Research Article Line Tester Mating Design Analysis for ...downloads.hindawi.com/journals/ija/2013/201851.pdf · Line × Tester Mating Design Analysis for Grain Yield and Yield

Hindawi Publishing CorporationInternational Journal of AgronomyVolume 2013 Article ID 201851 9 pageshttpdxdoiorg1011552013201851

Research ArticleLine times Tester Mating Design Analysis for Grain Yield and YieldRelated Traits in Bread Wheat (Triticum aestivum L)

Zine El Abidine Fellahi1 Abderrahmane Hannachi1

Hamenna Bouzerzour2 and Ammar Boutekrabt3

1 National Institute of Agricultural Research Setif Agricultural Research Unit Setif 19000 Algeria2 Faculty of Life and Natural Sciences Ecology and Plant Biology Department University of Ferhat Abbas Setif 1 19000 Algeria3 Faculty of Agro-Veterinary and Biological Sciences Agronomy Department University of Saad Dahlab Blida 09000 Algeria

Correspondence should be addressed to Zine El Abidine Fellahi zinouagrogmailcom

Received 6 March 2013 Revised 20 May 2013 Accepted 29 May 2013

Academic Editor David Clay

Copyright copy 2013 Zine El Abidine Fellahi 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

Nine bread wheat (Triticum aestivum L) genotypes were crossed in a line times tester mating designThe 20 F1rsquos and their parents were

evaluated in a randomized complete block design with three replications at the Field Crop Institute-Agricultural ExperimentalStation of Setif (Algeria) during the 20112012 cropping seasonThe results indicated that sufficient genetic variability was observedfor all characters studied A

899times Rmada A

899timesWifak and A

1135timesWifak hybrids had greater grain yield mean than the parents

A901

line and the tester Wifak were good combiners for the number of grains per spike MD is a good combiner for 1000-kernelweight and number of fertile tillers HD

1220is a good general combiner to reduce plant height Rmada is a good general combiner

to shorten the duration of the vegetative growth period A901timesWifak is a best specific combiner to reduce plant height to increase

1000-kernel weight and number of grains per spike AAtimesMD is a best specific combiner to reduce duration of the vegetative periodplant height and to increase the number of kernels per spike A

899timesWifak showed the highest heterosis for grain yield accompanied

with positive heterosis for the number of fertile tillers and spike length and negative heterosis for 1000-kernel weight and thenumber of days to heading 1205902gca120590

2

sca (1205902

1198631205902

119860)

12 low ratios and low to intermediate estimates of ℎ2ns supported the involvement ofboth additive and nonadditive gene effects The preponderance of non-additive type of gene actions clearly indicated that selectionof superior plants should be postponed to later generation

1 Introduction

Bread wheat (Triticum aestivum L) is an important staplefood in Algeria This crop ranks third after durum wheat(Triticum durum Desf) and barley (Hordeum vulgare L)with a yearly cropped area of 08 million hectares represent-ing 242 of the 33 million hectares devoted to small graincereals Algeria imported 30 million tons of bread wheat in20102011 to remedy the decline in the domestic productionand to build stocks to meet the needs

Increasing wheat production can be achieved by appli-cation of improved agronomic technics developing andadopting high yielding varieties Major emphasis in breedingprogram is put on the development of improved varietieswith superior qualitative and quantitative traits and resilience

to abiotic stresses In fact genetic improvement in breadwheat having better tolerance against terminal heat andwater stress has a good promise to improve grain yieldaverage and total wheat production

However to breed high yielding varieties breeders oftenface the problem of selecting parents and crosses In thiscontext various breeding approaches have been suggestedThe line times tester analysis method introduced by Kempthorne[1] is one of the powerful tools available to estimate thecombining ability effects and aids in selecting desirable par-ents and crosses for exploitation in pedigree breeding [2ndash4]Performances per se do not necessarily reveal which parentsare good or poor combiners To surmount this difficulty it isnecessary to gather information on the nature of gene actionsGeneral combining ability is attributed to additive type of

2 International Journal of Agronomy

gene effects while specific combining ability is attributed tononadditive type of gene actions Nonadditive gene type ofactions is not reliably fixable whereas additive type of geneactions or complementary type epistatic gene interactions arereliably fixable [5ndash7]

Heterosis estimates for differentmorphological and yieldrelated characters are attributed to both additive and non-additive gene actions Heritability gives information aboutgenetic variation it is useful for predicting the response toselection in the succeeding generations Heritability is depen-dent upon the nature of gene action [8ndash10] Better under-standing of the underlying genetic control of importanttraits in bread wheat is useful in breeding for higher grainyield Kamaluddin et al [11] reported high contribution ofgeneral combining ability for genetic control of bread wheatcharacters Kumar and Maloo [12] identified the best specificand general combiners that were efficient for breeding daysto flowering and grain yield in bread wheat Involvement ofboth additive and dominance gene actions was also reportedfor genetic control of heading time in wheat [7] grain yield[13] number of grains per spike 1000-kernel weight [14] andfertile tillers per plant [15] Ahmad et al [16] reported thatadditive gene effect was important for days to heading Khanand Habib [17] observed that grain weight was controlled byover dominance type of gene action

The objectives of this study are to assess the combiningability to determine the nature andmagnitude of gene actionsand to estimate heterosis and heritability for yield and yield-related traits in a line times tester mating design in bread wheat

2 Materials and Methods

The present investigation was carried out at the Field CropInstitute-Agricultural Experimental Station of Setif (ITGC-AES 36∘121015840N and 05∘241015840E Algeria) during the 20102011 and20112012 cropping seasons The soil of the experimental siteis silty clay with CaCO

3and organic matter contents of 35

and 135 respectivelyThe experimental material comprisesnine bread wheat (Triticum aestivum L) genotypes Fivegenotypes Acsad

901 Acsad

899 Acsad

1135 Acsad

1069 and Ain

Abid were used as females hereafter designated as linesand four genotypes Mahon Demias Rmada HD

1220Wifak

designated as testers were used as males Mahon Demias isa genealogical selection from a land race introduced fromBalearic Islands in the mid-forties of the past century Thiscultivar is widely adapted to the arid and semiarid highplateaus of Algeria HD

1220is a selection from CIMMYT

segregatingmaterial it was released as cultivar in the ninetiesDrought tolerant and earlymaturing this variety gained largeacceptance from farmers due to its high yielding potential[18]

The nine parents were crossed to produce 20 F1hybrids

according to the line times tester mating design developed byKempthorne [1] F

1seeds were sown in the field along with

their parents in a randomized complete block design withthree replications Each plot comprised one row of 25mlength with space of 30 cm between rows and seeds wereplaced 15 cm apart Recommended cultural practices werefollowed to raise a good crop Monoammonium phosphate

