing relevant genetic and genomicresources for spider plant breeding programs It is also clearthat indirect selection for improved spider plant accessionscould be based on the yield related traits like number ofleaves per plant plant height number of primary branchesand days to flowering which exhibited high heritability Thisstudy recommends the complementation of morphological


Table 5 Mean comparison of the quantitative traits of 49 spider plant accessions from Kenya and South Africa grown in the University ofNairobi Field at Kabete for the two combined seasons

Entry Accession No Origin DTF LL LW NPB NLPP PH SLA SPAD1 1 Kenya 397 51 142 63 105 383 87 5642 2 Kenya 457 64 115 57 588 315 88 5763 3 Kenya 453 67 17 7 1122 41 109 5034 4 Kenya 438 52 138 75 758 367 87 5695 5 Kenya 453 5 132 75 538 468 83 5766 6 Kenya 397 5 117 75 572 478 78 5697 7 Kenya 45 59 16 77 913 512 99 538 9 Kenya 455 62 143 65 943 39 96 5669 10 Kenya 415 56 117 57 68 407 83 58310 11 Kenya 45 36 106 55 1005 401 63 56811 12 Kenya 46 46 119 75 61 422 76 58812 13 Kenya 378 5 131 62 648 383 83 55913 14 Kenya 398 47 113 78 833 632 74 53314 15 Kenya 387 37 93 57 557 378 6 46715 16 Kenya 452 78 156 9 757 755 112 61616 1959 S Africa 438 58 12 67 892 343 85 54317 1988 S Africa 342 68 126 57 507 452 95 49918 2000 S Africa 328 65 136 42 197 308 96 53119 2232 S Africa 398 53 10 47 56 263 74 24320 2241 S Africa 453 78 154 78 41 423 112 4421 2249 S Africa 393 62 132 42 203 212 92 42922 2279 S Africa 532 63 123 67 232 221 9 39623 2289 S Africa 44 72 139 63 313 417 102 4324 2299 S Africa 40 51 98 73 78 303 72 43725 30316 Kenya 377 59 103 10 146 83 8 57126 31990 Kenya 41 57 132 85 1093 913 88 55427 31992 Kenya 42 8 11 92 201 917 99 59528 45426 Kenya 43 74 115 10 97 97 95 58429 45446 Kenya 423 88 13 83 68 1117 111 57330 45451 Kenya 477 72 161 92 2475 1093 109 53931 50259 Kenya 44 57 106 92 182 1013 79 58132 50264 Kenya 402 44 105 117 1722 1088 69 60833 50265 Kenya 443 55 116 103 787 922 82 60134 50273 Kenya 403 51 95 87 928 897 71 57535 50290 Kenya 403 48 104 93 1052 932 71 62236 50296 Kenya 40 105 183 107 1095 113 142 57937 50298 Kenya 40 44 79 115 140 827 61 59538 50299 Kenya 417 48 94 105 1013 998 69 58939 50307 Kenya 40 6 12 83 1407 938 87 58440 50319 Kenya 403 62 125 95 101 778 9 62541 50325 Kenya 40 68 121 10 948 95 93 57942 50326 Kenya 443 56 111 125 1658 1012 81 57543 50328 Kenya 402 54 86 87 965 757 71 58644 50330 Kenya 398 57 104 98 172 1043 78 60845 50332 Kenya 405 42 88 92 1158 1105 62 62546 50339 Kenya 427 52 128 125 1498 1017 83 61647 50353 Kenya 423 45 84 11 1468 86 63 57848 50584 Kenya 397 48 98 97 888 787 7 57949 50600 Kenya 388 59 88 95 788 988 75 562

Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33

DTF days to 50 flowering SLA single leaf area (cm2) LL leaf length (cm) LW leaf width (cm) NLPP number of leaves per plant NPB number of primarybranches PH plant height (cm) and SPAD soil plant analysis development


