Research Article Genetic Variability and Selection ...downloads.hindawi.com/journals/tswj/2014/190531.pdf · Research Article Genetic Variability and Selection Criteria in Rice Mutant

Research ArticleGenetic Variability and Selection Criteria inRice Mutant Lines as Revealed by Quantitative Traits

Yusuff Oladosu1 M Y Rafii12 Norhani Abdullah13 Mohammad Abdul Malek14

H A Rahim5 Ghazali Hussin6 Mohammad Abdul Latif27 and Isiaka Kareem1

1 Institute of Tropical Agriculture Universiti Putra Malaysia (UPM) Serdang 43400 Selangor Malaysia2 Department of Crop Science Faculty of Agriculture Universiti Putra Malaysia (UPM) Serdang 43400 Selangor Malaysia3 Department of Biochemistry Faculty of Biotechnology and Biomolecular Science Universiti Putra Malaysia (UPM)Serdang 43400 Selangor Malaysia

4 Bangladesh Institute of Nuclear Agriculture Mymensingh 2202 Bangladesh5 Bioscience and Agrotechnology Division Malaysian Nuclear Agency Bangi 43000 Selangor Malaysia6 Strategic Livestock Research Centre Malaysian Agricultural Research and Development Institute (MARDI)Serdang 43400 Selangor Malaysia

7 Bangladesh Rice Research Institute (BRRI) Gazipur Bangladesh

Correspondence should be addressed to M Y Rafii mrafiiupmedumy

Received 5 July 2014 Revised 8 September 2014 Accepted 8 September 2014 Published 5 November 2014

Academic Editor Jean Louis Hilbert

Copyright copy 2014 Yusuff Oladosu et alThis is an open access article distributed under the Creative Commons Attribution Licensewhich permits unrestricted use distribution and reproduction in any medium provided the original work is properly cited

Genetic based knowledge of different vegetative and yield traits play a major role in varietal improvement of rice Genetic variationgives room for recombinants which are essential for the development of a new variety in any crop Based on this background thiswork was carried out to evaluate genetic diversity of derived mutant lines and establish relationships between their yield and yieldcomponents using multivariate analysis To achieve this objective two field trials were carried out on 45 mutant rice genotypesto evaluate their growth and yield traits Data were taken on vegetative traits and yield and its components while genotypic andphenotypic coefficients variance components expected genetic advance and heritability were calculated All the genotypes showedvariations for vegetative traits and yield and its components Also there was positive relationship between the quantitative traitsand the final yield with the exception of number of tillers Finally the evaluated genotypes were grouped into five major clustersbased on the assessed traits with the aid of UPGMA dendrogram So hybridization of group I with group V or group VI could beused to attain higher heterosis or vigour among the genotypes Also this evaluation could be useful in developing reliable selectionindices for important agronomic traits in rice

1 Introduction

Rice (Oryza sativa) is an important staple food in the worldDespite its position among the highly rated cereals it feedsmore than half of the worldrsquos population The geometricgrowth rate of the global population has called for yieldimprovement of this very important cereal On this sev-eral methods have been tried by scientists to combat thisperennial problem Some researchers have tried nutritionalmethod physiological method and breeding as well ascontrol of pests and diseases Among these methods it isestablished that breeding for high yield traits is the most

sustainable because the traits are heritable To achieve thisbreeding objective changes at cellular level are employed toget traits that could improve yieldThese changes are referredto as mutations and may occur either in the genes or atchromosomal level to bring the desired improvement Atpresent improvement of major food crops in the world restsmajorly on mutation This comes either naturally or throughirradiation So crops with restriction in genetic variationrequire mutagenesis or induced mutation to create desirableand heritable variations in them [1] Following this the useof induced mutations has been extensively used for geneticenhancement of different crops [2 3] Ionizing radiation of

Hindawi Publishing Corporatione Scientific World JournalVolume 2014 Article ID 190531 12 pageshttpdxdoiorg1011552014190531

2 The Scientific World Journal

high frequencies and chemical mutagens are being employedto achieve genetic enhancement among various mutagenicagents available to induce favourable mutations The mech-anism of operation of ionizing radiation hinges on the levelof the energy absorbed by the biological system of theplants or plant parts The primary system which is the mostimportant target in energy absorption is the chromosome [4]Through this there will be chromosomal changes which willmanifest themselves in the subsequent progenies The use ofseed irradiation to achieve chromosomal changes has beenfound to increase frequencies of mutation and promote generecombination which in turn promotes recombination andleads to mutation spectrum widening [5] Based on radiationmutation plant breeding has officially released over 2700 newcrop cultivars in approximately 170 species [6]

Development of high-yielding cultivars requires a thor-ough knowledge of genetic variation in yield and its com-ponent among the existing cultivars This is sine qua non tohigh-yielding cultivar development Any variation observedin organisms is the result of combination of estimate of bothgenetic and environmental causes Out of these two onlygenetic cause is heritable To get complete expectation ofplant response to selection only heritability estimate is notenough So it should be combined with other estimates likegenotypic and phenotypic coefficients of variation geneticadvance and change in mean value between generations [7]Extensive uses of gamma and x-rays in inducingmutations incrop plants have been well established Therefore successfuladoption of this technology requires knowing the optimumdose for irradiating plant materials At the same time soundmanagement of variations resulting from induction has to beembarked upon [8]

MR219 rice variety was developed by the MalaysianAgricultural Research and Development Institute (MARDI)and officially released in January 2001 It was the first varietyto be developed bymeans of a direct seeding planting systemThe emphasis was on the panicle characteristics These aremainly the grain size and number of grains per panicle Asa result a single grain of MR219 variety can weigh as much as28ndash30mg and the number of grains can be as high as 200This is higher than the previously released varieties Othercharacteristics of this variety include short life cycle (105ndash111days) fairly long but strong culms and resistance to blast andbacterial leaf blightTherefore it could bemarketed as a long-grain variety In addition cooked form of MR219 has softtexture (amylase content of 214) This is the preference ofmost local consumers It is the most cultivated variety grownin Malaysia covering almost 90 of cultivated area [9]

Breeders are interested in evaluating genetic diversitybased on morphological characteristics because they areinexpensive rapid and simple to score The study of thesecharacteristics does not require sophisticated equipmentMoreover they can be inherited without specific biochemicalor molecular techniques Also this evaluation could beuseful in developing reliable selection indices for importantagronomic traits in riceTherefore the present studywas con-ducted to evaluate genetic diversity of derived mutant linesof MR219 and establish relationship between yield and yieldcomponents in the mutant lines using multivariate analysis

2 Materials and Methods

21 Experimental Site and Plant Husbandry Because MR219is the most cultivated rice variety in Malaysia coveringalmost 90 of the cultivated areas [9] efforts are constantlybeing made to increase its yield potentials Therefore itsseeds were irradiated with an ion beam for radiosensitivitydetermination A total of 100 seeds were subjected to 010 20 40 60 80 100 120 160 and 200 Gray (Gy) todetermine the optimum doses for the production of highmutant frequency and spectrum The optimum dose wasfound to be 60Gy The M

1seedlings were transplanted into

the field with 25 cm times 25 cm planting distance Ten thousandM1seedlings were planted to produce M

2seeds and a total

of 5250 plants were selected from which 2 panicles perhill were randomly harvested from each hill About 5 ofM2populations were selected for further screening in M

3

After several series of selection and fixation 31 potentiallines with the required adaptive traits were recovered at M