Table 1 ANOVA for line times tester analysis

Source of variation Degree of freedom (df) Mean squareReplication (119903) (119903 minus 1)Genotypes (119892) (119892 minus 1) MS

2

Parents (119901) (119901 minus 1)Parents versus crosses 1

Crosses (119888) (119888 minus 1)Lines (119897) (119897 minus 1) 119872

119897

Testers (119905) (119905 minus 1) 119872119905

Lines times testers (119897 minus 1)(119905 minus 1) 119872119897times119905

Error (119903 minus 1)(119905 minus 1) MS1

Where MS2119872119897119872119905119872119897times119905 andMS1 were genotypic mean square line meansquare testermean square linetimes testermean square and errormean squarerespectively

(52P2O5+ 12N) with 80 kg haminus1 was applied just before

sowing and 75 kg haminus1 of Sulfate (26 N + 35 SO3) was

spread at tillering stageWeeds were controlled by applicationof 12 g haminus1 of Granstar [Methyl Triberunon] herbicide mixedwith water

Five competitive plants (excluding border plants) weretagged before heading and data were recorded for the numberof days to heading plant height spike length number offertile tillers per plant number of grains per spike 1000-kernel weight and grain yield per plant Data recordedwere subjected to analysis of variance according to Steeland Torrie [19] to determine significant differences amonggenotypes Combining ability effects are very effective geneticparameters in deciding the next phase of breeding programsThey were computed according to the line times tester method[20] Line times tester analysis was performed as outlined in theformat of ANOVA table given in Table 1

The variances for general and specific combining abilitywere tested against their respective error variances derivedfrom the analysis of variance of the different traits as follows

Covariance of half-sib of line

= CovHS (line)

=

119872119897minus119872119897times119905

119903119905

(1)

Covariance of half-sib of tester

= CovHS (tester)

=

119872119905minus119872119897times119905

119903119897

(2)

Covariance of full sib

= CovFS

=

(119872119897minus119872119890) + (119872

119905minus119872119890) + (119872

119897times119905minus119872119890)

3119903

+

6119903CovHS minus 119903 (119897 + 119905)CovHS3119903

(3)

International Journal of Agronomy 3

Table 2 Analysis of variance for combining ability effects of different bread wheat characters

Source df DHE PHT SL FT TKW NG GYRep 2 57 541 03 13 12 523 62Gen 28 156lowast 2268lowast 29lowast 234lowast 332lowastlowast 1528lowastlowast 397lowast

Par (P) 8 264lowast 3480lowast 26lowast 299lowast 631lowastlowast 2791lowastlowast 427lowast

Crosses (C) 19 107lowast 1718lowast 25lowast 196lowast 223lowastlowast 1076lowastlowast 338lowast

P versus C 1 214lowast 3014lowast 119lowast 445lowast 09ns 23ns 1290lowast

Lines (L) 4 212lowast 890lowast 33lowast 281lowast 268lowastlowast 1205lowast 655lowast

Testers (L) 3 395lowast 7512lowast 25lowast 372lowast 911lowastlowast 3577lowastlowast 258lowast

L versus T 1 76ns 1747lowast 03ns 151lowast 453lowast 2676lowast 22ns

L times T 12 42ns 468lowast 19lowast 100lowast 36ns 408lowast 256lowast

Error 56 25 241 07 34 6 111 65DHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillers TKW thousand-kernel weight (g) NG numberof grains per spike GY grain yield (g) ns lowast and lowastlowast non-significant and significant effect at 005 and 001 probability P parents C crosses L lines and Ttesters

While CovHS (average) was calculated by the formula

CovHS (average)

=

1

119903 (2119897119905 minus 119897 minus 119905)

[

(119897 minus 1) (119872119897) + (119905 minus 1) (119872

119905)

119897 + 119905 minus 2

minus119872119897times119905]

(4)

Assuming no epistasis variance due to GCA (1205902gca) andvariance due to SCA (1205902sca) were calculated as follows

1205902

gca = CovHS = (1 + 119865

4

) 1205902

119860

1205902

sca = (1 + 119865

2

)

2

1205902

119863

(5)

Additive and dominance genetic variances (1205902119860and 1205902

119863) were

calculated by taking inbreeding coefficient (119865) equal to onethat is 119865 = 1 because both lines and testers were inbred

Significance test for general combining ability and spe-cific combining ability effects were performed using 119905-testMid-parent heterosis (119867PM) is defined as the increased vigorof the 119865

1over the mean of the parents It was estimated from

mean values and its significance was performed using 119905-test[21] Narrow sense heritability was estimated after derivationof the variance components [20] (1205902gca120590

2

sca) and (1205902

1198631205902

119860)

12

ratios were used to rate the relative weight of additive versusnonadditive type of gene actions [22]

3 Results and Discussion

31 Genetic Variability among Parents and Hybrids Theanalysis of variance revealed significant genotype effect forall the characters under study This provides evidence ofthe presence of sufficient genetic variability among linestesters and hybrids and allows further assessment of generalcombining ability analysis (Table 2) Differences between theextreme mean values for the measured traits were 87 days366 cm 34 cm 104 tillers 153 g 337 grains and 161 g for

days to heading plant height spike length fertile tillers 1000-kernel weight grains per spike and grain yield respectivelyThese differences were 3 to 5 times higher than the LSD005values Parents and crosses showed significant effects for alltraits Mean square of the contrast ldquoparents versus crossesrdquowas significant for days to heading plant height spike lengthfertile tillers and grain yield and nonsignificant for 1000-kernel weight and number of grains per spike (Table 2)The differences between overall mean of parents and that ofhybrids indicated that hybrids were 11 days earlier 40 cmtaller and had more effective tillers and a grain yield advan-tage of 27 g Parents and hybrids showed similar averages for1000-kernel weight and number of grains per spike (Table 3)Line and tester effects were significant for all traits (Table 2)

Among lines the differences between the extreme meanvalues for the measured traits were 54 days 147 cm 25 cm72 tillers 89 g 156 grains and 112 g for days to headingplant height spike length fertile tillers 1000-kernel weightgrains per spike and grain yield respectively (Table 3) Thebest grain yielding line is A

1135which is the tallest and

had also the highest average for the number of fertile tillersand 1000-kernel weight (Table 3) A

1069was the earliest

with an average of 1323 days while Ain Abid (AA) hadthe longest and most fertile spike (Table 3) Among testersthe differences between the extreme mean values for thecharacters under study were 73 days 366 cm 22 cm 84tillers 119 g 287 grains and 69 g for days to heading plantheight spike length fertile tillers 1000-kernel weight grainsper spike and grain yield respectively (Table 3) The bestgrain yielding tester was Rmada (204 g) which exhibitedalso the longest spike (136 cm) Wifak was the earliest withan average of 1327 days while Mahon Demias (MD) wasthe tallest with 998 cm and presented the highest mean forthe number of fertile tillers and 1000-kernel weight HD

