+ All Categories
Home > Documents > Analysis of Direct and Indirect Selection and Indices...

Analysis of Direct and Indirect Selection and Indices...

Date post: 19-Aug-2020
Category:
Upload: others
View: 1 times
Download: 0 times
Share this document with a friend
12
Research Article Analysis of Direct and Indirect Selection and Indices in Bread Wheat (Triticum aestivum L.) Segregating Progeny Zine El Abidine Fellahi , 1 Abderrahmane Hannachi, 2 and Hamenna Bouzerzour 3 1 Department of Agronomy, Faculty of Natural, Life and Earth Sciences and the Universe, University of Mohamed El Bachir El Ibrahimi, 34034 Bordj Bou Arr´ eridj, Algeria 2 National Agronomic Research Institute of Algeria (INRAA), Unit of S´ etif, 19000 S´ etif, Algeria 3 Department of Ecology and Plant Biology, Valorization of Natural Biological Resources Laboratory, Faculty of Natural and Life Sciences, University of Ferhat Abbas S´ etif-1, 19000 S´ etif, Algeria Correspondence should be addressed to Zine El Abidine Fellahi; [email protected] Received 27 December 2017; Accepted 5 March 2018; Published 19 April 2018 Academic Editor: Iskender Tiryaki Copyright © 2018 Zine El Abidine Fellahi et al. is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. ree selection methods including direct and indirect selection along with selection index based on the phenotypic values of eleven traits of agronomic interest were assessed for their application in F 4 bread wheat progenies. Significant genetic variation existed among parents and crosses for the traits measured. e following were the most efficient indices for simultaneous selection of superior lines for yield and its components: base index of Williams, followed by the sum of ranks index of Smith and Hazel. e selection-based index provided the highest grain yield gains as compared to the other selection criteria, except for flag leaf area, indicating that the direct and indirect monotrait selection were not appropriate in the situation analyzed in this work. PCA identified Ain Abid × Mahon-Demias, Ain Abid × Rmada, and Ain Abid × El-Wifak as the most promising populations. At 5% selection intensity, the top 30 lines selected were distinguished, in comparison with the standard check Hidhab, by significant improvements in yield and yield components. 1. Introduction In Algeria, most of wheat producing areas are located in the High Plateaus which are characterized by cold winters, insufficient and erratic rainfall, frequent spring frosts, and late-season sirocco occurrence [1, 2]. In addition to these cli- matic stresses, there are some other technical constraints that essentially arise from the use of unproductive varieties and oſten bad agronomic practices. Selection for a better adapta- tion to environmental stresses is, therefore, more promising outcome in the field of wheat breeding. Breeders are contin- ually seeking to improve the selection methods in order to develop superior wheat varieties with high grain yield, good end-use quality, and tolerance to biotic and abiotic stresses. Direct selection based on grain yield is mainly practiced in wheat breeding programs without considering the adaptive traits that are crucial production regulators under variable environments [3–5]. In these environments, the presence of genotype × environment interactions reduces the efficiency of using grain yield as the sole selection criterion and, thus, complicates the efforts of selection [6, 7]. In addition to the environmental effects, other factors such as polygenic nature, low heritability of grain yield, linkage, and nonadditive gene action may make the selection less efficient mainly in early segregating generations. In order to overcome these difficulties, breeders are focus- ing on other traits that can be used in parallel or indepen- dently of yield in a multitraits approach. Indirect selection uses some yield components that are more heritable than yield itself and more stable in relation to genetic and environ- mental factors affecting them. When these components are measured without error and expressed in appropriate units, their product is yield. is has created new opportunities for plant breeders to use certain morphological, physiological, and biochemical traits during selection for grain yield. In the literature, several authors have reported the use of many of Hindawi International Journal of Agronomy Volume 2018, Article ID 8312857, 11 pages https://doi.org/10.1155/2018/8312857
Transcript
Page 1: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