Table 6 Correlation in combined seasons

DTF LL LW NPB NLPP PH SLA SPADDTF -LL 012lowast -LW 028lowastlowast 062lowastlowast -NPB 009 002 -020lowastlowast -NLPP 01 -001 -011 063lowastlowast -PH -007 016lowast -019lowast 082lowastlowast 069lowastlowast -SLA 021lowastlowast 092lowastlowast 088lowastlowast -008 -006 001 -SPAD -004 -007 -011 054lowastlowast 045lowastlowast 059lowastlowast -01 -lowast implies significance difference at Plt005 lowastlowast implies significance difference at plt0001 (2-tailed) DTF- days to 50 flowering SLA- single leaf area (cm2)LL- leaf length (cm) LW- leaf width (cm) NLPP- number of leaves per plant NPB- number of primary branches PH- plant height (cm) SPAD- soil plantanalysis development

Table 7 Stepwise regression analysis of the 7 evaluated traits

Step Variable Partial R square Adjusted R square F-test1 Plant height 048 047 4300lowastlowastSignificant at ple 005 y = 1102x + 21947 y = number of leaves per plant and x = plant height

Table 8 Estimates of yield and yield related components of 49 spider plant accessions

Traits VE VG VP HBS ()

DTF Season 1 10 112 122 918Season 2 10 101 111 910

LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792

NPB Season 1 03 48 51 941Season 2 03 38 41 927

NLPP Season 1 134 22941 23075 994Season 2 134 23146 23280 994

PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991

SLA Season 1 04 26 30 865Season 2 04 26 30 865

SPAD Season 1 21 544 565 963Season 2 21 451 472 955

VE = environmental variance VG = genotypic variance VP = phenotypic variance HBS = broad sense heritability NLLP = number of leaves per plant NPB= number of primary branches LL = leaf length LW = leaf width PH = plant height SLA = single leaf area and SPAD = soil plant analysis development

characterization with the use of molecular markers forgermplasm characterization and genetic diversity since theyare under little influence from the environment

Conflicts of Interest

The authors declare no conflicts of interest regarding thepublication of this paper

Acknowledgments

The authors would like to acknowledge the University ofNairobi for the opportunity to undertake the research This

paper is part of the MS thesis entitled ldquoGenetic Characteri-zation and Nutrition Analysis of Eastern and South AfricanCleome gynandra (Spider Plant) Accessionsrdquo submitted to theUniversity of Nairobi towards achievement of an MS degreein plant breeding and biotechnology

References

[1] R R Schippers ldquoAfrican indigenous vegetables an overview ofthe cultivated speciesrdquo 2000

[2] J A Chweya and N A Mnzava Catrsquos whiskers gynandra LPromoting the conservation and use of underutilized and neglect-ed crops vol 11 1997


[3] Department of Agriculture Forestry and Fisheries (DAFF)Cleome Resource Centre Pretoria South Africa

[4] HCDA Horticulture Data 2011-2013 Validation Report Horti-cultural Crops Development authority

[5] F B Krsquoopondo ldquoMorphological characterization of selectedspider plant (Cleome gynandraL) types from western KenyardquoAnnals of Biological Research vol 2 no 2 pp 54ndash64 2011

[6] S A Mohammadi and B M Prasanna ldquoAnalysis of geneticdiversity in crop plantsmdashsalient statistical tools and consider-ationsrdquo Crop Science vol 43 no 4 pp 1235ndash1248 2003

[7] C O Omondi Variation and Yield Prediction Analyses of SomeMorphological Traits in Six Kenyan Landraces Population ofSpider flower (Gynandropsis gynandra (L) Briq) [PhD thesis]Univ Nairobi Nairobi Kenya

[8] S Shukla A Bhargava A Chatterjee A Srivastava and S PSingh ldquoGenotypic variability in vegetable amaranth (Amaran-thus tricolor L) for foliage yield and its contributing traits oversuccessive cuttings and yearsrdquo Euphytica vol 151 no 1 pp 103ndash110 2006

[9] R Jaetzold ldquoNatural conditions and farm management infor-mation of Central Kenya Farm management handbook ofKenyardquo 2004