1220

expressed the best average of the number of grains perspike (Table 3) Compared to testers lines showed shorterplant height (minus51 cm) lighter 1000-kernel weight (minus26 g)and higher number of grains per spike (+63 grains) Theinteraction lines times testers were significant for plant heightspike length fertile tillers number of grains per spike and


Table 3 Means of the measured characters for 9 bread wheat parents and their 20 F1 hybrids

DHE PHT SL FT TKW NG GYLinesA901 1330efg 662jk 118hij 78h 239i 551ab 105i

A899 1373b 696hijk 115ij 137abcdef 327abcdefgh 449cdef 200abcdefgh

A1135 1327efg 809bcdef 135abcde 150abcd 328abcdefg 444def 217abcdefg

A1069 1323efg 731efghij 127efghij 118bcdefgh 324abcdefgh 499bcdef 188abcdefgh

AA 1377ab 716fghijk 140abcde 88efgh 261ghi 600a 137ghi

TestersMD 1400a 998a 114j 168ab 392a 263h 173bcdefghi

Rmada 1327efg 760efghi 136abcde 141abcd 295abcdefgh 476bcdef 204abcdefgh

HD1220 1373b 632k 122fghij 124bcdefgh 273abcdefgh 550ab 188abcdefgh

Wifak 1327efg 706ghijk 128defghi 84fgh 328hi 493bcdef 135hi

Lines times testersA901 timesMD 1343def 876bc 132bcdefg 147abcd 321cdef 426efg 202abcdefgh

A901 times Rmada 1327efg 752fghij 135abcdef 149abcd 270hij 518abcd 206abcdefgh

A901 timesHD1220 1347cde 698hijk 119ghij 109cdefgh 250j 506bcdef 146fghi

A901 timesWifak 1337defg 671ijk 119ghij 83gh 293fghij 600a 154efghi

A899 timesMD 1370bc 896b 131cdefgh 177a 331bcde 349gh 204abcdefgh

A899 times Rmada 1320fg 758efghi 141abcd 165ab 297defghi 493bcdef 242abc

A899 timesHD1220 1357bcd 706ghijk 118ghij 139abcde 294efghi 502bcdef 202abcdefgh

A899 timesWifak 1323efg 780cdefgh 140abcde 182a 295efghi 505bcdef 266a

A1135 timesMD 1380ab 822bcde 116ij 140abcde 352bc 347gh 173bcdefghi

A1135 times Rmada 1320fg 803bcdef 141abc 133abcdefg 307defgh 475bcdef 194abcdefgh

A1135 timesHD1220 1337defg 745efghij 140abcde 151abcd 320cdef 478bcdef 227abcde

A1135 timesWifak 1313g 800cdefg 145ab 161abc 314cdefg 480bcdef 243ab

A1069 timesMD 1343def 992a 133bcdef 151abcd 368ab 424fg 236abcd

A1069 times Rmada 1320fg 777defgh 137abcde 139abcde 299defghi 537abc 222abcdef

A1069 timesHD1220 1337defg 740efghij 134bcdef 121bcdefgh 290fghi 502bcdef 174bcdefghi

A1069 timesWifak 1320fg 784cdefgh 140abcde 137abcde 303defgh 481bcdef 197abcdefgh

AA timesMD 1357bcd 861bcd 138abcde 121bcdefgh 336bcd 515abcdef 200abcdefgh

AA times Rmada 1340def 775defgh 148a 105defgh 297defghi 522abcd 162cdefghi

AA timesHD1220 1370bc 717fghijk 139abcde 111cdefgh 278ghij 521abcd 157defghi

AA timesWifak 1340def 765efghi 136abcde 108cdefgh 301defghi 516abcde 166bcdefghi

Over all mean 1343 773 132 132 306 484 190LSD005 19 69 10 22 29 66 30DHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillers TKW thousand-kernel weight (g) NG numberof grains per spike and GY grain yield (g)Means within each column followed by the same letter are not significantly different from each other based on the 005 probability level of LSD

grain yield suggesting that hybrids perform better than theparents for these traits (Table 2)

Mean values of the hybrids were within the limits of themeans of the parents for the number of days to headingplant height spike length and number of grain per spikeFor grain yield A

899times Rmada A

899times Wifak and A

1135times

Wifak cross-combinations presented higher mean valuesthan the parents (Table 3) Genetic variability and meanperformance of parents and hybrids are important criteria forgenotypic evaluation however the parents with high meanvalue may not transmit this characteristic to their hybridsThese parental and hybrid abilities are estimated in termsof general combining ability (GCA) and specific combiningability (SCA) effects

32 General and Specific Combining Ability Effects The sig-nificance of mean squares due to lines and testers for thenumber of days to heading and 1000-kernel weight suggestedthe prevalence of additive genetic effects for these traitsWhile the simultaneous significance of mean squares dueto lines testers and lines times testers for plant height spikelength fertile tillers number of grains per spike and grainyield indicated that both additive and nonadditive type ofgene action were involved in the genetic control of thesecharacters A

901 A899

and AA lines and the tester HD1220

exhibited significant but negative GCA for grain yield A901

line and the tester Wifak presented significant and positiveGCA effects for the number of grains per spike (Table 4)MD is the best combiner for 1000-kernel weight number


Table 4 General combining ability (gi) effects for characters in bread wheat parents

Parents DHE PHT SL FT TKW NG GYLinesA901 minus025 minus553lowastlowast minus120lowastlowast minus217lowast minus334lowastlowast 417lowastlowast minus326lowast

A899 038 minus012 minus023 441lowastlowast minus023 minus341lowast minus447lowastlowast

A1135 minus038 1 02 147 264lowastlowast minus598lowastlowast 161A1069 minus150lowastlowast 564lowastlowast 028 009 134lowast 016 128AA 175lowastlowast minus099 095lowastlowast minus380lowastlowast minus04 505lowastlowast minus411lowastlowast

se (119892119897) 045 164 023 09 061 045 135

TestersMD 280lowastlowast 1552lowastlowast minus061lowastlowast 164lowast 538lowastlowast minus109lowastlowast 063Rmada minus220lowastlowast minus192 096lowastlowast 027 minus176lowastlowast 36lowastlowast 097HD1220 140lowastlowast minus974lowastlowast minus063lowastlowast minus158lowast minus292lowastlowast 255 minus258lowast

Wifak minus200lowastlowast minus386lowastlowast 028 minus033 minus071 47lowastlowast 098se (119892119905) 039 142 02 078 053 129 117

DHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillers TKW thousand-kernel weight (g) NG numberof grains per spike GY grain yield (g) se (119892119897) standard error for GCA effects for line se (119892119905) standard error for GCA effects for tester ns and lowast and lowastlowast non-significant and significant effect at 005 and 001 probability

of fertile tillers and the number of days to heading withGCA effect of 538 164 and 280 respectively HD