Research ArticleAnalysis of Direct and Indirect Selection and Indices inBread Wheat (Triticum aestivum L) Segregating Progeny

Zine El Abidine Fellahi 1 Abderrahmane Hannachi2 and Hamenna Bouzerzour3

1Department of Agronomy Faculty of Natural Life and Earth Sciences and the UniverseUniversity of Mohamed El Bachir El Ibrahimi 34034 Bordj Bou Arreridj Algeria2National Agronomic Research Institute of Algeria (INRAA) Unit of Setif 19000 Setif Algeria3Department of Ecology and Plant Biology Valorization of Natural Biological Resources LaboratoryFaculty of Natural and Life Sciences University of Ferhat Abbas Setif-1 19000 Setif Algeria

Correspondence should be addressed to Zine El Abidine Fellahi zinouagrogmailcom

Received 27 December 2017 Accepted 5 March 2018 Published 19 April 2018

Academic Editor Iskender Tiryaki

Copyright copy 2018 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

Three selectionmethods including direct and indirect selection along with selection index based on the phenotypic values of eleventraits of agronomic interest were assessed for their application in F4 bread wheat progenies Significant genetic variation existedamong parents and crosses for the traits measured The following were the most efficient indices for simultaneous selection ofsuperior lines for yield and its components base index of Williams followed by the sum of ranks index of Smith and Hazel Theselection-based index provided the highest grain yield gains as compared to the other selection criteria except for flag leaf areaindicating that the direct and indirectmonotrait selectionwere not appropriate in the situation analyzed in thiswork PCA identifiedAin Abid times Mahon-Demias Ain Abid times Rmada and Ain Abid times El-Wifak as the most promising populations At 5 selectionintensity the top 30 lines selected were distinguished in comparison with the standard check Hidhab by significant improvementsin yield and yield components

1 Introduction

In Algeria most of wheat producing areas are located inthe High Plateaus which are characterized by cold wintersinsufficient and erratic rainfall frequent spring frosts andlate-season sirocco occurrence [1 2] In addition to these cli-matic stresses there are some other technical constraints thatessentially arise from the use of unproductive varieties andoften bad agronomic practices Selection for a better adapta-tion to environmental stresses is therefore more promisingoutcome in the field of wheat breeding Breeders are contin-ually seeking to improve the selection methods in order todevelop superior wheat varieties with high grain yield goodend-use quality and tolerance to biotic and abiotic stresses

Direct selection based on grain yield is mainly practicedinwheat breeding programswithout considering the adaptivetraits that are crucial production regulators under variableenvironments [3ndash5] In these environments the presence of

genotype times environment interactions reduces the efficiencyof using grain yield as the sole selection criterion and thuscomplicates the efforts of selection [6 7] In addition to theenvironmental effects other factors such as polygenic naturelow heritability of grain yield linkage and nonadditive geneaction may make the selection less efficient mainly in earlysegregating generations

In order to overcome these difficulties breeders are focus-ing on other traits that can be used in parallel or indepen-dently of yield in a multitraits approach Indirect selectionuses some yield components that are more heritable thanyield itself andmore stable in relation to genetic and environ-mental factors affecting them When these components aremeasured without error and expressed in appropriate unitstheir product is yield This has created new opportunities forplant breeders to use certain morphological physiologicaland biochemical traits during selection for grain yield In theliterature several authors have reported the use of many of

HindawiInternational Journal of AgronomyVolume 2018 Article ID 8312857 11 pageshttpsdoiorg10115520188312857

2 International Journal of Agronomy

these traits to improve grain yield in diverse environments[8ndash10]