4

generation during the 2009ndash2012 seasons (M0ndashM4) In total

forty-five rice genotypes which formed three populations(Malaysia Vietnam and Bangladesh) were analysed Ofthese 45 genotypes 31 weremutant lines derived fromMR2194 were other commercial varieties from Malaysia 8 werecommercial mutant varieties from Bangladesh and 2 weremutant varieties from Vietnam (Table 1) The field evaluationwas carried out repeatedly in two locations onM4 generationThe first location was field 10 of the Universiti Putra Malaysia(UPM) It was 31m above the sea level and is located betweenlatitude 3∘021015840N and longitude 101∘421015840E The second locationwas the farmersrsquo field in Melor Kelantan Malaysia It waslocated between latitude 59833∘N and longitude 1023∘ETheclimate of the two locations was a hot humid tropic Theareas fell under the rain forestThe periods of cultivationwereDecember 2012 to April 2013 and February to June 2013 30 gof each rice accession was placed in plastic petri dishes andoven-dried at 50∘C for 24 hours to break seed dormancyTheseeds were then pregerminated and finally transferred into anursery for proper establishment 21-day-old seedlings werethen transplanted into a field of size 22m by 18m The plantspacing was 25 cm by 30 cmwith density of two seedlings perstand This experiment was laid out in randomized completeblock (RCB) with three replications The fields were irrigatedthroughout the experiment with average of 10 cm waterabove the soil level Regular hand weeding was embarkedupon to free the plant of interspecific competition Fertilizerwas applied in splits using 57 kgha triple super phosphate42 kghamuriate of potash and 80 kgha urea at 15 35 55 and75 days after planting Insecticides (Malathion and HopperGun) were applied for controlling insect pests as required atrecommended rates

22 Data Collection From each variety five plants wererandomly sampled from each block for data collection Datacollection was on plant height flag leaf length to width rationumber of tillers per hill days to flowering days to maturitynumber of panicles per hill panicle length total number of

The Scientific World Journal 3

Table 1 List of rice genotypes studied

Genotypecode

Name ofaccession State Mutagenesis source

GN1 ML-1 Mutant line Ion beamGN2 ML-2 Mutant line Ion beamGN3 ML-3 Mutant line Ion beamGN4 ML-4 Mutant line Ion beamGN5 ML-5 Mutant line Ion beamGN6 ML-6 Mutant line Ion beamGN7 ML-7 Mutant line Ion beamGN8 ML-8 Mutant line Ion beamGN9 ML-9 Mutant line Ion beamGN10 ML-10 Mutant line Ion beamGN11 ML-11 Mutant line Ion beamGN12 ML-12 Mutant line Ion beamGN13 ML-13 Mutant line Ion beamGN14 ML-14 Mutant line Ion beamGN15 ML-15 Mutant line Ion beamGN16 ML-16 Mutant line Ion beamGN17 ML-17 Mutant line Ion beamGN18 ML-18 Mutant line Ion beamGN19 ML-19 Mutant line Ion beamGN20 ML-20 Mutant line Ion beamGN21 ML-21 Mutant line Ion beamGN22 ML-22 Mutant line Ion beamGN23 ML-23 Mutant line Ion beamGN24 ML-24 Mutant line Ion beamGN25 ML-25 Mutant line Ion beamGN26 ML-26 Mutant line Ion beamGN27 ML-27 Mutant line Ion beamGN28 ML-28 Mutant line Ion beamGN29 ML-29 Mutant line Ion beamGN30 ML-30 Mutant line Ion beamGN31 ML-31 Mutant line Ion beam

GN32 MR 219 Released variety Conventionalmethod




GN36 Binadhan-4 Released variety Gamma rayGN37 Binadhan-5 Released variety Gamma rayGN38 Binadhan-6 Released variety Gamma rayGN39 Binadhan-7 Released variety Gamma rayGN40 Binadhan-10 Released variety Gamma rayGN41 Binadhan-8 Released variety Gamma rayGN42 IRATOM-38 Released variety Gamma rayGN43 Binasail Released variety Gamma rayGN44 VN-124 Released variety Gamma rayGN45 VN-121 Released variety Gamma rayNB ML mutant line

grain per panicle 100 grain weight total grain weight perhill seed length seed length to width ratio and yield in tha(Table 2)

23 Variance Components Variance components were esti-mated to determine genetic variation among genotypes andto assess genetic and environmental effects on different traitsThe variance components were calculated as follows

24 Genotypic Variance (1205902119892) Consider

1205902

119892=

(MSG minusMSE)119903

(1)

whereMSG ismean square of genotypesMSE ismean squareof the error and 119903 is number of replications

25 Phenotypic Variance (1205902119901) Consider

1205902

119901= 1205902

119892+ 1205902

119890 (2)

where 1205902119892is genotypic variance and 1205902

119890is the mean squares of

error

26 Error Variance (1205902119890) Consider

1205902

119890= MSE (3)

where MSE is the mean square of error

27 Phenotypic and Genotypic Coefficient of Variation (119875119862119881and 119866119862119881) The estimates of phenotypic and genotypic coef-ficient of variation were calculated according to Singh andChoudhary [10] as follows

PCV =radic1205902

119875

119883

times 100

GCV =radic1205902

119892

119883

times 100

(4)

1205902

119875is the phenotypic variance 1205902

119892is the genotypic variance

119883 is the mean of the trait GCV and PCV values werecategorized as low (0ndash10) moderate (10ndash20) and high(20 and above) following Sivasubramanian and MadhavaMenon [11]

28 Heritability Estimate This heritability ℎ2119861(broad sense) is

the ratio of genetic variance (1205902119892) to phenotypic variance (1205902

119901)

[12] It is calculated as

ℎ2

119861=

1205902

119892

1205902

119901

(5)

where 1205902119892is the genotypic variance and 1205902

119901is the phenotypic

varianceThe heritability percentage was categorized as low (0ndash

30) moderate (30ndash60) and high (ge60) in accordancewith Robinson et al [13]


Table 2 List of quantitative traits

Growth traityield trait Denotation Method of evaluationPlant height PH (cm) The average height from the base to the tip of last leaf (flag leaf)Flag leaf length to width ratio FLWR (cm) Divide the flag leaf length by widthNumber of tillers per hill NT (number) Count the number of tillers per hillDays to flowering DF (days) Count the number of days from seeding to floweringDays to maturity DM (days) Count the number of days from seeding to maturityNumber of panicles per hill NPH (number) Count the number of panicles per hill

Panicle length PL (cm) Measure the length from the node below the lowest branch on the panicle to thetop of first superior spikelet

Total number of grain per panicle TNGP (number) Count the number of spikelets per panicleHundred grain weight 100GW (g) Weigh any 100 filled grainsTotal grain weight per hill TGWH (g) Weigh total grains produced per hill

Seed length SL (cm) Measure the length of the seeds using microscopic picture and Leica applicationsuit software

Seed length to width ratio SLWR (cm) Divide the seed length by widthYield in tha Yha (ton) Number of tillers per square meter times average total grains per tillers divided by 100

29 Estimated and Expected Genetic Advance Expectedgenetic advance (GA) (as percentage of the mean) wascalculated using the method of Assefa et al [14] and selectionintensity (119870) was assumed to be 5 Genetic advance wascategorized as low (0ndash10) moderate (10ndash20) and high(gt20) by following [15]

GA = 119870 times radic1205902119901times ℎ2

119861

GA () = 119870 timesradic1205902

119875

119883

times ℎ2

119861times 100

(6)

119870 is a constant which represents the selection intensityWhen119896 is 5 the value is 206 radic1205902

119875is phenotypic standard devia-

tion ℎ2119861is the heritability and119883 is the mean of traits

210 Cluster Analysis In this study cluster and principalcomponent analysis (PCA) were used to assess the geneticdiversity of quantitative traits [16] Cluster analysis groupedindividuals on the basis of their characteristics So individualshaving similar characteristics were mathematically clusteredtogether using distance similarity and relatedness of varietiesas the basis On the basis of distance clustering was done

211 Data Analysis All the morphological and yield datacollected were subjected to analysis of variance (ANOVA)while significant means were separated with least significantdifference (LSD) using SAS 91 software Also mean rangestandard deviation and coefficient of variation (CV) wererecorded for each trait measured The relationships amongthe traits were also determined using correlation analysisGenetic variance data generated were analysed based onEuclidian distance method Dicersquos and Jaccardrsquos similaritycoefficient Genetic relationships among the rice genotypeswere determined usingUPGMA algorithm and SAHNmeth-ods All these were achieved with NTSYS-pc 21 software