1220is

a good general combiner to reduce plant height (Table 4)Significant negative GCA effect for plant height is useful forthe development of dwarf plant material The tester Rmadahad significant and negative GCA for the number of days toheadingThis tester is a good general combiner to shorten theduration of the vegetative growth period

Even though SCA effects do not contribute tangibly inthe improvement of self-pollinated crops except in situationswhere exploitation of heterosis is feasible best hybrids areexpected to generate transgressive segregants which couldbe selected as potent homozygous lines A

901timesMD hybrid

exhibited significant SCA effects simultaneously for thenumber days to heading and spike length This cross-combination is best suited to select among its offspringrsquosthe earliest ones bearing long spike (Table 5) Accordingto Kenga et al [23] cross-combinations with high meansfavorable SCA estimates and involving at least one of the par-ents with high GCA would likely enhance the concentrationof favorable alleles to improve target traits In the presentstudy it is worth noting that A

901presented a significant and

negative GCA effect for spike length while MD presentedsignificant GCA negative for spike length and positive fordays to heading (Table 4) The positive SCA effect forspike length of this cross-combination resulted from parentshaving both significant and negative GCA One parent of thishybrid presented a positive and significant GCA for days toheading while the hybrid exhibited a significant and negativeSCA for this trait (Tables 4 and 5) A

901times Rmada presented a

significant and positive SCA effect for the number of fertiletillers per plant One parent of this cross-combination theline A

901 had a significant and negative GCA for the number

of fertile tillers A901times Wifak exhibited significant SCA

effects negative for plant height spike length fertile tillersand positive for 1000-kernel weight and number of grainsper spike (Table 5) Both parents A

901and Wifak of this

cross presented significant GCA for the number of grains perspike and plant height and at least one parent had significantGCA effect for days to heading (Wifak) spike length andthe number of fertile tillers (A

901) (Table 4) This hybrid is a

best specific combiner to reduce plant height and to increase1000-kernel weight and number of grains per spike Howeverthis cross-combination presents the disadvantage to reducethe number of fertile tillers which is a strong determinant ofgrain yield under semiarid growth conditions [24]

A1069times MD cross-combination has significant and pos-

itive SCA for plant height and 1000-kernel weight Thiscross is a best specific combiner to increase plant heightand 1000-kernel weight AA timesMD presented significant andnegative SCA effect for days to heading and plant height andsignificant and positive SCA effect for the number of kernelsper spike This cross is a best specific combiner to reduceduration of the vegetative period plant height and to increasethe number of kernels per spike (Table 5) None of the hybridsexhibited significant SCA effect for grain yield suggestingeven though the parents varied widely for grain yield theygenerated hybrids with grain yield averages within the grainyield limits of the parents

33 Gene Action Degree of Dominance Heterosis Heritabilityand Contribution to the Total Variance The variance due togeneral combining ability (1205902gca)was lower than specific com-bining ability variance (1205902sca) for all traits studied suggestingthe preponderance of nonadditive gene action controllingthese characters (Table 6) Dominance genetic variance waslarger than additive genetic variance for all traits Theseresults are supported by ratio of variance of general to specificcombining ability (1205902gca120590

2

sca) which was smaller than unity

and by the degree of dominance (12059021198631205902

119860)

12 which takesvalues greater than unity (Table 6)Therefore it appeared thatthe inheritance of all the studied characters was controlledby a preponderance of nonadditive gene effects Such type


Table 5 Specific combining ability (sij) effects for characters in bread wheat hybrids

Crosses DHE PHT SL FT TKW NG GYA901 timesMD minus205lowast 35 144lowastlowast 217 032 minus205 31A901 times Rmada 045 232 036 374lowast minus033 minus281 339A901 timesHD1220 minus015 207 minus049 minus041 minus214lowast minus351 minus2A901 timesWifak 175 minus789lowastlowast minus131lowastlowast minus550lowastlowast 214lowast 836lowastlowast minus45A899 timesMD 132 115 041 009 minus133 minus611lowast minus431A899 times Rmada minus117 minus214 031 minus034 074 094 1A899 timesHD1220 072 minus217 minus150lowastlowast minus25 132 35 minus135A899 timesWifak minus087 317 077 276 minus074 168 467A1135 timesMD 357lowastlowast minus1114lowastlowast minus230lowastlowast minus257 minus111 minus384 37A1135 times Rmada minus042 356 minus006 minus22 minus064 09 122A1135 timesHD1220 minus152 258 minus126 226 244 239 minus243A1135 timesWifak minus162 5 minus111 25 minus069 055 minus249A1069 timesMD minus08 979lowastlowast 023 052 256lowast 159 61A1069 times Rmada 07 minus497 minus082lowast minus002 minus065 404 137A1069 timesHD1220 minus04 minus281 029 minus087 minus08 minus016 minus383A1069 timesWifak 05 minus201 029 038 minus111 minus547lowast minus364AA timesMD minus205lowast minus330lowastlowast 022 minus02 minus045 1041lowastlowast 362AA times Rmada 045 123 02 minus118 088 minus308 minus23AA timesHD1220 135 034 044 152 minus082 minus221 049AA timesWifak 025 173 minus086lowast minus013 039 minus512 minus18Se (sij) 077 284 039 284 106 258 234DHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillers TKW thousand-kernel weight (g) NG numberof grains per spike GY grain yield (g) se (sij) standard error for SCA effects for crosses ns and lowast and lowastlowast non-significant and significant effect at 005 and001 probability

Table 6 Estimates of genetic components for the measured characters in bread wheat

DHE PHT SL FT TKW NG GY1205902

gca 193 2765 007 168 410 1469 1491205902

sca 1217 17348 084 1227 2380 9803 15281205902

119860387 5530 015 336 820 2938 297

1205902

1198631217 17348 084 1227 2380 9803 1528

1205902

gca1205902

sca 016 016 009 014 017 015 010[1205902

1198631205902

119860]12 177 177 239 191 170 183 227

ℎ2

ns 5630 6030 1030 3330 5440 5040 16901205902

119860 additive genetic variance 1205902

119863 dominance genetic variance ℎ2ns narrow sense heritability 1205902gca estimate of GCA variance 1205902sca estimate of SCA variance

and 1205902gca1205902

sca average degree of dominance DHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillersTKW thousand-kernel weight (g) NG number of grains per spike and GY grain yield (g)

of gene action clearly indicated that selection of superiorplants in terms of grain yield plant height fertile tillers andduration of the vegetative growth period should be postponedto later generation where these traits can be improvedby making selections among the recombinants within thesegregating populations