Selection-based index is another approach certainlycomplex but can avoid the limits of the single-trait selec-tion particularly the undesirable between-trait relationsthat present an additional nuisance in breedersrsquo work [11]Selection-based index approach targets the simultaneousimprovement of several traits at the same time includingthe grain yield [12 13] The indices allow the use of a singlevalue in the selection process since the analysis is carriedout by means of linear combinations of phenotypic dataof different traits of agronomic interest with the geneticproperties of a population [14] The objective is to guaranteethe improvement of the population genotypic values and con-sequently the efficiency of the selection process maximizingthe expected genetic gain In this purpose many selectionindices have been used as an effective selection criterion inplant breeding programs on different crops [15] Howeverthe conditions determining the usefulness of an appropriateselection index may vary with individual plant breeder Theobjective of this research paper was to evaluate the efficiencyand applicability of different selection criteria based on theestimation of genetic gain in F4 segregating populations ofbread wheat evaluated under semiarid conditions

2 Materials and Methods

21 Plant Material and Experimental Design This experi-ment had 609 genotypes comprising 20 F4-derived familiesand their 9 parents The history of these families is thatafter the initial crosses in 201011 [18] 30 F2 lines wereselected in each family by using pedigree method and wereevaluated during the 201213 cropping season [19] A totalof 600 lines were planted and harvested in bulk during twoconsecutive growing seasons 201314 and 201415 The F4lines along with their parents were planted in Setif ResearchUnit (36∘151015840N 5∘871015840E 1081 masl) of the Algerian NationalInstitute of Agronomic Research (INRAA) in a modifiedrandomized complete block design with three replicationsThe experimental unit consisted of a single row plot of 2mlength with rows spaced 02m apartThe plots were fertilizedwith 100 kg haminus1 of 46 superphosphate before sowing and75 kg haminus1 of 35 urea at tillering stage Weed control wasperformed chemically using 12 g haminus1 of the Granstar [tribe-nuron methyl] herbicide According to Chennafi et al [20]the climate of the region is a semiarid type continental withcold winter and hot and dry summer Total annual rainfallis around 350mm The soil used was classified as a silty claywith high CaCO3 content [20]

22Measurements The following traits weremeasured as perplot basis Chlorophyll content (CHL Spad) was determinedat heading stage using the SPAD-502 chlorophyll meter(Minolta Camera Co Osaka Japan) Flag leaf area (FLAcm2) was obtained using the formula described by Spagno-letti Zeuli and Qualset [21] FLA (cm2) = 119871 (cm) times 119897 (cm)where 119871 is the flag leaf length and 119897 refers to the flag leafwidth Heading date (HD days) was recorded as the number

Table 1 The skeleton of the analysis of variance

Source of variation df MS 119865-testBlock 119887 minus 1 M1 M1M6Genotype 119892 minus 1 M2 M2M6

Parents 119875 minus 1 M3 M3M6F4 119899 minus 1 M4 M4M6Parents vs F4 1 M5 M5M6

Residual (119892 minus 1)(119887 minus 1) M6 - -Total 119887119892 minus 1 - - - -

of calendar days from January first to the date when 50of the spikes were half way out from flag leaf Plant height(PH cm) was measured at maturity from the soil surface tothe top of the spike awns excluded Above ground biomass(BIO gmminus2) was estimated from a harvested area of 05mlong times 020m interrow spacing which served also to obtainthe grain yield (GY gmminus2) number of spikes (SN mminus2)and spikes weight (SW gmminus2) 1000-kernel weight (TKW g)was obtained from the count and weight of 250-kernel Thenumber of grains per spike (GN) was derived from estimatedvalues of grain yield number of spikes and 1000-kernelweight Harvest index (HI ) was estimated by the ratio be-tween grain yield and above ground biomass

23 Data Analysis Data collected were subjected to analysisof variance following the procedures described by McIntosh[22] the skeleton of the analysis of variance is shown inTable 1 The statistical model used considered the completerandomized block design as 119884119894119895 = 120583 + 119892119894 + 119887119894 + 120576119894119895 where 119884119894119895is the observation of the 119894th genotype evaluated in the 119895threplicate 120583 is the overall mean of the experiment 119892119894 is theeffect of the 119894th genotype 119887119895 is the effect of the 119895th block and120576119894119895 is the effect of the 119894119895th plot