[10] F Koyuncu ldquoBreaking seed dormancy in black mulberry(Morus nigra L) by cold stratification and exogenous appli-cation of gibberellic acidrdquo Acta Biologica Cracoviensia SeriesBotanica vol 47 no 2 pp 23ndash26 2005

[11] Food and Agricultural Organisation (FAO) Production yearbook vol 49 FoodandAgriculturalOrganisation (FAO)RomeItaly 1995

[12] X Perrier and J P Jacquemoud-Collet ldquoDARwin softwarerdquo[13] RW Payne D A Murray S A Harding D B Baird and D

Soutar ldquoAn introduction to GenStat for Windowsrdquo vol 2011VSN International Hemel Hempstead UK

[14] CHHansonH F Robinson andR EComstock ldquoBiometricalStudies of Yield in Segregating Populations of Korean Lespedeza1rdquo Agronomy journal vol 48 no 6 pp 268ndash272 1956

[15] A Masuka M Goss and U Mazarura ldquoMorphological char-acterization of four selected spider plant (Cleome gynandraL) morphs from Zimbabwe and Kenyardquo Asian Journal ofAgriculture and Rural Development vol 2 no 4 p 646 2012

[16] K S Gould D A Dudle and H S Neufeld ldquoWhy some stemsare red Cauline anthocyanins shield photosystem II againsthigh light stressrdquo Journal of Experimental Botany vol 61 no 10pp 2707ndash2717 2010

[17] H E Khoo A Azlan S T Tang and S M Lim ldquoAnthocyani-dins and anthocyanins colored pigments as food pharmaceu-tical ingredients and the potential health benefitsrdquo in Food ampNutrition Research vol 61 2017

[18] S Venter W S Jansen van Rensburg H J Vorster E Vanden Heever and J J B Zijl ldquoPromotion of African leafy vege-tables within the Agricultural Research Council-Vegetable andOrnamental Plant Institute The impact of the projectrdquo

[19] H-X Chang A E Lipka L L Domier and G L Hart-man ldquoCharacterization of disease resistance loci in the USDAsoybean germplasm collection using genome-wide associationstudiesrdquo Journal of Phytopathology vol 106 no 10 pp 1139ndash11512016

[20] R Aruna D M Rao L J Reddy H D Upadhyaya and H CSharma ldquoInheritance of trichomes and resistance to pod borer(Helicoverpa armigera) and their association in interspecificcrosses between cultivated pigeonpea (Cajanus cajan) and its

wild relative C scarabaeoidesrdquo Euphytica vol 145 no 3 pp247ndash257 2005

[21] U Alahakoon J Adamson L Grenkow J Soroka P Bonham-Smith andMGruber ldquoField growth traits and insect-host plantinteractions of two transgenic canola (Brassicaceae) lines withelevated trichome numbersrdquo The Canadian Entomologist vol148 no 5 pp 603ndash615 2016

[22] D O Wasonga J L Ambuko G N Cheminingwa D AOdeny and B G Crampton ldquoMorphological Characterizationand Selection of Spider Plant (Cleome Gynandra) Accessionsfrom Kenya and South Africardquo Asian Journal of AgriculturalSciences vol 7 no 4 pp 36ndash44 2015

[23] C O Ojiewo O Mbwambo I Swai et al ldquoSelection evaluationand release of varieties from genetically diverse African night-shade germplasmrdquo International Journal of Plant Breeding vol7 no 2 pp 76ndash89 2013

[24] Z Kiebre P Bationo-Kando N Sawadogo M Sawadogo and JD Zongo ldquoSelection of phenotypic interests for the cultivationof the plant Cleome gynandra L in the vegetable gardens inBurkina Fasordquo Journal Of Experimental Biology and AgricultureSciences vol 3 pp 288ndash297 2015

[25] E D Sogbohossou E G Achigan-Dako P Maundu et al ldquoAroadmap for breeding orphan leafy vegetable species a casestudy of Gynandropsis gynandra (Cleomaceae)rdquo HorticultureResearch vol 5 no 1 p 2 2018