Selection efficiency is related to the magnitude of her-itability In this study low estimates of narrow-sense her-itability were observed for grain yield and spike lengthintermediate for the number of fertile tillers per plant andhigh for the number of days to heading plant height 1000-kernel weight and number of grains per spike (Table 6)

Heterosis is the process by which the performance of anF1is superior to that of the mean of the crossed parents Nine

hybrids among 20 exhibited a significant mid-parent hetero-sis for grain yield Besides heterosis for grain yield A

901times

MD and A899times Rmada expressed significant heterosis for

the number of fertile tillers spike length and the number ofdays to heading (Table 7) A

899timesWifak presented the highest

heterosis for grain yield accompanied with positive heterosisfor the number of fertile tillers and spike length and negativeheterosis for 1000-kernel weight and the number of days toheading (Table 7) Besides heterosis for grain yield AA timesMD hybrid exhibited significant and positive heterosis for


Table 7 Significant mid-parent heterosis () for seven traits in bread wheat genotypes

Crosses DHE PHT SL FT TKW NG GYA901 timesMD minus16 138 195 453A901 times Rmada 361 333A901 timesHD1220

A901 timesWifak 149 283A899 timesMD 144 161A899 times Rmada minus22 124 187 198A899 timesHD1220

A899 timesWifak minus20 113 152 647 minus99 588A1135 timesMD minus90A1135 times RmadaA1135 timesHD1220

A1135 timesWifak 103 376 381A1069 timesMD 147 104 113 307A1069 times Rmada 102A1069 timesHD1220

A1069 timesWifak 98 356 220AA timesMD minus23 87 194 290AA times RmadaAA timesHD1220 minus94AA timesWifakDHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillers TKW thousand-kernel weight (g) NG numberof grains per spike and GY grain yield (g)

Table 8 Proportional () contribution of lines testers and lines times testers to total hybrids variation in bread wheat

DHE PHT SL FT TKW NG GYLines (L) 3343 1123 3034 3267 2530 2358 4080Testers (T) 4671 7106 1724 3244 6450 5249 1205L times T 1987 1771 5241 3488 1020 2395 4784DHE number of days to heading PHT plant height (cm) SL spike length (cm) FT number of fertile tillers TKW thousand-kernel weight (g) NG numberof grains per spike and GY grain yield (g)

the number of grains per spike spike length and negativeheterosis for the number of days to heading (Table 7) A

1069times

MDhybrid exhibited significant and positive heterosis for thenumber of grains per spike spike length and plant height(Table 7)

Testers contributedmore to the total sum square for num-ber of days to heading plant height thousand kernel weightand number of grains per spike The contribution of lineswas lower compared to the testers and lines times testers inter-action for all traits under study All three sources of varia-tion contributed equally for the number of fertile tillers perplant Contribution of line times tester was slightly greater thanthat of testers and lines for grain yield and spike length(Table 8)These results showed that testers and the interactionlinestimes testers broughtmuch variation in the expression of thestudied traits The results of the present study revealed largevariation between parents and hybrids for the seven traitsunder study Compared to the parents hybrids were earliertaller bearing more effective tillers and had higher grainyielding A

1135line and Rmada tester were high grain yield-

ing Compared to testers lines were shorter and had low

1000-kernel weight and higher number of grains per spikeMean values of the hybrids were within the limits of the par-ental means for the number of days to heading plant heightspike length and number of grain per spike A

899times Rmada

A899timesWifak and A

1135timesWifak cross-combinations exhib-

ited higher mean values for grain yield than the parentsConcomitant significance of mean squares due to lines

testers and linestimes testers for plant height spike length fertiletillers number of grains per spike and grain yield suggestedthat both additive and nonadditive types of gene actions wereinvolved in the genetic control of the characters A

901line

and the tester Wifak were good combiners for the numberof grains per spike MD is a good combiner for 1000-kernelweight and number of fertile tillers HD

1220is a good general

combiner to reduce plant height while the tester Rmadais a good general combiner to shorten the duration of thevegetative growth period

A901times MD hybrid exhibited significant SCA effects

simultaneously for the number of days to heading andspike length This cross-combination is best suited and offersthe opportunity to select among the progenies early plant


with long spike This cross-combination resulted from Ltimes Lparents for spike length andHtimes L for days to heading Vermaand Srivastava [22] mentioned that positive SCA effect wasusually associated with crosses where at least one parentwas a good general combiner According to Singh et al [25]the desirable performance of combination like Htimes L may beascribed to the interaction between dominant alleles fromgood combiners and recessive alleles from poor combinersA901times Rmada presented a significant and positive SCA effect

for the number of fertile tillers per plant One parent ofthis cross-combination had significant and negative GCAfor the number of fertile tillers A

901times Wifak exhibited

significant SCA effects negative for plant height spike lengthfertile tillers and positive for 1000-kernel weight and numberof grains per spike Both parents of this cross presentedsignificant GCA for the number of grains per spike and plantheight and at least one parent had significant GCA effect fordays to heading spike length and the number of fertile tillersThis hybrid is a best specific combiner to reduce plant heightand to increase 1000-kernel weight and number of grainsper spike A

1069times MD cross-combination is a best specific

combiner to increase plant height and 1000-kernel weightAA times MD is a best specific combiner to reduce duration ofthe vegetative period plant height and to increase the numberof kernels per spike None of the hybrids exhibited significantSCA effect for grain yield suggesting even though the parentsvaried widely for grain yield they generated hybrids withgrain yield average within the grain yield limits of the parentsTiwari et al [26] mentioned that hybrid combinations whereat least one parent is a good general combiner could be usedto developing high yielding pure lines due to presence of addi-tive gene action even if these crosses showed nonsignificantSCA effects though In this study A

901 A899

AA and HD1220

were poor general combiners for grain yield 1205902gca was lowerthan 1205902sca and 120590

2

119863was larger than 1205902

119860for all traits suggesting

the preponderance of nonadditive gene action These resultsare supported by 1205902gca120590

2

sca ratio which was smaller than unity

and by (12059021198631205902

119860)

12 ratio which takes values greater than unityPremlatha et al [27] reported the importance of nonadditivegene action for plant height and grain yield Gnanasekaranet al [28] reported nonadditive gene action for seed weightand plant height while Sharma [29] reported an additive geneeffect Similar results of predominance of 1205902sca variance over1205902

gca have been reported by Verma et al [30] for barley Theresults of this study do not corroborated findings reported byBorghi et al [31] and Borghi and Perenzin [32] who observedthat 1205902gca was of greater importance than 1205902sca for majority ofcharacters Lucken [33] noted that nonadditive 1205902sca is bestexpressed in space planting The difference in the results ofvarious pieces of research may be attributed to differences ofbreeding material and to genotype times environments Betranet al [34] observed significant interactions for combiningabilities under low and high nitrogen in maize The prepon-derance of nonadditive type of gene actions clearly indicatedthat selection of superior plants in terms of grain yield plantheight fertile tillers and duration of the vegetative growthperiod should be postponed to later generation