In case 119865-test was significant standard error and criticaldifferences were calculated by using the least significantdifference test at 005 probability level (LSD005) accordingto Steel and Torrie [23] LSD005 = 119905005radic21205902119890 119887 where 119905005is the tabulated value of the 119905-test at 005 probability level for(119892 minus 1)(119887 minus 1) residual degrees of freedom 1205902119890 is the residualvariance and 119887 refers to the number of replications or blocks

The genotypic variance (1205902119892) and residual variance (1205902119890)were calculated and served to determine the following geneticand nongenetic parameters for each trait The coefficient ofexperimental variation (CV) was calculated by CV () =radic1205902119890 119883 where1205902119890 is the residual variance and119883 is the generalmean of the trait The coefficient of genetic variation (CV119892)was calculated by the following equation CV119892 () = radic1205902119892119883where1205902119892 is the genotypic variance and119883 is the generalmeanof the trait The variation index was determined as the ratioof CV119892CV119890 Broad-sense heritability of average progenies(ℎ2bs) was estimated by the expression [24] ℎ2bs () = 100 times[1205902119892(1205902119892+1205902119890 )]Narrow-sense heritability of individualswithinfamilies (ℎ2ns) was determined based on the parent-offspring

International Journal of Agronomy 3

regression To do so two methods were used the first wasby the linear regression of F4 on the parental F3 individualvalues [16] while the second approach was performed withstandardized data of offspring (F4) versus standardized of thecorresponding parent (F3) according to Frey andHorner [17]

The selection gains were estimated among families basedon three selection criteria direct selection indirect selectionand selection-based index considering the selection intensityof 5 of top families The expected gains by direct selectionfor each trait evaluated were estimated by the expression [25]Δ119866119894 = ℎ2119894times119878119894 = ℎ2119894times(119883119904119894minus1198830119894) whereΔ119866119894 is the gainwith thedirect selection carried for the 119894th trait ℎ2119894 is the heritability ofthe 119894th trait 119878119894 refers to the differential selection of the 119894th trait119883119904119894 is themean of the 119894th trait for the selected individuals and1198830119894 is the mean of the 119894th trait in the base population Theexpected gain of direct selection expressed as a percentage ofthe population mean is given by Δ119866119894119894 = (Δ119866119894 times 100)1198830119894

Gains from indirect response to selection were calculatedusing the following expression [25] GS119895(119894) = ℎ2119895 times (119883119894119895 minus1198830119895) = ℎ2119895 times DS119895(119894) where GS119895(119894) is the gain on the 119895th traitwith selection based on the 119894th trait 119883119904119895 is the mean of the119895th trait for the selected individuals based on the 119894th trait1198830119895 is the mean of the 119895th trait ℎ2119895 is the heritability ofthe 119895th trait and DS119895(119894) refers to the differential selection ofthe 119895th trait in which the selected lines presented the bestperformance for the 119894th trait The expected gain of indirectselection expressed as a percentage of the population meanis given by GS119895(119894) = (GS119895(119894) times 100)1198830119894

For the selection-based index the following methodolo-gieswere used for gains estimation the classic index proposedby Smith [26] and Hazel [27] the base index of Williams[28] the free weights and parameters index of Elston [29]the index of desired gains of Pesek and Baker [30] the multi-plicative index of Subandi et al [31] the sum of ranks index ofMulamba andMock [32] and the genotype-ideotype distanceindex proposed by Cruz [25] Each index displays certainparticularities in its calculations and as such applicationis generally laborious due to the need to assign adequateeconomic weights to each trait Based on the differentanalytical procedures of selection the best populations wereidentified and the gains from selection were calculated Allstatistical analyses were carried out using Genes software [33]and a Microsoft Excelcopy spreadsheet