[26] N Ali F Javidfar J Y Elmira and M Y Mirza ldquoRelation-ship among yield components and selection criteria for yieldimprovement in winter rapeseed (Brassica napus L)rdquo PakistanJournal of Botany vol 35 no 2 pp 167ndash174 2003

[27] Z Kiebre P Bationo-Kando A Barro et al ldquoEstimates ofgenetic parameters of spider plant (Cleome gynandra L) ofBurkina Fasordquo International Journal of Agricultural Policy andResearch vol 5 no 9 pp 138ndash144 2017

[28] S Guler and H Ozcelik ldquoRelationships between leaf chloro-phyll and yield related characters of dry bean (Phaseolusvulgaris L)rdquo Journal of Plant Sciences vol 6 no 4 pp 700ndash7032007

[29] N Sabaghnia H Dehghani B Alizadeh andMMohghaddamldquoInterrelationships between seed yield and 20 related traits of49 canola (Brassica napus L) genotypes in non-stressed andwater-stressed environmentsrdquo Spanish Journal of AgriculturalResearch vol 8 no 2 pp 356ndash370 2010

[30] C C Nwangburuka O J Olawuyi K Oyekale K O Ogun-wenmo O A Denton and E Nwankwo ldquoGrowth and yieldresponse of Corchorus olitorius in the treatment of Arbuscularmycorrhizae (AM Poultry manure (PM)rdquoCombination of AM-PM and Inorganic Fertilizer (NPK) in Applied Science Researchvol 3 no 3 pp 1466ndash1471 2012

Nutrition and Metabolism

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BotanyJournal of


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Veterinary Medicine International


Cell BiologyInternational Journal of



BioMed Research International

Agronomy



Submit your manuscripts atwwwhindawicom


to exploit and develop land divert water flow and changethe environment Secondly as human population continuesto increase there is pressure on natural land being clearedby human activity The need for cultivation conservationand characterization of spider plant remains imperative inmaintaining the integrity of the genetic information anddiversity

Mendelian analysis of discrete morphological traits canbe used to estimate genetic diversity in plants [6] and hasbeen successfully used in spider plant [5] Some of the keytraits that have been used as a guide in selection for goodgenotypes in previous studies included high heritability traitssuch as days to flowering plant height and number of leavesper plant [7] Omondi [7] observed that higher leaf yieldplant uniformity longer vegetative phase late flowering anddrought tolerance could form the best criterion in selectionof good performing spider plant accessions However foran efficient crop improvement program information onestimates of heritability for these desirable traits must beestablished [8] Due to limited knowledge on the geneticvariability more research remains of the essence to elucidatethe genetic and phenotypic diversity of existing spider plantaccessions Thus the main thrust of this study was tounderstand the extent of phenotypic diversity and heritabilityof qualitative traits among 49 spider plant accessions assem-bled from Kenya and South Africa Promising spider plantaccessions can be utilized in various breeding programs andhave the potential of enhancing its utilization while aiding tofight hidden hunger in Kenya

2 Materials and Methods

21 Plant Materials The study used 49 spider plant acces-sions mainly local landraces assembled from 3 sources Genebank of Kenya (25) Gene bank of South Africa (9) andKenyan farmersrsquo landraces (15) (Table 1)

22 Experimental Design and Study Site The experimentswere carried out at the University of Nairobirsquos Kabete Fieldstation (Nairobi Kenya) for two seasons fromMarch 2014 toMay 2014 and October 2014 to January 2015The experimentswere laid out in a randomized complete block design withthree replications Kabete Field station lies at 36∘411015840E and01∘15rsquoSwith an altitude of 1737m above sea level It receives anaverage temperature of 23∘C with a bimodal rainfall patternand an annual precipitation of 600mm to 1800mm The soiltype is well drained very dark reddish brown to dark redfriable clay locally known as Kikuyu red clay loam with anaverage pH of 62 [9]