Low estimates of h2ns were observed for grain yield andspike length intermediate for the number of fertile tillersper plant and high for the number of days to heading plantheight 1000-kernel weight and number of grains per spikeThis supported the involvement of both additive and nonad-ditive gene effects Medium to high narrow sense heritabilityestimates were reported by Yadav et al [35] for differenttraits Nine hybrids among 20 exhibited a significant mid-parent heterosis for grain yield Besides heterosis for grainyield A

901times MD and A

899times Rmada expressed significant

heterosis for the number of fertile tillers spike length and thenumber of days to heading A

899timesWifak showed the highest

heterosis for grain yield accompanied with positive heterosisfor the number of fertile tillers and spike length and negativeheterosis for 1000-kernel weight and the number of days toheading AA timesMD hybrid exhibited significant and positiveheterosis for the number of grains per spike spike lengthand negative heterosis for the number of days to headingThe results indicated that testers and the interaction lines timestesters contributed more to the variation in the expression ofthe studied traits

4 Conclusion

A899times Rmada A


1135times Wifak hybrids

had greater grain yield mean than the parents 1205902gca1205902

sca

(1205902

1198631205902

119860)

12 low ratios and low to intermediate estimates of h2nssupported the involvement of both additive and nonadditivegene effects with preponderance of nonadditive type ofgene actions The testers and the interaction lines times testerscontributed more to the variation of the expression of thedifferent traits A

901line and the tester Wifak were good

combiners for the number of grains per spike MD is agood combiner for 1000-kernel weight and number of fertiletillers HD

1220is a good general combiner to reduce plant

height Rmada is a good general combiner to shorten theduration of the vegetative growth period A

901times Wifak is

a best specific combiner to reduce plant height to increase1000-kernel weight and number of grains per spike AA timesMD is a best specific combiner to reduce duration of thevegetative period plant height and to increase the number ofkernels per spike A

899times Wifak showed the highest heterosis

for grain yield accompanied with positive heterosis forthe number of fertile tillers and spike length and negativeheterosis for 1000-kernel weight and the number of daysto heading The preponderance of nonadditive type of geneactions clearly indicated that selection of superior plantsshould be postponed to later generations

References

[1] O Kempthorne An Introduction to Genetic Statistics JohnWiley amp Sons New York NY USA 1957

[2] M Rashid A A Cheema andM Ashraf ldquoLine x tester analysisin basmati ricerdquo Pakistan Journal of Botany vol 39 no 6 pp2035ndash2042 2007

[3] S Basbag R Ekinci and O Gencer ldquoCombining ability andheterosis for earliness characters in line x tester population of


Gossypium hirsutum Lrdquo Hereditas vol 144 no 5 pp 185ndash1902007

[4] S K Jain and E V D Sastry ldquoHeterosis and combining abilityfor grain yield and its contributing traits in bread wheat(Triticum aestivum L)rdquo RRJAAS vol 1 pp 17ndash22 2012

[5] B Xiang and B Li ldquoA new mixed analytical method for geneticanalysis of diallel datardquoTheCanadian Journal of Forest Researchvol 31 no 12 pp 2252ndash2259 2001

[6] W Yan and L A Hunt ldquoBiplot analysis of diallel datardquo CropScience vol 42 no 1 pp 21ndash30 2002

[7] M Iqbal A Navabi D F Salmon et al ldquoGenetic analysis offlowering and maturity time in high latitude spring wheatgenetic analysis of earliness in springwheatrdquoEuphytica vol 154no 1-2 pp 207ndash218 2007

[8] P G Swati and B R Ramesh ldquoThe nature and divergence inrelation to yield traits in rice germplasmrdquoAnnals of AgriculturalResearch vol 25 pp 598ndash5602 2004

[9] Z Hasnain G Abbas A Saeed A Shakeel A Muhammadand M A Rahim ldquoCombining ability for plant height andyield related traits in wheat (Triticum aestivum L)rdquo Journal ofAgricultural Research vol 44 pp 167ndash1175 2006

[10] M A Chowdhary M Sajad and M I Ashraf ldquoAnalysis oncombining ability of metric traits in bread wheat (Triticumaestivum L)rdquo Journal of Agricultural Research vol 45 pp 11ndash118 2007

[11] K Kamaluddin R M Singh L C Prasad M Z Abdin and AK Joshi ldquoCombining ability analysis for grain filling durationand yield traits in spring wheat (Triticum aestivum L EmThell)rdquo Genetics and Molecular Biology vol 30 no 2 pp 411ndash416 2007

[12] V Kumar and S R Maloo ldquoHeterosis and combining abilitystudies for yield components and grain protein content in breadwheat (Triticum aestivum L)rdquo Indian Journal of Genetics andPlant Breeding vol 71 no 4 pp 363ndash366 2011

[13] N Mahmood and M A Chowdhry ldquoInheritance of flag leaf inbread wheat genotypesrdquoWheat Information Service vol 90 pp7ndash12 2000

[14] J Ahmadi A A Zali B Y Samadi A Talaie M R Ghannadhaand A Saeidi ldquoA study of combining ability and gene effect inbread wheat under stress conditions by diallel methodrdquo IranianJournal of Agricultural Sciences vol 34 pp 1ndash18 2003

[15] M A ChowdhryM TMahmood and I Khaliq ldquoGenetic anal-ysis of some drought and yield related characters in Pakistanispring wheat varietiesrdquo Wheat Information Service vol 82 pp11ndash118 1996

[16] F Ahmad S Khan A Latif H Khan A Khan and A NawazldquoGenetics of yield and related traits in breadwheat over differentplanting dates using diallel analysisrdquo African Journal of Agricul-tural Research vol 6 no 6 pp 1564ndash1571 2011

[17] A S Khan and I Habib ldquoGene action in a five parent diallelcross of spring wheat (Triticum aestivum L)rdquo Pakistan Journalof Biological Sciences vol 6 pp 1945ndash11948 2003

[18] L Benderradji F Brini S B Amar et al ldquoSodium transportin the seedlings of two bread wheat (Triticum aestivum L)Genotypes showing contrasting salt stress tolerancerdquoAustralianJournal of Crop Science vol 5 no 3 pp 233ndash241 2011

[19] R G D Steel and J H Torrie Principles and Procedures ofStatistics A Biometrical Approach McGrawHill New York NYUSA 1980

[20] R K Singh and B D Chaudhary Biometrical Methods inQuantitative Genetic Analysis Kalyani New Delhi India 1985

[21] G Oettler S H Tams H F Utz E Bauer and A E MelchingerldquoProspects for hybrid breeding in winter triticale I Heterosisand combining ability for agronomic traits in European elitegermplasmrdquo Crop Science vol 45 no 4 pp 1476ndash1482 2005