3 Results and Discussion

31 Genetic Variability and Heritability The analysis of vari-ance revealed significant genotype effect for almost all thetraits under study except for flag leaf area and grain yieldwhich were not significant at 005 probability level (Table 2)This provides evidence of the presence of sufficient geneticvariability among parents and hybrids that can be exploitedin wheat breeding program through selection Partitioningthe genotype effect indicated significant differences betweenall parents for HD PH SN TKW SW GN BIO and HIand significant interpopulation differences for HD PH andGN The contrast ldquoParents versus F4rdquo was highly significant

for HD PH and SN while the F4 populations exhibitedsignificant differences for nearly all the observed quantitativephenotypic traits except for CHL FLA GN and HI whichwere not significant at the 5 probability level The resultsof this study corroborate those of Abd El-Shafi [34] whoreported significant and highly significant differences amonggenotypes (families + parents) and families for all studiedtraits across three segregating generations F2 F3 and F4Thisauthor also reported that greater response to selection canbe expected from selection in cross having greater phenotypicand genotypic variances

The coefficient of variation (CV) presented values be-tween 07 and 268 for heading date and grain yield respec-tively (Table 2) The CV above 20 is considered high indi-cating high dispersion of the experimental data which mayhave been caused by the genetic and phenotypic differencesbetween the studiedmaterials HighCV estimate obtained forgrain yield can be explained by the fact that it is quantitativetrait governed by several genes and highly influenced by theenvironment

The variances values coefficient of variation and geneticparameters estimates for wheat traits studied are presentedin Table 3 Broad-sense heritability is the proportion of totalphenotypic variation due to all genetic effects The knowl-edge of the genotypic determination coefficient (ℎ2bs) allowsestablishing an estimate of the genetic gain to be obtained anddefines the best strategy to be used in the plant breeding pro-gram [35] In this study the estimated broad-sense heritabil-ity varied from 000 to 9180The highest values were foundfor heading date (9180) followed by plant height (8082)1000-kernel weight (7290) biomass (6533) and numberof grains per spike (6164) indicating that these traits arehighly heritable among the genotypes evaluatedThese resultscan be confirmed with the values obtained by the CV119892CV119890ratio that were close to or greater than 1 for these traits sug-gesting satisfactory conditions for selection [36] Moderateestimates of ℎ2bs occurred for the number of chlorophyllcontents (4031) number of spikes (5200) spikes weight(5485) and harvest index (4067) On the other handthe lowest values of ℎ2bs were found for the flag leaf area(000) and grain yield (1939) These traits exhibited alsolow CV119892CV119890 ratio values indicating the dominant effect ofthe environment on crop

Generally literature indicates widely varying narrow-sense heritability estimates Mesele et al [37] reported highheritability values for days to heading days to maturityand 1000-kernel weight moderate estimates for grain fillingperiod spike length number of spikelets per plant grainsper spike and harvest index and low values for number oftillers per plant biomass yield and grain yield Evaluatingseven F2 populations derived through cross combinations offive parental varietieslines of bread wheat Saleem et al [38]found low to high broad-sense heritability values rangingfrom 475 to 926 depending on the trait and the crossThefindings of Yaqoob [39] showed that heritability estimateswere low for number of tillers per plant (20) grainsper spike (2681) days to maturity (3013) spike length(3666) and 1000-kernel weight (3868) moderate forplant height (4579) and high for heading date (8473)

4 International Journal of Agronomy

Table 2 Analysis of variance of different bread wheat traits studied

SV Bloc Genotypes Parents F4 Parents versus F4 Error CV ()df 2 28 8 19 1 56CHL 86 190lowastlowast 95ns 238lowastlowast 36ns 86 66HD 15 90lowastlowast 183lowastlowast 25lowastlowast 587lowastlowast 07 07FLA 894 111