23 Crop Husbandry Pregermination for each accession wasdone for 72 hours under treatment with 02 gibberellicacid to break seed dormancy and enhance germination [10]Each individual accession was planted by hand in two rowscomprising ten seeding holes per row (20 plants in a plot)Row plots were 3m in length with inter-row spacing of30 cm and intra-row spacing of 30 cm Farmyard manurewas applied to rows at the rate of 105 gaccession and mixedwith soil at planting Hand weeding was done throughout the

Table 1 List of Kenyan and South African spider plant accessionsevaluated in the study

Entry Accession no Country of origin Region1 1ke Kenya Siaya2 2ke Kenya Bungoma3 3ke Kenya Kakamega4 4ke Kenya Kitale5 5ke Kenya Mbale6 6ke Kenya Bomet7 7ke Kenya Busia8 9ke Kenya Marakwet9 10ke Kenya Kisumu10 11ke Kenya Homabay11 12ke Kenya Nandi12 13ke Kenya Kakamega13 14ke Kenya Kisii14 15ke Kenya Mbale15 16ke Kenya Meru16 1959sa South Africa Mpumalanga17 1988sa South Africa Mpumalanga18 2000sa South Africa Mpumalanga19 2232sa South Africa Northern province20 2241sa South Africa Northern province21 2249sa South Africa Northern province22 2279sa South Africa Northern province23 2289sa South Africa Mpumalanga24 2299sa South Africa Mpumalanga25 30316ke Kenya Western26 31990ke Kenya Western27 31992ke Kenya Western28 45426ke Kenya Western29 45446ke Kenya Central30 45451ke Kenya Central31 50259ke Kenya Kisii32 50264ke Kenya Nyamira33 50265ke Kenya Nyamira34 50273ke Kenya Nyamira35 50290ke Kenya Nyamira36 50296ke Kenya Nyamira37 50298ke Kenya Nyamira38 50299ke Kenya Nyamira39 50307ke Kenya Kisii40 50319ke Kenya Nyamira41 50325ke Kenya Kisii42 50326ke Kenya Nyamira43 50328ke Kenya Nyamira44 50330ke Kenya Nyamira45 50332ke Kenya Kisii46 50339ke Kenya Nyamira47 50353ke Kenya Nyamira48 50584ke Kenya Nyamira49 50600ke Kenya Kisiike=originated from Kenya sa=originated from South Africa













1205902119892 = 119866119872119878 minus 119864119872119878119903 (2)

where



1205902119901 = 1205902119890 + 1205902119892 (3)