[22] O P Verma and H K Srivastava ldquoGenetic component andcombining ability analyses in relation to heterosis for yield andassociated traits using three diverse rice-growing ecosystemsrdquoField Crops Research vol 88 no 2-3 pp 91ndash102 2004

[23] R Kenga S O Alabi and S C Gupta ldquoCombining ability stud-ies in tropical sorghum (Sorghum bicolor (L) Moench)rdquo FieldCrops Research vol 88 no 2-3 pp 251ndash260 2004

[24] A Adjabi H Bouzerzour C Lelarge A BenmahammedA Mekhlouf and A Hanachi ldquoRelationships between grainyield performance temporal stability and carbon isotope dis-crimination in durum wheat (Triticum durum Desf) underMediterranean conditionsrdquo Journal of Agronomy vol 6 no 2pp 294ndash301 2007

[25] N B Singh V P Singh and N Singh ldquoVariation in physiolog-ical traits in promising wheat varieties under late sown con-ditionrdquo Indian Journal of Plant Physiology vol 19 pp 171ndash1752005

[26] D K Tiwari P Pandey S P Giri and J L Dwivedi ldquoPredictionof gene action heterosis and combining ability to identifysuperior rice hybridsrdquo International Journal of Botany vol 7 no2 pp 126ndash144 2011

[27] M Premlatha A Kalamani and A Nirmalakumari ldquoHeterosisand combining ability for grain yield and quality in maize (Zeamays L)rdquo Advances in Environmental Biology vol 5 no 6 pp1264ndash1266 2011

[28] M Gnanasekaran P Vivekanandan and SMuthuramu ldquoCom-bining ability and heterosis for yield and grain quality in twoline rice (Oryza sativa L) hybridsrdquo Indian Journal of HumanGenetics vol 66 pp 6ndash69 2006

[29] R K Sharma ldquoStudies on gene action and combining ability foryield and its component traits in rice (Oryza sativa L)rdquo IndianJournal of Human Genetics vol 66 pp 227ndash2228 2006

[30] A K Verma S R Vishwakarma and P K Singh ldquoLine x testeranalysis in barley (Hordeum vulgare L) across environmentsrdquoBarley Genetics Newsletter vol 37 pp 29ndash233 2007

[31] B Borghi M Perenzin and R J Nash ldquoCombining abilityestimates in bread wheat and performances of 100 F1 hybridsproduced using a chemical hybridizing agentrdquo Journal of Genet-ics and Breeding vol 43 pp 11ndash116 1989

[32] B Borghi andM Perenzin ldquoDiallel analysis to predict heterosisand combining ability for grain yield yield components andbread-making quality in bread wheat (Triticum aestivum L)rdquoTheoretical and Applied Genetics vol 89 no 7-8 pp 975ndash9811994

[33] K A Lucken ldquoThe breeding and production of hybrid wheatin USA genetic improvement in yield of wheatrdquo in CSSA SpecPub Crop Science Society of America and American Society ofAgronomy vol 13 pp 87ndash107 Madison Wis USA 1986

[34] F J Betran D BeckM Banziger andGO Edmeades ldquoGeneticanalysis of inbred and hybrid grain yield under stress andnonstress environments in tropical maizerdquoCrop Science vol 43no 3 pp 807ndash817 2003

[35] A K Yadav R K Maan S Kumar and P Kumar ldquoVariabilityheritability and genetic advance for quantitative characters inhexaploid wheat (Triticum aestivum L)rdquo Electronic Journal ofPlant Breeding vol 2 pp 405ndash4408 2011

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014


Food ScienceInternational Journal of

Agronomy


International Journal of



Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in


PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of


ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of


Plant GenomicsInternational Journal of


Biotechnology Research International


Forestry ResearchInternational Journal of


Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of


Veterinary Medicine International


Cell BiologyInternational Journal of


Evolutionary BiologyInternational Journal of



gene effects while specific combining ability is attributed tononadditive type of gene actions Nonadditive gene type ofactions is not reliably fixable whereas additive type of geneactions or complementary type epistatic gene interactions arereliably fixable [5ndash7]

Heterosis estimates for differentmorphological and yieldrelated characters are attributed to both additive and non-additive gene actions Heritability gives information aboutgenetic variation it is useful for predicting the response toselection in the succeeding generations Heritability is depen-dent upon the nature of gene action [8ndash10] Better under-standing of the underlying genetic control of importanttraits in bread wheat is useful in breeding for higher grainyield Kamaluddin et al [11] reported high contribution ofgeneral combining ability for genetic control of bread wheatcharacters Kumar and Maloo [12] identified the best specificand general combiners that were efficient for breeding daysto flowering and grain yield in bread wheat Involvement ofboth additive and dominance gene actions was also reportedfor genetic control of heading time in wheat [7] grain yield[13] number of grains per spike 1000-kernel weight [14] andfertile tillers per plant [15] Ahmad et al [16] reported thatadditive gene effect was important for days to heading Khanand Habib [17] observed that grain weight was controlled byover dominance type of gene action

The objectives of this study are to assess the combiningability to determine the nature andmagnitude of gene actionsand to estimate heterosis and heritability for yield and yield-related traits in a line times tester mating design in bread wheat

2 Materials and Methods

The present investigation was carried out at the Field CropInstitute-Agricultural Experimental Station of Setif (ITGC-AES 36∘121015840N and 05∘241015840E Algeria) during the 20102011 and20112012 cropping seasons The soil of the experimental siteis silty clay with CaCO

3and organic matter contents of 35

and 135 respectivelyThe experimental material comprisesnine bread wheat (Triticum aestivum L) genotypes Fivegenotypes Acsad

901 Acsad

899 Acsad

1135 Acsad

1069 and Ain

Abid were used as females hereafter designated as linesand four genotypes Mahon Demias Rmada HD

1220Wifak

designated as testers were used as males Mahon Demias isa genealogical selection from a land race introduced fromBalearic Islands in the mid-forties of the past century Thiscultivar is widely adapted to the arid and semiarid highplateaus of Algeria HD

1220is a selection from CIMMYT

segregatingmaterial it was released as cultivar in the ninetiesDrought tolerant and earlymaturing this variety gained largeacceptance from farmers due to its high yielding potential[18]

The nine parents were crossed to produce 20 F1hybrids

according to the line times tester mating design developed byKempthorne [1] F

1seeds were sown in the field along with

their parents in a randomized complete block design withthree replications Each plot comprised one row of 25mlength with space of 30 cm between rows and seeds wereplaced 15 cm apart Recommended cultural practices werefollowed to raise a good crop Monoammonium phosphate

Table 1 ANOVA for line times tester analysis

Source of variation Degree of freedom (df) Mean squareReplication (119903) (119903 minus 1)Genotypes (119892) (119892 minus 1) MS