ns69ns 66ns 01ns 149 204

PH 91 863lowastlowast 1030lowastlowast 800lowastlowast 726lowast 166 63SN 11831 111875lowastlowast 260924lowastlowast 26578ns 539945lowastlowast 53711 159TKW 335 282lowastlowast 394lowastlowast 127ns 2321lowastlowast 77 74SW 1170346 448736lowastlowast 744904lowastlowast 182488ns 3137463lowastlowast 202598 209GN 366 447lowastlowast 613lowastlowast 370lowast 410ns 171 168BIO 811895 1310774lowastlowast 2431421lowastlowast 436017ns 8965592lowastlowast 454408 185GY 341492 157279ns 268354ns 110965ns 562059lowast 126778 268HI 1222 623lowast 1366lowastlowast 361ns 214ns 37 168CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) HI harvest index () and nslowast and lowastlowast nonsignificant and significant effect at 005 and 001 probability

Table 3 Genetic and nongenetic parameters of different bread wheat traits studied

Traits 1205902119901 1205902119890 1205902119892 ℎ2bs ℎ2ns (SK) ℎ2ns (FH) CV119892 () CV119892CV119890Chl 336 200 135 4031 2855 3013 264 047HD 301 025 276 9180 1669 3432 131 193FLA 368 498 000 000 000 000 000 000PH 2878 552 2326 8082 2727 2727 743 119SN 372917 179037 193879 5200 000 000 959 060TKW 941 255 686 7290 3164 2599 698 095SW 1495791 675325 820467 5485 866 1070 1319 064GN 1467 563 904 6164 1670 714 1228 073BIO 4369245 1514693 2854552 6533 337 593 1448 079GY 524264 422592 101671 1939 1013 1231 758 028HI 2076 1231 845 4067 1410 1497 803 048CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY Grain yield (gmminus2) HI harvest index () ℎ2bsbroad-sense heritability ℎ2ns (SK) narrow-sense heritability according to Smith and Kinman [16] and ℎ2ns (FH) narrow-sense heritability according to Freyand Horner [17]

and grain yield (9983) His results also indicated that mostof these traits exhibited low heritability under drought stressconditions suggesting the presence of high genotype times envi-ronment interactions that affected the crop behavior

Narrow-sense heritability is the proportion of the totalphenotypic variation that is due to the additive effects ofgenes This component of variation is important because it isthe only variation that natural selection can act on Henceℎ2ns determines the resemblance of offspring to their parentsand the populationrsquos evolutionary response to selectionTherecan be considerable nonadditive genetic variance but thisdoes not contribute to the resemblance between parents andoffspring or the response to selection Low to moderatenarrow-sense heritability values were recorded in this studyChlorophyll content heading date plant height 1000-kernelweight and number of grains per spike recorded the highestestimates These traits were less influenced by the environ-mental factors and would respond positively to a selection

pressure in the current breeding programThe lowheritabilityvalues can be explained by the change occurring in thesegregating lines behavior from the precedent to the currentgeneration This change may be due to nonadditive geneaction andor high environmental factors effects

Means of the variables measured showed that the bestvalues varied depending on the cross and the trait and thefew populations had the best performances for several traitsat the same time (Table 4)The best grain yielding population(5575 g) was Ain Abid times El-Wifak which had also the highestaverage for the number of grains per spike (3147 grains)spikes weight (8573 g) above ground biomass (13503 g) andharvest index (413) Ain Abid timesHidhab cross combinationhad the longest vegetative cycle with an average of 1278days and presented the highest mean for the flag leaf area(210 cm2) Acsad1069 times El-Wifak had the highest average forthe chlorophyll content (540 Spad) Acsad1069 times Mahon-Demias was the tallest (7523 cm) while Acsad1135 times Rmada