Table3Morph

ologicaldescrip

torsrecorded

forthe

49field

grow

nspider

plantaccessio

nsforthe

combinedseason

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leaf

hairiness

Growth

habit

11

Kenya

Purple

Purple

Green

Profuse

Medium

dark

green

Medium

Erect

22

Kenya

White

Purple

Pink

Profuse

Medium

light

green

Sparse

Erect

33

Kenya

Pink

Purple

Pink

Profuse

Medium

dark

green

Sparse

Erect

44

Kenya

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

55

Kenya

White

Purple

Purple

Profuse

Profuse

light

green

Profuse

Erect

66

Kenya

Pink

Purple

Pink

Profuse

Profuse

dark

green

Medium

Erect

77

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

89

Kenya

White

Purple

Purple

Profuse

Profuse

dark

green

Sparse

Erect

910

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1011

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Medium

Erect

1112

Kenya

Pink

Purple

Purple

Profuse

Medium

light

green

Sparse

Erect

1213

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1314

Kenya

Purple

Green

Purple

Profuse

Profuse

dark

green

Medium

Erect

1415

Kenya

Pink

Purple

Purple

Medium

Sparse

dark

green

Sparse

Erect

1516

Kenya

Pink

Purple

Pink

Medium

Sparse

light

green

Sparse

Erect

161959

SAfrica

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

171988

SAfrica

White

Green

Green

Sparse

Sparse

light

green

Sparse

Semierect

182000

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

ssemierect

192232

SAfrica

White

Green

Green

Glabrou

sGlabrou

sdark

green

Glabrou

sErect

202241

SAfrica

White

Green

Pink

Sparse

Sparse

light

green

Sparse

Erect

212249

SAfrica

White

Green

Pink

Glabrou

sGlabrou

slig

htgreen

Glabrou

sErect

222279

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

sSemierect

232289

SAfrica

White

Green

Pink

Medium

Sparse

dark

green

Sparse

Erect

242299

SAfrica

White

Green

Green

Glabrou

sSparse

light

green

Glabrou

sErect

2530316

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect


Table3Con

tinued

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leafha

iriness

Growth

habit

2631990

Kenya

Purple

green

Green

Medium

Sparse

light

green

Sparse

Erect

2731992

Kenya

Pink

purple

Green

Profuse

Medium

dark

green

Medium

Erect

2845426

Kenya

Purple

Purple

Green

Profuse

Sparse

dark

green

Sparse

Erect

294544

6Ke

nya

White

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3045451

Kenya

Pink

Purple

Purple

Profuse

Profuse

dark

green

Medium

Erect

3150259

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3250264

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3350265

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3450273

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3550290

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Semierect

3650296

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3750298

Kenya

Purple

Purple

Purple

Medium

Sparse

light

green

Sparse

Semierect

3850299

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3950307

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

Erect

4050319

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4150325

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

4250326

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4350328

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4450330

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4550332

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4650339

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4750353

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4850584

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

495060

0Ke

nya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect











Cluster 2

Cluster 1


50353

50332

50319

50296

50265

7

50290

11

50600

50339

50330

50328

50326

50299

50273

50264

30316

50259

45446

12

50584

50307

13

10

50325

16

15

50298

45451

14

6

31992

1

1959

4

45426

9

5

3

2

31990

10

2289

2241

1988

2249

2232

2279

2000

2299







Table 4
















Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy
















1205902119892 = 119866119872119878 minus 119864119872119878119903 (2)

where



1205902119901 = 1205902119890 + 1205902119892 (3)