3 Results

31 Genetic Variation for Vegetative Characters From thepooled data of the two sites there were significant differencesamong yield components Five parameters related to plantvegetative growth were analysed for variation assessment(Table 3) The varietal impact on plant height was significantfor all the genotypes (119875 lt 001) The plant height variedfrom 14767 to 7174 cm The tallest plant (14767 cm) wasfrom Binasail whereas the shortest plant (7174 cm) was fromIRATOM-38 The genotypes MR 219 MR 220 ML-16 ML-11 ML-24 ML-7 ML-18 ML-19 ML-22 ML-25 ML-4 ML-27 ML-31 ML-30 ML-5 and ML-29 were similar in heightTheir height was an intermediate one The values for flag leaflength to width ratio (FLWR) ranged from 7289 to 2232The Binasail genotype had the highest value whereas VN121had the lowest value In the case of number of tillers (NT)per hill the values were between 26 and 14 The highestnumber of tillers (26) was from Binasail whereas the lowestnumber (14)was fromML-7 andML-12HoweverML-3ML-16 ML-24 ML-17 ML-6 ML-9 ML-30 ML-15 and ML-28were statistically equal to one another The number of daysto 50 flowering was between 53 and 77 days as observed inIRATOM-38 and ML7 respectively Days to maturity variedsignificantly (119875 lt 001) among the genotypes and range wasfrom 85 to 124 daysThe earliest maturing (85 days) genotypewas IRATOM-38 while genotype Binasail matured last (124days) ML-2 ML-5 ML-17 ML-29 ML-31 ML-1 ML-2ML-3 ML-16 ML-13 ML-18 ML-6 ML-21 ML-27 ML-20MR 220 and ML-9 had similar average number of days tomaturity

32 Yield and Its Components Eight parameters on rice yieldwere analysed for genetic variability (Table 3) The numberof panicles per hill varied from 21 to 13 The highest number


Table 3 Morphological traits yield and yield components of rice accession for the two location

Accession PH(cm)

FLR(cm) NT (no) DF

(days)DM(days)

NPH(no)

PL(cm)

TNGP(no)

100GW(g)

TGWH(g)

SL(mm)

SLWR(mm)

Yha(tha)

ML-1 8880 3417 195 725 1115 178 2417 1412 255 3326 752 222 532ML-2 8613 3927 158 720 1110 157 2283 1457 231 2660 718 213 426ML-3 8907 4032 178 728 1118 170 2417 1570 285 3022 806 234 484ML-4 8427 4737 145 718 1067 140 2317 2047 267 4221 765 229 672ML-5 8347 3886 162 722 1112 155 2400 1425 247 2112 776 225 338ML-6 9127 4109 173 737 1127 162 2433 1523 241 4048 795 232 648ML-7 8527 3965 140 765 1155 132 2467 1577 242 2880 768 226 461ML-8 8273 4044 185 753 1143 162 2467 1563 252 3227 763 227 516ML-9 8887 4589 173 757 1133 162 2450 1878 272 4455 777 225 713ML-10 9040 4826 158 733 1103 162 2383 2068 281 4079 760 230 653ML-11 8653 3879 152 753 1143 153 2417 1525 248 2946 732 214 471ML-12 8707 3398 140 758 1148 128 2350 1302 251 1791 729 215 287ML-13 8867 3554 183 733 1123 165 2383 1600 263 3965 762 219 634ML-14 8927 4584 167 700 1090 150 2283 1412 242 2947 786 240 472ML-15 8800 4222 172 718 1108 165 2433 1573 251 2681 745 220 429ML-16 8673 3537 178 733 1123 172 2533 1723 254 3291 778 232 527ML-17 8487 4395 173 722 1112 148 2417 1600 255 2871 776 227 459ML-18 8527 3673 152 735 1125 145 2500 1632 255 3168 788 236 507ML-19 8440 4082 163 730 1103 158 2433 1427 244 3372 767 230 540ML-20 8773 3997 197 735 1132 197 2467 1465 262 2797 811 234 448ML-21 8027 4415 188 755 1128 178 2467 1992 275 3979 788 234 637ML-22 8433 3866 165 748 1138 158 2450 1608 246 2732 782 218 437ML-23 8760 4231 167 732 1122 167 2467 1660 241 2950 761 221 472ML-24 8613 3646 177 748 1138 170 2417 1588 242 2854 758 224 457ML-25 8427 4054 180 760 1150 187 2417 1635 263 3848 757 225 616ML-26 8987 3495 193 765 1155 195 2450 1585 251 3207 801 234 513ML-27 8413 3762 188 740 1130 183 2450 1530 248 3683 773 226 589ML-28 9180 4172 172 750 1140 168 2467 1528 250 2950 799 240 472ML-29 8333 3937 183 722 1112 158 2467 1552 259 3173 795 235 508ML-30 8347 3950 173 753 1143 153 2467 1558 242 2734 831 236 438ML-31 8367 3613 168 723 1113 163 2450 1678 243 2923 800 231 468MR 219 8527 4039 157 760 1150 157 2333 1438 238 3052 785 234 488MR 220 8587 4108 168 743 1133 147 2467 1425 271 2874 787 233 460MR 253 8260 2929 168 672 1062 155 2467 1293 266 2662 785 222 426MR 264 8140 3667 158 717 1107 157 2450 1113 246 3484 791 232 558Binadhan-4 12850 5254 198 648 958 172 2667 1487 263 3427 752 206 548Binadhan-5 9947 4390 198 592 982 183 2283 1028 240 3118 665 198 499Binadhan-6 11050 4276 142 625 1015 155 2550 1392 227 3483 587 173 557Binadhan-7 7320 2763 160 555 945 155 2367 1227 224 2062 693 198 330Binadhan-10 8180 3408 210 612 1002 197 2317 912 254 3016 639 184 483Binadhan-8 9693 5431 183 553 943 195 2367 997 281 2838 568 171 454IRATOM-38 7147 3141 220 532 858 188 2267 1275 248 2133 645 183 341Binasail 14770 7289 262 642 1247 217 3133 2203 173 2484 512 167 398VN-124 7840 2298 225 577 967 213 2233 1112 239 2138 765 226 342VN-121 7900 2232 193 562 952 195 2350 1355 227 2929 736 214 469LSD (119875 le 005) 685 392 21 315 36 28 140 257 014 918 044 028 147


of panicles (21) was found in Binasail which was statisticallysimilar to Binadhan-5 Binadhan-8 IRATOM-38 VN121VNI24 and Binadhan-10The lowest number of panicles (13)was observed in ML12 (1283) which was statistically similarto ML7 However genotypes ML-29 ML-19 ML-22 ML-2MR 219 MR 264 MR 253 Binadhan-6 Binadhan-7 ML-5ML-30 ML-11 and ML-14 had similar number of paniclesThe differences in the number of panicles produced amongthe genotypes were significantly significant (119875 lt 001) Thepanicle length ranged from 3133 to 2233 cm and the highest(3133 cm) was observed in Binasail followed by Binadhan-4 (2667 cm) while the shortest panicle length (2233 cm)was recorded in VN124 which was statistically similar toBinadhan-5 ML-14 ML-2 and IRATOM-38 The highestnumber of filled grains (181) was from ML-21 while thelowest (65) was from Binadhan-7 A significant differencewas observed for grain per panicle among all varieties withvalues that ranged from 220 to 91 The highest number ofgrain per panicle (220) was observed in Binasail which wasstatistically similar to ML-10 and ML-4 but differed from therest of the genotypesThe lowest number of grains per panicle(91) was observed in Binadhan-10 The highest amount oftotal grain weight per hill belonged to ML-9 whereas ML-12 had the lowest amount of total grain weight per hill The100 grain weight varied significantly among the genotypeswith the weight ranging from 285 to 173 g ML-3 had thehighest value of 100 grain weight (285 g) which was notsignificantly different from ML-10 ML-21 ML-9 MR 220and Binadhan-8 The lowest 100 grain weight (173 g) wasrecorded in Binasail (Table 3) The seed length also variedsignificantly among the genotypes ranging from 831 to511mm ML-30 had the longest (831mm) seeds whereasBinasail had the shortest (511mm) seeds The grain shapevaried significantly among the genotypes with values rangingfrom 24 to 166mm The highest seed length to width ratio(24mm) was found in ML-14 whereas the lowest value(166mm) was observed in Binasail The highest yieldha(712 tonha) was found in genotype ML-9 followed by ML-4 (672 tonha) whereas the lowest value (287 tonha) wasobserved in ML-12 (Table 3)