International Journal of Agronomy 5

Table4Means

ofthem

easuredtraitsfor2

0F 4

breadwheatpo

pulatio

ns

Popu

latio

nCH

LHD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Acsad 901timesM

ahon

-Dem

ias

430

1260

201

737

4483

386

6340

220

11350

3792

334

Acsad 901timesR

mada

445

1252

199

608

4424

346

6336

281

10497

4255

405

Acsad 901timesH

idhab

425

1259

198

657

4267

334

6167

230

10983

3255

296

Acsad 901timesE

l-Wifa

k451

1249

204

704

4673

332

6843

304

11490

4604

401

Acsad 899timesM

ahon

-Dem

ias

437

1257

169

708

4760

364

6260

205

11517

3668

319

Acsad 899timesR

mada

434

1250

166

595

4797

378

6257

200

10327

3688

357

Acsad 899timesH

idhab

444

1269

182

566

4060

350

5980

228

9970

3360

337

Acsad 899timesE

l-Wifa

k454

1250

181

631

4147

355

5667

242

9657

3577

370

Acsad 1135timesM

ahon

-Dem

ias

416

1261

170

681

4567

398

6040

210

10737

3834

357

Acsad 1135timesR

mada

422

1252

168

615

5037

369

6853

240

11137

4404

396

Acsad 1135timesH

idhab

415

1270

175

688

4770

395

7447

232

12687

4509

355

Acsad 1135timesE

l-Wifa

k455

1252

170

646

3970

346

4893

221

8790

3099

353

Acsad 1069timesM

ahon

-Dem

ias

438

1258

182

752

4390

392

6233

217

11937

3713

311

Acsad 1069timesR

mada

442

1250

173

617

4147

348

5810

245

9617

3608

375

Acsad 1069timesH

idhab

467

1259

185

605

4113

350

6170

266

9767

3953

405

Acsad 1069timesE

l-Wifa

k540

1250

184

665

4233

346

6110

279

9940

4033

406

Ain

AbidtimesM

ahon

-Dem

ias

420

1277

200

727

4547

388

7473

272

12867

4818

375

Ain

AbidtimesR

mada

403

1263

205

658

4357

365

6993

282

11717

4581

391

Ain

AbidtimesH

idhab

432

1278

210

602

4050

368

6933

304

11307

4628

409

Ain

AbidtimesE

l-Wifa

k436

1269

203

651

4693

377

8573

315

13503

5575

413

ParentsR

ange

391

1250

87

530

2700

300

3200

975300

1850

141

522

1320

391

820

8000

490

12600

394

23300

8090

636

F 4Ra

nge

334

1240

76420

800

190

800

931500

430

126

538

1360

364

1000

8500

510

14800

568

24200

9210

700

LSD005

48

1463

67

1197

45

2352

68

3532

1851

99CH

Lchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)andHIharvestind

ex(

)

6 International Journal of Agronomy

exhibited the highest average for the number of spikes (5037spikes) For 1000-kernel weight Acsad1135 timesMahon-Demiaswas the best population with an average of 398 g

As the contrast ldquoParents versus F4rdquo was significant forPH SN TKW SW BIO and GY and compared with theleast significant difference (LSD005) significant differencesexisted between the parents and hybrids values for thesemea-sured traits These differences were 1 to 3 times higher thanthe LSD005 and were in favor of some F4 lines indicatingthat they perform better than the parents and suggestingthe possibilities of making significant changes through effec-tive selection These findings were in harmony with thoseobtained by Abd El-Shafi [34] Loffler and Busch [40] andAlexander et al [41]

32 Genetic Gain from Direct and Indirect Selection andSelection-Based Index The gains of direct and indirect selec-tion are shown inTable 5The results showed a big variation inthe percentages of gains among the measured traitsThe totalsum of gains per selection varied from minus3611 for headingdate to 3475 for flag leaf area Generally the direct selectionbased on chlorophyll content heading date spikes weightgrain yield and harvest index resulted in negative total gainsOn the other hand flag leaf area plant height number ofspikes 1000-kernel weight number of grains per spike andbiomass recorded positive total gains