2

Parents (119901) (119901 minus 1)Parents versus crosses 1

Crosses (119888) (119888 minus 1)Lines (119897) (119897 minus 1) 119872

119897

Testers (119905) (119905 minus 1) 119872119905

Lines times testers (119897 minus 1)(119905 minus 1) 119872119897times119905

Error (119903 minus 1)(119905 minus 1) MS1

Where MS2119872119897119872119905119872119897times119905 andMS1 were genotypic mean square line meansquare testermean square linetimes testermean square and errormean squarerespectively

(52P2O5+ 12N) with 80 kg haminus1 was applied just before

sowing and 75 kg haminus1 of Sulfate (26 N + 35 SO3) was

spread at tillering stageWeeds were controlled by applicationof 12 g haminus1 of Granstar [Methyl Triberunon] herbicide mixedwith water

Five competitive plants (excluding border plants) weretagged before heading and data were recorded for the numberof days to heading plant height spike length number offertile tillers per plant number of grains per spike 1000-kernel weight and grain yield per plant Data recordedwere subjected to analysis of variance according to Steeland Torrie [19] to determine significant differences amonggenotypes Combining ability effects are very effective geneticparameters in deciding the next phase of breeding programsThey were computed according to the line times tester method[20] Line times tester analysis was performed as outlined in theformat of ANOVA table given in Table 1

The variances for general and specific combining abilitywere tested against their respective error variances derivedfrom the analysis of variance of the different traits as follows

Covariance of half-sib of line

= CovHS (line)

=

119872119897minus119872119897times119905

119903119905

(1)

Covariance of half-sib of tester

= CovHS (tester)

=

119872119905minus119872119897times119905

119903119897

(2)

Covariance of full sib

= CovFS

=

(119872119897minus119872119890) + (119872

119905minus119872119890) + (119872

119897times119905minus119872119890)

3119903

+

6119903CovHS minus 119903 (119897 + 119905)CovHS3119903

(3)













CovHS (average)

=

1

119903 (2119897119905 minus 119897 minus 119905)

[

(119897 minus 1) (119872119897) + (119905 minus 1) (119872

119905)

119897 + 119905 minus 2

minus119872119897times119905]

(4)


1205902

gca = CovHS = (1 + 119865

4

) 1205902

119860

1205902

sca = (1 + 119865

2

)

2

1205902

119863

(5)


119863) were





2

sca) and (1205902

1198631205902

119860)

12










1220




































899times Rmada A


1135times



901 A899









se (119892119897) 045 164 023 09 061 045 135





1220is



901timesMD hybrid
















2



119860)







gca 193 2765 007 168 410 1469 1491205902

sca 1217 17348 084 1227 2380 9803 15281205902

119860387 5530 015 336 820 2938 297

1205902

1198631217 17348 084 1227 2380 9803 1528

1205902

gca1205902

sca 016 016 009 014 017 015 010[1205902

1198631205902

119860]12 177 177 239 191 170 183 227

ℎ2

ns 5630 6030 1030 3330 5440 5040 16901205902



and 1205902gca1205902






901times















1069times






899times Rmada





901line













901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014




































CovHS (average)

=

1

119903 (2119897119905 minus 119897 minus 119905)

[

(119897 minus 1) (119872119897) + (119905 minus 1) (119872

119905)

119897 + 119905 minus 2

minus119872119897times119905]

(4)


1205902

gca = CovHS = (1 + 119865

4

) 1205902

119860

1205902

sca = (1 + 119865

2

)

2

1205902

119863

(5)


119863) were





2

sca) and (1205902

1198631205902

119860)

12










1220




































899times Rmada A


1135times



901 A899









se (119892119897) 045 164 023 09 061 045 135





1220is



901timesMD hybrid
















2



119860)







gca 193 2765 007 168 410 1469 1491205902

sca 1217 17348 084 1227 2380 9803 15281205902

119860387 5530 015 336 820 2938 297

1205902

1198631217 17348 084 1227 2380 9803 1528

1205902

gca1205902

sca 016 016 009 014 017 015 010[1205902

1198631205902

119860]12 177 177 239 191 170 183 227

ℎ2

ns 5630 6030 1030 3330 5440 5040 16901205902



and 1205902gca1205902






901times















1069times






899times Rmada





901line













901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014


























































899times Rmada A


1135times



901 A899









se (119892119897) 045 164 023 09 061 045 135





1220is



901timesMD hybrid
















2



119860)







gca 193 2765 007 168 410 1469 1491205902

sca 1217 17348 084 1227 2380 9803 15281205902

119860387 5530 015 336 820 2938 297

1205902

1198631217 17348 084 1227 2380 9803 1528

1205902

gca1205902

sca 016 016 009 014 017 015 010[1205902

1198631205902

119860]12 177 177 239 191 170 183 227

ℎ2

ns 5630 6030 1030 3330 5440 5040 16901205902



and 1205902gca1205902






901times















1069times






899times Rmada





901line













901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014





























se (119892119897) 045 164 023 09 061 045 135





1220is



901timesMD hybrid
















2



119860)







gca 193 2765 007 168 410 1469 1491205902

sca 1217 17348 084 1227 2380 9803 15281205902

119860387 5530 015 336 820 2938 297

1205902

1198631217 17348 084 1227 2380 9803 1528

1205902

gca1205902

sca 016 016 009 014 017 015 010[1205902

1198631205902

119860]12 177 177 239 191 170 183 227

ℎ2

ns 5630 6030 1030 3330 5440 5040 16901205902



and 1205902gca1205902






901times















1069times






899times Rmada





901line













901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014





























gca 193 2765 007 168 410 1469 1491205902

sca 1217 17348 084 1227 2380 9803 15281205902

119860387 5530 015 336 820 2938 297

1205902

1198631217 17348 084 1227 2380 9803 1528

1205902

gca1205902

sca 016 016 009 014 017 015 010[1205902

1198631205902

119860]12 177 177 239 191 170 183 227

ℎ2

ns 5630 6030 1030 3330 5440 5040 16901205902



and 1205902gca1205902






901times















1069times






899times Rmada





901line













901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014


































1069times






899times Rmada





901line













901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014































901 A899

AA and HD1220


2




2


and by (12059021198631205902

119860)




901times MD and A





4 Conclusion

A899times Rmada A




sca

(1205902

1198631205902

119860)






901times Wifak is




References








































Journal of




Agronomy





Microbiology




Volume 2014




























































Journal of




Agronomy





Microbiology




Volume 2014


























Journal of




Agronomy





Microbiology




Volume 2014
























Documents

Research Article Line Tester Mating Design Analysis for ...downloads.hindawi.com/journals/ija/2013/201851.pdf · Line × Tester Mating Design Analysis for Grain Yield and Yield