33 Heritability Selection Gain and Coefficient of VariationIn this study broad sense heritability was high for all thetraits except total grainweight (28) yield per hectare (44)and seed length to width ratio (45) (Table 4) The highestheritability value (98) was found in 100 grain weightwhereas days to flowering days to maturity and plant heighthad 91 91 and 89 values respectively In additionhigh level of variability was observed in genotypic coefficientof variation (GCV) and phenotypic coefficient of variation(PCV) The high values of GCV were recorded from flag leaflength to width ratio (2831) and yield per hectare (2253)whereas the low values were found in panicle length (72)100 grain weight (949) and days to maturity (966) Thehighest PCV value was found in yield per hectare (3414)whereas the lowest value was recorded from panicle length(878) Furthermore the genetic advance (GA) for 13 traitshad its peak with the flag leaf length to width ratio (5577)

followed by the plant height (3857) whereas the lowestvalue were recorded from the panicle length (1215) andfrom seed length to width ratio (1418) (Table 4)

34 Relationship between Traits Correlation coefficientsamong morphological traits and yield and its componentsshowed that all the traits had a positive correlation with yieldexcept for the number of tillers (Table 5) The total grainweight significantly correlated with the flag leaf length towidth ratio days to flowering days to maturity leaf area andpanicle length (Table 5)

35 Cluster Analysis of Morphological Traits The standard-ized morphological data were employed to calculate theEuclidean distances among the 45 genotypes and a UPGMAdendrogram was constructed using these values as indicatedin Table 6 and Figure 1 In this dendrogram the 45 ricegenotypes were grouped into 5 primary clusters based on13 morphological traits Among the five clusters cluster Ihad the largest number of genotypes (35) cluster II had 2genotypes and cluster III had 3 genotypes Cluster IV had 4genotypes whereas cluster V contained only a genotype

4 Discussion

The traits assessed in 45 rice accessions showed significantdifferences among themselves This pointed at the existenceof variation in the population The differences exhibitedby the genotypes could be because they originated fromdifferent areas In this direction several reports have beenpublished on phenotypic variation among rice genotypes [17ndash20] Similarly Pandey et al [21] reported highly significantdifferences among 40 rice accessions with the use of 12quantitative characters In the same vein [22] discovered 95differences among five rice populations by using 20 morpho-logical characters The high-yielding genotypes were shortThis feature was as a result of short internode This couldequally be attributed to very effective assimilate partitioningat the expense of vegetative growth So instead of having tallplants high yield came as a compensation for the vegetativedeficiency This trait is also advantageous in protectionagainst lodging Though plant height is mostly governed bythe genetic makeup of the genotype it is highly influencedby environmental factors As indirectly pointed out earlierrice yield is indirectly related to its height This is due tosink competition for the limited photosynthates produced bylimited sources So what will be used for yield increase will beunnecessarily used for somatic cell enlargement that results inluxuriant vegetative growth and enhanced height Thereforetall varieties normally have lower yield than the short onesAnother serious disadvantage of tallness rice is lodgingwhichsignificantly lowers the final yield and makes the plantsprone to some other natural attacks In this experiment allthe high-yielding varieties were found to be of intermediateheight This implies that moderate plant height is desirablewhen breeding for high-yielding varieties Flag leaf plays asignificant role in enhancing rice yield because it remains theonly source of assimilate production for the filling spikelets


Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob

abilitylevellowastlowasthigh

lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh

eritabilityGAgeneticadvance(aspercentage

ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)


Table5Correlationcoeffi

cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA

PH062741lowastlowast028782lowastlowastminus01204lowast

00958

016758lowast041755lowastlowast01498lowastminus035534lowastlowast

006722minus042857lowastlowastminus013489lowast

006

656

FLR

013341lowast

006756

02601lowastlowast

006

049041032lowastlowast031058lowastlowastminus012583lowast018743lowastminus029327lowastlowastminus012007lowast01891lowast

NT

minus029697lowastlowastminus013746lowast074554lowastlowast025826lowastlowast

0114

84minus019125lowastminus000905minus02171lowastlowastminus030109lowastlowastminus000893

DF

085622lowastlowastminus024661lowastlowastminus005954025831lowastlowast020724lowastlowast019197lowast050885lowastlowast017302lowast019181lowast

DM

minus013274lowast013885lowast030822lowastlowastminus009126012728lowast029584lowastlowast

011109

012724

NP

031088lowastlowast02133lowast

minus01037

005901minus021643lowastminus042423lowastlowast

005928

PL054372lowastlowastminus024504lowastlowast011983lowastminus014073lowastminus040921lowastlowast

0117

2TN

Gminus004898022772lowast

009622minus024709lowastlowast022528lowast

GW

021795039876lowastlowast

0118


TGW

015764lowastlowast

006


SL033344lowastlowast015588lowast

SLWR

006269

Notelowastsig

nificanta

t005prob

ability

levellowastlowasthigh

lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha


Table 6 Grouping of 45 genotypes according to cluster analysis

Cluster number Number of genotypes Genotypes

I 35GN1 GN2 GN3 GN4 GN5 GN6 GN7 GN8 GN9 GN10 GN11 GN12 GN13GN14 GN15 GN16 GN17 GN18 GN19 GN20 GN21 GN22 GN23 GN24GN25 GN26 GN27 GN28 GN29 GN30 GN31 GN32 GN33 GN34 GN35

II 2 G36 G38III 3 G37 G40 G41IV 4 G39 G42 G44 G45V 1 G43Note GN1 = ML-1 GN2 = ML-2 GN3 = ML-3 GN4 = ML-4 GN5 = ML-5 GN6 = ML-6 GN7 = ML-7 GN8 = ML-8 GN9 = ML-9 GN10 = ML-10 GN11 =ML-11 GN12 = ML-12 GN13 = ML-13 GN14 = ML-14 GN15 = ML-15 GN16 = ML-16 GN17 = ML-17 GN18 = ML-18 GN19 = ML-19 GN20 = ML-20 GN21= ML-21 GN22 =ML-22 GN23 =ML-23 GN24 =ML-24 GN25 =ML-25 GN26 =ML-26 GN27 =ML-27 GN28 =ML-28 GN29 =ML-29 GN30 =ML-30GN31 = ML-31 GN32 = MR 219 GN33 = MR 220 GN34 = MR 253 GN35 = MR 264 GN36 = Binadhan-4 GN37 = Binadhan-5 GN38 = Binadhan-6 GN39= Binadhan-7 GN40 = Binadhan-10 GN41 = Binadhan-8 GN42 = IRATOM-38 GN43 = Binasail GN44 = VN-124 and GN45 = VN-121

GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient

Figure 1 Clustering pattern of the morphological trait at dissimilarity coefficient of 143

during grain-filling stage [23 24]The larger the leaf area themore the solar interception and photosynthate productionprovided that all other factors of production are not limitingTherefore flag leaf area was found to be directly related tothe yield components number of panicles panicle lengthnumber of grains per panicle 100 grain weight total grainweight per hill and yield per hectare Furthermore the flagleaf has been found to be metabolically active to supporthigher grain yield Corroborating our finding in this workAshrafuzzaman et al [25] have made it clear that yield

components positively correlate with flag leaf area Numberof tillers plays a significant role in determining yield of therice grain since it is directly related to panicle number thatwill be produced per unit ground area Fewer tillers resultin fewer panicles excess tillers cause high tiller abortionssmall panicles poor grain filling and reduction in grainyield [21] It has been observed that leaf area index andplant nitrogen status are the two major factors that affecttiller production in rice crops [21] When there is adequatenutrient supply mitotic cell division will be enhanced and


growth of tillers and plant general vegetative life will receivea boost In this work the tiller production was betweenmoderate and low levels So the case of tiller abortion wasnot a problem during production period The number ofpanicles per hill was between moderate (21) and low (13)This correlates with the number of tillers produced So thenumber of effective tillers rests on the number of tillersproduced and this is directly proportional to the paniclesproduced per unit area and finally depends on variety [26]The panicle number is an important character which directlyinfluences the yieldTherefore yield could be increased whenagronomic manipulation is used to increase the number ofpanicles produced per unit area Among yield contributingcharacters is panicle length This is proportional to thenumber of potential spikelets to be filled during grain-fillingstage

Rice plants will naturally have the superior spikeletsfilled before the inferior ones The superior spikelets arelocated on the primary branches located at the apex ofthe rice plant So they fill faster and produce heavier andlarger grain weight On the other hand inferior spikelets areusually located on secondary branches and are usually slowin filling They also produce poorly filled spikelets or sterileones [27] Slow grain-filling rate and low grain weight ofinferior spikelets have often been attributed to limitation incarbohydrate supply [27] According to [28] the fundamentalfactors responsible for variations in grain filling between thesuperior and inferior spikelets remain unknown As it couldbe seen from this study some varieties flower earlier thanthe others Those that flowered earlier matured early whilethose that flowered late had a delay in their maturity Earlyflowering indicates short life cycle and is considered a positivecharacter for rice improvement Early maturing varieties areadvantageous in areas with short rainfall duration becausethey grow faster during the vegetative phase and are thusmore competitive withweedsThey reduceweed control costsand utilize less water [29] It is a known fact that droughtfrequently impedes production of rain-fed lowland rice [30]So when drought occurs towards the reproductive stage ofrice production pollination and fertilization as well as grainfilling are severely affected and panicle blanking may resultIn the situation early maturing variety will give remedialmeasures in lieu of establishment of irrigation facilities anddevelopment of drought-tolerant varieties [30]

The number of filled grains among the varieties rangedfromhigh (181) to low (65) Increase in number of filled grainscould be attributed to efficient translocation of carbohydratesfrom the sources to the spikelets (sinks) which consequentlyleads to increase in grain yield [31] The varietal yield inthis work was between high and low Yield differencesamong different rice varieties have been reported anytime acomparison ismade between different varieties of rice in bothfield and glasshouse trials [32]The differences are geneticallybased though environment has a great contribution in themanifestation of the inherent potential In this work thegenotypes with higher number of effective tillers as well ashigher number of grains per panicle also had higher yieldThis is in line with the observation of [33] The weight of100 or 1000 grain weight contributes significantly to the final

yield per unit area It represents the weight of individual seedswhich could not be directly measured because of the sizeof individual seeds The result of the present study showedthat 100 grain weight varied significantly among the testedvarieties This could also be due to their differences in originand genetic makeup Similar reports have been published bya BRRI scientist [34] as well as Ashrafuzzaman et al [25]Panicle length determines how many spikelets will be foundin a panicle and therefore filled spikelets and consequentlyfinal grain yieldThe longer the panicle themore the spikeletsand the filled grains if other environmental conditionsare not limiting As found here panicle length correlatedpositively with the final yield Credence to this researchfinding has been laid by [35] who also found a significantpositive association between panicle length and grain yieldper hill

Heritability is the proportion of phenotypic traits (phys-ical appearance) or total variance that is inherited from theparents Higher genotypic coefficient of variation togetherwith high heritability as well as high genetic advance givesbetter clues than individual parameters Thus the traitswith high genotypic coefficient of variation heritability andgenetic advance are selected In this study flag leaf length towidth ratio plant height and the total number of grains perpanicle had higher values for genotypic coefficient of vari-ation heritability and genetic advance Therefore selectionwith a view to develop one trait whichwill positively influenceother traits is of paramount importance

Having positive correlation coefficients between the veg-etative traits and the yield parameters in this work wasan indication that the quantitative traits measured wereappropriate for yield prediction and selection for breedingprogrammes to obtain better vigour or heterosisThe positivecorrelation between the final yield and plant height and totalnumber of grain per panicle and days to maturity indicatesthat better exploration of these three traits could be used todevelop desired genotypes

The contribution of individual panicle grain yield sumsup to produce the final yield Therefore high panicle grainyield could be successfully used as an important selectionindex for grain yield [36] Furthermore when the panicleyield is correlated with the yield per unit area positive cor-relation coefficient will result [37] In the present study yieldper panicle had the best relationship with yield productionHence more attention is given to this trait for final yielddetermination in all the assessed genotypes

The UPGMA dendrogram broadly clustered the ricegenotypes into five major groups at 143 dissimilarity coef-ficients This implies a high level of morphological diversityin the rice genotypes This study revealed effectiveness ofquantitative or morphological traits in grouping rice geno-types It has been established that genetic divergence analysisamong rice genotypes on the basis of morphological traitscan be used to classify and differentiate different genotypesin a population [38] This genetic divergence analysis alsoplays a vital role in selection of diverse genotypes for thefurther improvement of rice varieties through breeding [39]We have succeeded in clustering 45 genotypes into 5 majorgroups using some quantitative traits for better handling of


the diverse genotype as well as better selection for acquisitionof higher heterotic vigour in the resulting offspring aftercrossing Similarly Ahmadikhah et al [40] clustered 58 ricevarieties into four groups using 18morphological traits whileMazid et al [41] had his 41 rice genotypes clustered into 6on the basis of 13 morphological traits Therefore to attainsuccess in breeding for higher vigour among the genotypesgroup I could be hybridized with group V or group VI in thiswork and as corroborated by findings of [41 42]

As observed in this study the experimented rice geno-types showed notable genetic diversity in terms of morpho-logical traitsThis shows the efficacy of ion beam radiation ininducing high performing agronomic traits in some mutantlines as compared to the control High heritability togetherwith high genetic advance was obtained for plant height flagleaf length to width ratio days to maturity and total numbersof grains per panicle

5 Conclusion

The development of rice genotypes that could lead to attain-ment of food security rests on exploring genetic aspects ofits quantitative traits This is because genetic variations giveroom for recombinants which are essential in development ofnew genotype Field trials were carried out on 45 mutant ricegenotypes and their genetic diversitieswere determined usingtheir quantitative traits All the genotypes showed variationsfor vegetative and yield traits Also all the traits relatedpositively with the final yield with the exception of numberof tillers Finally the evaluated genotypes were grouped intofive major clusters based on the assessed traits with theaid of UPGMA dendrogram So hybridization of group Iwith group V or group VI could be used to attain higherheterosis or vigour among the genotypes This evaluationcould be useful in developing reliable selection indices forimportant agronomic traits in rice It is recommended thatfuture research explore molecular means to further confirmthe outcome of this research and establish the relationshipbetween the two methods

Conflict of Interests

The authors declare that there is no conflict of interestsregarding the publication of this paper

Acknowledgments

The authors would like to acknowledge the financial sup-port of IAEArsquos Coordinated Research Project (CRP) forFood and Feed [CRP Code D23030] We also appreciateMalaysia Ministry of Education for Long-Term ResearchGrant Scheme (LRGS) on enhancing sustainable rice produc-tion and MOSTI for Science Fund (06-03-01-SF0110)