Table3Morph

ologicaldescrip

torsrecorded

forthe

49field

grow

nspider

plantaccessio

nsforthe

combinedseason

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leaf

hairiness

Growth

habit

11

Kenya

Purple

Purple

Green

Profuse

Medium

dark

green

Medium

Erect

22

Kenya

White

Purple

Pink

Profuse

Medium

light

green

Sparse

Erect

33

Kenya

Pink

Purple

Pink

Profuse

Medium

dark

green

Sparse

Erect

44

Kenya

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

55

Kenya

White

Purple

Purple

Profuse

Profuse

light

green

Profuse

Erect

66

Kenya

Pink

Purple

Pink

Profuse

Profuse

dark

green

Medium

Erect

77

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

89

Kenya

White

Purple

Purple

Profuse

Profuse

dark

green

Sparse

Erect

910

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1011

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Medium

Erect

1112

Kenya

Pink

Purple

Purple

Profuse

Medium

light

green

Sparse

Erect

1213

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1314

Kenya

Purple

Green

Purple

Profuse

Profuse

dark

green

Medium

Erect

1415

Kenya

Pink

Purple

Purple

Medium

Sparse

dark

green

Sparse

Erect

1516

Kenya

Pink

Purple

Pink

Medium

Sparse

light

green

Sparse

Erect

161959

SAfrica

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

171988

SAfrica

White

Green

Green

Sparse

Sparse

light

green

Sparse

Semierect

182000

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

ssemierect

192232

SAfrica

White

Green

Green

Glabrou

sGlabrou

sdark

green

Glabrou

sErect

202241

SAfrica

White

Green

Pink

Sparse

Sparse

light

green

Sparse

Erect

212249

SAfrica

White

Green

Pink

Glabrou

sGlabrou

slig

htgreen

Glabrou

sErect

222279

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

sSemierect

232289

SAfrica

White

Green

Pink

Medium

Sparse

dark

green

Sparse

Erect

242299

SAfrica

White

Green

Green

Glabrou

sSparse

light

green

Glabrou

sErect

2530316

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect


Table3Con

tinued

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leafha

iriness

Growth

habit

2631990

Kenya

Purple

green

Green

Medium

Sparse

light

green

Sparse

Erect

2731992

Kenya

Pink

purple

Green

Profuse

Medium

dark

green

Medium

Erect

2845426

Kenya

Purple

Purple

Green

Profuse

Sparse

dark

green

Sparse

Erect

294544

6Ke

nya

White

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3045451

Kenya

Pink

Purple

Purple

Profuse

Profuse

dark

green

Medium

Erect

3150259

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3250264

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3350265

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3450273

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3550290

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Semierect

3650296

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3750298

Kenya

Purple

Purple

Purple

Medium

Sparse

light

green

Sparse

Semierect

3850299

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3950307

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

Erect

4050319

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4150325

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

4250326

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4350328

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4450330

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4550332

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4650339

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4750353

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4850584

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

495060

0Ke

nya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect











Cluster 2

Cluster 1


50353

50332

50319

50296

50265

7

50290

11

50600

50339

50330

50328

50326

50299

50273

50264

30316

50259

45446

12

50584

50307

13

10

50325

16

15

50298

45451

14

6

31992

1

1959

4

45426

9

5

3

2

31990

10

2289

2241

1988

2249

2232

2279

2000

2299







Table 4
















Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy





Table3Morph

ologicaldescrip

torsrecorded

forthe

49field

grow

nspider

plantaccessio

nsforthe

combinedseason

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leaf

hairiness

Growth

habit

11

Kenya

Purple

Purple

Green

Profuse

Medium

dark

green

Medium

Erect

22

Kenya

White

Purple

Pink

Profuse

Medium

light

green

Sparse

Erect

33

Kenya

Pink

Purple

Pink

Profuse

Medium

dark

green

Sparse

Erect

44

Kenya

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

55

Kenya

White

Purple

Purple

Profuse

Profuse

light

green

Profuse

Erect

66

Kenya

Pink

Purple

Pink

Profuse

Profuse

dark

green

Medium

Erect

77

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

89

Kenya

White

Purple

Purple

Profuse

Profuse

dark

green

Sparse

Erect

910

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1011

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Medium

Erect

1112

Kenya

Pink

Purple

Purple

Profuse

Medium

light

green

Sparse

Erect

1213

Kenya

Purple

Purple

Purple

Medium

Medium

light

green

Sparse

Erect

1314

Kenya

Purple

Green

Purple

Profuse

Profuse

dark

green

Medium

Erect

1415

Kenya

Pink

Purple

Purple

Medium

Sparse

dark

green

Sparse

Erect

1516

Kenya

Pink

Purple

Pink

Medium

Sparse

light

green

Sparse

Erect

161959

SAfrica

Pink

Purple

Pink

Profuse

Medium

light

green

Medium

Erect

171988

SAfrica

White

Green

Green

Sparse

Sparse

light

green

Sparse

Semierect

182000

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

ssemierect

192232

SAfrica

White

Green

Green

Glabrou

sGlabrou

sdark

green

Glabrou

sErect

202241

SAfrica

White

Green

Pink

Sparse

Sparse

light

green

Sparse

Erect

212249

SAfrica

White

Green

Pink

Glabrou

sGlabrou

slig

htgreen