References

[1] M A J Parry P J Madgwick C Bayon et al ldquoMutationdiscovery for crop improvementrdquo Journal of ExperimentalBotany vol 60 no 10 pp 2817ndash2825 2009

[2] L Velasco J M Fernandez-Martınez and A De Haro ldquoInheri-tance of reduced linolenic acid content in the Ethiopianmustardmutant N2-4961rdquo Plant Breeding vol 121 no 3 pp 263ndash2652002

[3] A M R Ferrie D C Taylor S L MacKenzie G Rakow J PRaney and W A Keller ldquoMicrospore mutagenesis of Brassicaspecies for fatty acid modifications a preliminary evaluationrdquoPlant Breeding vol 127 no 5 pp 501ndash506 2008

[4] A M van Harten Mutation Breeding Theory and PracticalApplications Cambridge University Press Cambridge UK1998

[5] A S Larik S Memon and Z A Soomro ldquoRadiation inducedpolygenic mutations in Sorghum bicolor Lrdquo Journal of Agricul-tural Research vol 47 pp 11ndash19 2009

[6] P J L Lagoda ldquoNetworking and fostering of cooperationin plant mutation genetics and breeding role of the jointFAOIAEA programme FAOIEAErdquo in Proceedings of the Inter-national Symposium on Induced Mutations in Plants pp 2ndash3Vienna Austria August 2008

[7] R M Wani and S Khan ldquoEstimates of genetic variabilityin mutated populations and the scope of selection for yieldattributes in Vigna radiata (L) Wilczekrdquo Egyptian Journal ofBiology vol 8 pp 1ndash6 2006

[8] B S Ahloowalia and M Maluszynski ldquoInduced mutationsmdashanew paradigm in plant breedingrdquo Euphytica vol 118 no 2 pp167ndash173 2001

[9] Z U Bashar A Wayayok and A M S Mohd ldquoDeterminationof some physical properties of common Malaysian rice MR219seedsrdquo Australian Journal of Crop Science vol 8 no 3 pp 332ndash337 2014

[10] R K Singh and B D Choudhary Biometrical Methods inQuantitative Genetic Analysis Kalyani Publishers New DelhiIndia 1985

[11] S S Sivasubramanian and P Madhava Menon ldquoGenotypic andphenotypic variability in ricerdquoMadras Agricultural Journal vol60 pp 1093ndash1096 1973

[12] D S Falconer Introduction to Quantitative Genetics LongmanNew York NY USA 3rd edition 1989

[13] H F Robinson R E Comstock and P H Harvey ldquoGenotypicand phenotypic correlation in corn and their implications inselectionrdquo Agronomy Journal vol 43 pp 282ndash287 1949

[14] K Assefa S Ketema H Tefera et al ldquoDiversity amonggermplasm lines of the Ethiopian cereal tef [Eragrostis tef(Zucc) Trotter]rdquo Euphytica vol 106 no 1 pp 87ndash97 1999

[15] H W Johnson H F Robinson and R E Comstock ldquoEsti-mation of genetic and environmental variability in soybeansrdquoAgronomy Journal vol 47 pp 314ndash318 1955

[16] A Melchinger M Messmer M Lee W Woodman and KLamkey ldquoDiversity and relationships amongUSmaize inbreedsreveal by restriction fragment length polymorphismsrdquo CropScience vol 31 no 3 pp 621ndash678 1991

[17] T Shehzad A Allah A El Naby A Allah M Ammar and AAbdelkhalik ldquoAgronomic and molecular evaluation of inducedmutant rice (Oryza sativa L) lines in Egyptrdquo Pakistan Journal ofBotany vol 43 no 2 pp 1183ndash1194 2011

[18] A Babaei G-A Nematzadeh and H Hashemi ldquoAn evaluationof genetic differentiation in rice mutants using semi-randommarkers and morphological characteristicsrdquo Australian Journalof Crop Science vol 5 no 13 pp 1715ndash1722 2011

[19] K Elayaraja M Prakash K Saravanan S B Kumar andJ Ganesan ldquoStudies on variability heritability and genetic


advance for certain quantitative characters in mutant popula-tion of rice (Oryza sativa L)rdquo Crop Research Hisar vol 1 pp134ndash137 2005

[20] S S Meheter A P Patil R CMahajan and B R Shinde ldquoVari-ability heritability character association and genetic divergencestudies in M2 generation of gamma irradiation upland paddyrdquoCrop Research Hisar vol 12 no 2 pp 155ndash161 1996

[21] P Pandey P John Anurag D K Tiwari S K Yadav andB Kumar ldquoGenetic variability diversity and association ofquantitative traits with grain yield in rice (Oryza sativa L)rdquoJournal of Bio-Science vol 17 no 1 pp 77ndash82 2009

[22] S PadmajaRao ldquoInfluence of source and sink on the productionof high density grain and yield in ricerdquo Indian Journal of PlantPhysiology vol 34 pp 339ndash348 1991

[23] A Raj and M P Tripathi ldquoVarietal variations in flag leaf areaand yield in deep water ricerdquo Indian Journal of Plant Physiologyvol 5 no 3 pp 293ndash295 2000

[24] J-L Wang Y-B Gao N-X Zhao et al ldquoMorphological andRAPD analysis of the dominant species Stipa krylovii Roshevin Inner Mongolia stepperdquo Botanical Studies vol 47 no 1 pp23ndash35 2006

[25] M Ashrafuzzaman M R Islam M R Ismail S M Shahidul-lah andMM Hanafi ldquoEvaluation of six aromatic rice varietiesfor yield and yield contributing charactersrdquo International Jour-nal of Agriculture and Biology vol 11 no 5 pp 616ndash620 2009

[26] M B Hossain M O Islam and M Hasanuzzaman ldquoInfluenceof different nitrogen levels on the performance of four aromaticrice varietiesrdquo International Journal of Agriculture and Biologyvol 10 no 6 pp 693ndash696 2008

[27] J Yang and J Zhang ldquoGrain filling of cereals under soil dryingrdquoNew Phytologist vol 169 no 2 pp 223ndash236 2006

[28] A Mostajeran and V Rahimi-Eichi ldquoEffects of drought stresson growth and yield of rice (Oryza sativa L) cultivars andaccumulation of proline and soluble sugars in sheath and bladesof their different ages leavesrdquo American-Eurasian Journal ofAgricultural amp Environmental Science vol 5 pp 264ndash272 2009

[29] G S Khush ldquoModern varietiesmdashtheir real contribution to foodsupply and equityrdquoGeoJournal vol 35 no 3 pp 275ndash284 1995

[30] S M Haefele and B A M Bouman ldquoDrought-prone rainfedlowland rice in Asia limitations and management options inrdquoin Drought Frontiers in Rice Crop Improvement for IncreasedRainfed Production J Serraj J Bennett and B Hardy Eds pp211ndash232 World Scientific Publishing Singapore 2009

[31] Z-Z Xu and G-S Zhou ldquoPhotosynthetic recovery of aperennial grass Leymus chinensis after different periods of soildroughtrdquo Plant Production Science vol 10 no 3 pp 277ndash2852007

[32] J K Biswas M A Hossain B C Sarker M Hassan and MZ Haque ldquoYield performance of several rice varieties seededdirectly as late Aman cropsrdquo Bangladesh Journal of Life Sciencesvol 10 pp 47ndash52 1998

[33] R K DuttaM A Basetmai and S Khanam ldquoPlant architectureand growth characteristics of fine grain and aromatic rice andtheir relation with grain yieldrdquo IRC Newsletter vol 51 pp 105ndash111 2001

[34] BRRI Annual Report 1996-1997 Bangladesh Rice ResearchInstitute Gazipur Bangladesh 1997