Glabrou

sErect

222279

SAfrica

White

Green

Green

Glabrou

sGlabrou

slig

htgreen

Glabrou

sSemierect

232289

SAfrica

White

Green

Pink

Medium

Sparse

dark

green

Sparse

Erect

242299

SAfrica

White

Green

Green

Glabrou

sSparse

light

green

Glabrou

sErect

2530316

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect


Table3Con

tinued

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leafha

iriness

Growth

habit

2631990

Kenya

Purple

green

Green

Medium

Sparse

light

green

Sparse

Erect

2731992

Kenya

Pink

purple

Green

Profuse

Medium

dark

green

Medium

Erect

2845426

Kenya

Purple

Purple

Green

Profuse

Sparse

dark

green

Sparse

Erect

294544

6Ke

nya

White

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3045451

Kenya

Pink

Purple

Purple

Profuse

Profuse

dark

green

Medium

Erect

3150259

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3250264

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3350265

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3450273

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3550290

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Semierect

3650296

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3750298

Kenya

Purple

Purple

Purple

Medium

Sparse

light

green

Sparse

Semierect

3850299

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3950307

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

Erect

4050319

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4150325

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

4250326

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4350328

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4450330

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4550332

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4650339

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4750353

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4850584

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

495060

0Ke

nya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect











Cluster 2

Cluster 1


50353

50332

50319

50296

50265

7

50290

11

50600

50339

50330

50328

50326

50299

50273

50264

30316

50259

45446

12

50584

50307

13

10

50325

16

15

50298

45451

14

6

31992

1

1959

4

45426

9

5

3

2

31990

10

2289

2241

1988

2249

2232

2279

2000

2299







Table 4
















Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy





Table3Con

tinued

Entry

Accession

no

Origin

Flow

ercolour

Stem

colour

Petio

lecolour

Stem

hairiness

Petio

leha

iriness

Leafcolour

Leafha

iriness

Growth

habit

2631990

Kenya

Purple

green

Green

Medium

Sparse

light

green

Sparse

Erect

2731992

Kenya

Pink

purple

Green

Profuse

Medium

dark

green

Medium

Erect

2845426

Kenya

Purple

Purple

Green

Profuse

Sparse

dark

green

Sparse

Erect

294544

6Ke

nya

White

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3045451

Kenya

Pink

Purple

Purple

Profuse

Profuse

dark

green

Medium

Erect

3150259

Kenya

Pink

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3250264

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3350265

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3450273

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3550290

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Semierect

3650296

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

Erect

3750298

Kenya

Purple

Purple

Purple

Medium

Sparse

light

green

Sparse

Semierect

3850299

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

Erect

3950307

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

Erect

4050319

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4150325

Kenya

Pink

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

4250326

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4350328

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4450330

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4550332

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4650339

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect

4750353

Kenya

Purple

Purple

Purple

Profuse

Medium

dark

green

Medium

erect

4850584

Kenya

Purple

Purple

Purple

Medium

Medium

dark

green

Sparse

erect

495060

0Ke

nya

Purple

Purple

Purple

Profuse

Medium

dark

green

Sparse

erect











Cluster 2

Cluster 1


50353

50332

50319

50296

50265

7

50290

11

50600

50339

50330

50328

50326

50299

50273

50264

30316

50259

45446

12

50584

50307

13

10

50325

16

15

50298

45451

14

6

31992

1

1959

4

45426

9

5

3

2

31990

10

2289

2241

1988

2249

2232

2279

2000

2299







Table 4
















Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy














Cluster 2

Cluster 1


50353

50332

50319

50296

50265

7

50290

11

50600

50339

50330

50328

50326

50299

50273

50264

30316

50259

45446

12

50584

50307

13

10

50325

16

15

50298

45451

14

6

31992

1

1959

4

45426

9

5

3

2

31990

10

2289

2241

1988

2249

2232

2279

2000

2299







Table 4
















Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy





Table 4
















Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy







Mean 418 58 12 8 97 684 85 551LSD 15 07 16 1 63 44 09 29

(plt005)CV 22 74 81 78 4 4 65 33










LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy












LL Season 1 02 17 19 893Season 2 02 17 19 893

LW Season 1 14 50 64 780Season 2 14 53 67 792



PH Season 1 82 9086 9168 991Season 2 82 9058 9140 991







Acknowledgments



References


































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy



































Journal of






Microbiology



Volume 2018



Volume 2018















BotanyJournal of










Agronomy




heritability analysis and phenotypic characterization of ...oar.icrisat.org/10826/1/heritability...

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