[35] R Chakraborty and S Chakraborty ldquoGenetic variability andcorrelation of some morphometric traits with grain yield inbold grained rice (Oryza sativa L) gene pool of Barak valleyrdquoAmerican-Eurasian Journal of Sustainable Agriculture vol 4 no1 pp 26ndash29 2010

[36] T Meenakshi A A D Ratinam and S Backiyarani ldquoCor-relation and path analysis of yield and some physiologicalcharacters in rain fed ricerdquo Oryza vol 6 pp 154ndash156 1999

[37] M A Mustafa and M A Y Elsheikh ldquoVariability correlationand path cominusefficient analysis for yield and its components inricerdquo African Crop Science Journal vol 15 no 4 pp 183ndash1892007

[38] J Franco J Crossa J M Ribaut J Bertran M L Warburtonand M Khairallah ldquoA method for combining molecular mark-ers and phenotypic attributes for classifying plant genotypesrdquoTheoretical and Applied Genetics vol 103 no 6-7 pp 944ndash9522001

[39] S M Shahidullah M M Hanafi M Ashrafuzzaman M RaziIsmail M A Salam and A Khair ldquoBiomass accumulationand energy conversion efficiency in aromatic rice genotypesrdquoComptes Rendus Biologies vol 333 no 1 pp 61ndash67 2010

[40] A Ahmadikhah S Nasrollanejad and O Alishah ldquoQuantita-tive studies for investigating variation and its effect on heterosisof ricerdquo International Journal of Plant Production vol 2 pp 297ndash308 2008

[41] M SMazidM Y RafiiMMHanafiHA RahimM Shaban-imofrad andMA Latif ldquoAgro-morphological characterizationand assessment of variability heritability genetic advance anddivergence in bacterial blight resistant rice genotypesrdquo SouthAfrican Journal of Botany vol 86 pp 15ndash22 2013

[42] M A Latif M M Rahman M S Kabir M A Ali M T Islamand M Y Rafii ldquoGenetic diversity analyzed by quantitativetraits among rice (Oryza sativa L) genotypes resistant to blastdiseaserdquo African Journal of Microbiology Research vol 5 no 25pp 4383ndash4391 2011

Submit your manuscripts athttpwwwhindawicom

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Anatomy Research International

PeptidesInternational Journal of


Hindawi Publishing Corporation httpwwwhindawicom

International Journal of

Volume 2014

Zoology


Molecular Biology International

GenomicsInternational Journal of


The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014


BioinformaticsAdvances in

Marine BiologyJournal of



Signal TransductionJournal of


BioMed Research International

Evolutionary BiologyInternational Journal of



Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014


Genetics Research International


Advances in

Virolog y

Hindawi Publishing Corporationhttpwwwhindawicom

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational



Enzyme Research



Microbiology


high frequencies and chemical mutagens are being employedto achieve genetic enhancement among various mutagenicagents available to induce favourable mutations The mech-anism of operation of ionizing radiation hinges on the levelof the energy absorbed by the biological system of theplants or plant parts The primary system which is the mostimportant target in energy absorption is the chromosome [4]Through this there will be chromosomal changes which willmanifest themselves in the subsequent progenies The use ofseed irradiation to achieve chromosomal changes has beenfound to increase frequencies of mutation and promote generecombination which in turn promotes recombination andleads to mutation spectrum widening [5] Based on radiationmutation plant breeding has officially released over 2700 newcrop cultivars in approximately 170 species [6]

Development of high-yielding cultivars requires a thor-ough knowledge of genetic variation in yield and its com-ponent among the existing cultivars This is sine qua non tohigh-yielding cultivar development Any variation observedin organisms is the result of combination of estimate of bothgenetic and environmental causes Out of these two onlygenetic cause is heritable To get complete expectation ofplant response to selection only heritability estimate is notenough So it should be combined with other estimates likegenotypic and phenotypic coefficients of variation geneticadvance and change in mean value between generations [7]Extensive uses of gamma and x-rays in inducingmutations incrop plants have been well established Therefore successfuladoption of this technology requires knowing the optimumdose for irradiating plant materials At the same time soundmanagement of variations resulting from induction has to beembarked upon [8]

MR219 rice variety was developed by the MalaysianAgricultural Research and Development Institute (MARDI)and officially released in January 2001 It was the first varietyto be developed bymeans of a direct seeding planting systemThe emphasis was on the panicle characteristics These aremainly the grain size and number of grains per panicle Asa result a single grain of MR219 variety can weigh as much as28ndash30mg and the number of grains can be as high as 200This is higher than the previously released varieties Othercharacteristics of this variety include short life cycle (105ndash111days) fairly long but strong culms and resistance to blast andbacterial leaf blightTherefore it could bemarketed as a long-grain variety In addition cooked form of MR219 has softtexture (amylase content of 214) This is the preference ofmost local consumers It is the most cultivated variety grownin Malaysia covering almost 90 of cultivated area [9]

Breeders are interested in evaluating genetic diversitybased on morphological characteristics because they areinexpensive rapid and simple to score The study of thesecharacteristics does not require sophisticated equipmentMoreover they can be inherited without specific biochemicalor molecular techniques Also this evaluation could beuseful in developing reliable selection indices for importantagronomic traits in riceTherefore the present studywas con-ducted to evaluate genetic diversity of derived mutant linesof MR219 and establish relationship between yield and yieldcomponents in the mutant lines using multivariate analysis

2 Materials and Methods

21 Experimental Site and Plant Husbandry Because MR219is the most cultivated rice variety in Malaysia coveringalmost 90 of the cultivated areas [9] efforts are constantlybeing made to increase its yield potentials Therefore itsseeds were irradiated with an ion beam for radiosensitivitydetermination A total of 100 seeds were subjected to 010 20 40 60 80 100 120 160 and 200 Gray (Gy) todetermine the optimum doses for the production of highmutant frequency and spectrum The optimum dose wasfound to be 60Gy The M

1seedlings were transplanted into

the field with 25 cm times 25 cm planting distance Ten thousandM1seedlings were planted to produce M

2seeds and a total

of 5250 plants were selected from which 2 panicles perhill were randomly harvested from each hill About 5 ofM2populations were selected for further screening in M

3

After several series of selection and fixation 31 potentiallines with the required adaptive traits were recovered at M

4

generation during the 2009ndash2012 seasons (M0ndashM4) In total

forty-five rice genotypes which formed three populations(Malaysia Vietnam and Bangladesh) were analysed Ofthese 45 genotypes 31 weremutant lines derived fromMR2194 were other commercial varieties from Malaysia 8 werecommercial mutant varieties from Bangladesh and 2 weremutant varieties from Vietnam (Table 1) The field evaluationwas carried out repeatedly in two locations onM4 generationThe first location was field 10 of the Universiti Putra Malaysia(UPM) It was 31m above the sea level and is located betweenlatitude 3∘021015840N and longitude 101∘421015840E The second locationwas the farmersrsquo field in Melor Kelantan Malaysia It waslocated between latitude 59833∘N and longitude 1023∘ETheclimate of the two locations was a hot humid tropic Theareas fell under the rain forestThe periods of cultivationwereDecember 2012 to April 2013 and February to June 2013 30 gof each rice accession was placed in plastic petri dishes andoven-dried at 50∘C for 24 hours to break seed dormancyTheseeds were then pregerminated and finally transferred into anursery for proper establishment 21-day-old seedlings werethen transplanted into a field of size 22m by 18m The plantspacing was 25 cm by 30 cmwith density of two seedlings perstand This experiment was laid out in randomized completeblock (RCB) with three replications The fields were irrigatedthroughout the experiment with average of 10 cm waterabove the soil level Regular hand weeding was embarkedupon to free the plant of interspecific competition Fertilizerwas applied in splits using 57 kgha triple super phosphate42 kghamuriate of potash and 80 kgha urea at 15 35 55 and75 days after planting Insecticides (Malathion and HopperGun) were applied for controlling insect pests as required atrecommended rates

22 Data Collection From each variety five plants wererandomly sampled from each block for data collection Datacollection was on plant height flag leaf length to width rationumber of tillers per hill days to flowering days to maturitynumber of panicles per hill panicle length total number of



Genotypecode











1205902

119892=


(1)



1205902

119901= 1205902

119892+ 1205902

119890 (2)



error


1205902

119890= MSE (3)



PCV =radic1205902

119875

119883

times 100

GCV =radic1205902

119892

119883

times 100

(4)

1205902






119901)


ℎ2

119861=

1205902

119892

1205902

119901

(5)














119861

GA () = 119870 timesradic1205902

119875

119883

times ℎ2

119861times 100

(6)






3 Results





Accession PH(cm)

FLR(cm) NT (no) DF

(days)DM(days)

NPH(no)

PL(cm)

TNGP(no)

100GW(g)

TGWH(g)

SL(mm)

SLWR(mm)

Yha(tha)








4 Discussion



Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob


lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh


ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology



Genotypecode











1205902

119892=


(1)



1205902

119901= 1205902

119892+ 1205902

119890 (2)



error


1205902

119890= MSE (3)



PCV =radic1205902

119875

119883

times 100

GCV =radic1205902

119892

119883

times 100

(4)

1205902






119901)


ℎ2

119861=

1205902

119892

1205902

119901

(5)














119861

GA () = 119870 timesradic1205902

119875

119883

times ℎ2

119861times 100

(6)






3 Results





Accession PH(cm)

FLR(cm) NT (no) DF

(days)DM(days)

NPH(no)

PL(cm)

TNGP(no)

100GW(g)

TGWH(g)

SL(mm)

SLWR(mm)

Yha(tha)








4 Discussion



Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob


lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh


ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology










119861

GA () = 119870 timesradic1205902

119875

119883

times ℎ2

119861times 100

(6)






3 Results





Accession PH(cm)

FLR(cm) NT (no) DF

(days)DM(days)

NPH(no)

PL(cm)

TNGP(no)

100GW(g)

TGWH(g)

SL(mm)

SLWR(mm)

Yha(tha)








4 Discussion



Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob


lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh


ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology



Accession PH(cm)

FLR(cm) NT (no) DF

(days)DM(days)

NPH(no)

PL(cm)

TNGP(no)

100GW(g)

TGWH(g)

SL(mm)

SLWR(mm)

Yha(tha)








4 Discussion



Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob


lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh


ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology







4 Discussion



Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob


lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh


ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology


Table4Meansquareso

fmorph

ologicaltraits

yieldandyieldcompo

nent

ofther

icea

ccessio

n

SOV

Df

PHcm

FLWR

cmNT(no)

DFdays

DMdays

NPH(no

)PL

(cm

)TN

GP(no

)TG

WH

(g)

100G

W(g)

SL(mm)

SLWR

(mm)

Yha(th

a)Re

psSite

(RS)

496843lowastlowast

5810

2lowastlowast

3290lowastlowast

8446lowastlowast

8092lowastlowast

512lowastlowast

35

28037

672

001lowastlowast

042

023

1Genotypes

(G)44

95561lowastlowast

39316lowastlowast

3295lowastlowast

26579lowastlowast

34421lowastlowast

232lowastlowast

107lowastlowast

28899lowastlowast

1549

017lowastlowast

264lowastlowast

021lowastlowast

53lowastlowast

Sites(S)

1432

41323lowastlowast

21690lowastlowast

48267lowastlowast

2296

3lowastlowast

6045lowastlowast

5808lowastlowast

1255395lowastlowast

2901

0001

0001

3167lowastlowast

25

GtimesS

44001

619

1635lowastlowast

5238lowastlowast

3037lowastlowast

142lowastlowast

29

8674

03

0001

0001

082lowastlowast

04

Error

176

3617

1184

334

766

987

58

154594

706

0001

015

006

16Mean

8844

3983

1765

70109

1673

2433

151

3079

25

749

219

493

GV

30648

12711

987

8604

1114

558

307

81017

281

006

083

005

123

PV34265

13895

1321

937

1213

2116

457

126957

987

006

098

011

283

PCV(

)2093(h)

2959(h)

2059(h)

1376

(m)

1008(m

)2036(h)

878

(l)236(h)

3227(h)

958(l)

1322(m

)1514

(m)

3414

(h)

GCV

()

1979(m

)2831

(h)

178(m

)1318

(m)

966(l)

144(m

)72

(l)1885(m

)1722

(m)

949(l)

1216

(m)

1021(m)

2253

(h)

ℎ2 119861(

)89

(h)

91(h)

75(h)

92(h)

92(h)

50(m

)67

(h)

64(h)

28(l)

98(h)

85(h)

45(m

)44

(m)

GA(

)3857

5577

3169

2602

1907

2097

1215

3102

1892

1939

2306

1418

3062

lowastSign

ificant

at005

prob


lysig

nificantat0

01p

robabilitylevel

SOVsou

rceo

fvariatio

ndfdegreeo

ffreedom

PHplant

heightFLR

flag

leafleng

thto

width

ratio

NT

numbero

ftillersp

erhill

DFdays

toflo

weringDMd

aystomaturity

NPHn

umbero

fpaniclesp

erhillPL

panicleleng

thN

PBPn

umbero

fprim

arybrachesp

erpanicle

TNGPtotalnu

mbero

fgrain

perp

anicleN

UFG

Pn

umber

ofun

filledgrainperp

anicleT

GWH

totalgrainweightp

erhill100G

Wone

hund

redgrainweightSL

seedleng

thSLW

Rseed

leng

thto

width

ratio

Yhayieldin

thaGV

geno

typicv

ariancePV

pheno

typic

varia

ncePC

Vph

enotypiccoeffi

cientofvariatio

nGCV

genotypiccoeffi

cientofvariatio

nℎ2 119861broad

senseh


ofmean)lowastsig

nificantat005level

andlowastlowasthigh

lysig

nificant

at001

level(hhighmm

oderatellow)



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology



cientamon

g16

quantitativetraits

for4

5ric

eaccessio

ns

FLR

NT

DF

DM

NP

PLTN

GGW

TGW

SLSLWR

YTHA


00958



006

656

FLR

013341lowast

006756

02601lowastlowast

006


NT


0114


DF


DM


011109

012724

NP


minus01037


005928


0117

2TN



GW


0118


TGW

015764lowastlowast

006



SLWR

006269

Notelowastsig

nificanta

t005prob

ability


lysig

nificanta

t001p

robabilitylevelPH

plant

heightF

LRfl

agleaf

leng

thto

width

ratio

NT

numbero

ftillersp

erhill

DF

days

toflo

wering

DMd

aysto

maturity

LAleafareaNPHn

umbero

fpaniclesp

erhillPL

panicleleng

thT

NGPtotalnu

mbero

fgrain

perp

anicle100

GWone

hund

redgrainweightTG

WH

totalgrainweightp

erhillSLseedleng

th

SLWR

seed

leng

thwidth

ratio

Yhayieldin

tha






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology






GN1

GN27

GN25GN13

GN6

GN20

GN26

GN2GN5

GN3

GN33

GN17

GN8

GN29

GN19

GN16

GN15

GN23GN22

GN24

GN31GN28

GN30

GN14

GN32GN7GN11

GN18GN35

GN34GN4

GN9

GN10GN21

GN12

GN36

GN38

GN37

GN40

GN41GN39

GN42

GN44

GN45GN43

Group I

Group II

Group III

Group IV

Group V361 277 193 143 110 026

Coefficient
















5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology













5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology




5 Conclusion




Acknowledgments


References




















































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology

































Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology








Volume 2014

Zoology






















Advances in

Virolog y



Volume 2014




Enzyme Research



Microbiology

Documents

Research Article Genetic Variability and Selection ...downloads.hindawi.com/journals/tswj/2014/190531.pdf · Research Article Genetic Variability and Selection Criteria in Rice Mutant