The gains obtained by direct selection were higher thanthose of indirect selection But sometimes the indirect selec-tion may be more efficient especially if the secondary traitis highly correlated with yield and is easily measurable [42]The number of grains per spike (1586) followed by aboveground biomass (1230) plant height (1210) and numberof spikes (1084) exhibited the greatest gains from directselection Inversely heading date (minus113) spikes weight(minus499) and harvest index (minus381) showed negative gainvalues Negative gain for heading date is desired in the case ofthis study as the Algerian wheat breeding program designedfor semiarid regions precocity is a crucial criterion adoptedfor selection It is related to the ability of plants to shortentheir cycle so as to decrease their exposure to the late-seasonsirocco weathering

The direct selection for chlorophyll content resulted innegative indirect gains for nearly all the other traits exceptfor the flag leaf area number of grains per spike and harvestindex that showed positive responses For heading date theindirect selection gains were only positive for chlorophyllcontent number of spikes and harvest indexThe correlativeeffects of the selection based on the flag leaf area were desir-able for grain yield although the indirect gain for the remain-ing traits was practically negative

The direct selection for plant height resulted in positiveindirect gains for heading date number of spikes 1000-kernelweight and spikes weight In addition positive responsesof the selection based on yield and yield components wereobserved for grain yield itself heading date 1000-kernelweight and spikes weight However negative indirect gainswere exhibited for other traits including chlorophyll contentflag leaf area and number of grains per spike indicatingthat indirect selection for one variable for gain in another

is unfeasible because there will be a loss in the indirectlyselected variable In cases of negative gains the model is con-sidered inappropriate for selection in this plant material

The highest correlated responses for grain yield weregenerated though indirect selection on the base of theflag leaf area (804 ie 3387 gmminus2) followed by thespike fertility (365 ie 1539 gmminus2) 1000-kernel weight(363 ie 1531 gmminus2) and above ground biomass (274 ie1154 gmminus2)These results indicated that the indirect selectionwas to be more effective in improving the primary trait thanthe indirect selection based on other traits andor on thedirect selection based on the grain yield itself

Several authors have estimated the genetic gains of traitsinvolved in yield determination The results are often incon-sistent and scarce Our results were consistent with thoseof DePauw and Shebeski [43] and Inagaki et al [44] whomentioned that direct selection on the basis of yield is ineffec-tive in early generations Benmahammed et al [45] reportedthe same findings in barley crop Their results showed thatbiomass-based direct selection appearedmore discriminatingthan yield-based selection in their plant material Howeverthe results of this study do not corroborate findings reportedby Lalic et al [11] Mitchell et al [46] Lungu et al [47] andEl-Morshidy et al [48] and who observed the effectiveness ofthe direct selection on grain yieldThedifference in the resultsmay be attributed to differences of breeding material and togenotype times environments

The gains of selection-based index are shown in Table 6The total sum of gains from the selection-based index rangedfrom minus937 to 4286 Five out of seven used indicesshowed positive total gains The base index of Williamsrecorded the highest total gain (4286) followed by theclassic index of Smith and Hazel the index desired gains ofPesek and Baker (2746) and the genotype-ideotype dis-tance index of Cruz (2207) The sum of ranks index ofMulamba and Mock had a very low gain of 374 Subandirsquosand Elstonrsquos indices exhibited negative total gains

The base index of Williams yielded positive responses forall measured traits except for chlorophyll content flag leafarea and harvest index which showed negative gains Thisindex also achieved the highest grain yield response (627ie 2641 gmminus2) compared to the other indices employed inthis study The classic index of Smith and Hazel ranked sec-ond followed by the index of desired gains of Pesek and Bakerthe genotype-ideotype distance index proposed by Cruz andthe sum of ranks index of Mulamba andMock with selectionresponses of 3693 2746 2207 and 374 respectivelyThe single-trait responses varied from one index to anotherwith a more or less balanced distribution of positive andnegative estimates among the traits This shows that themonotrait selection was inadequate because it led to a higherfinal product when considering the grain yield and generatedunfavorable responses in other traits These results indicatedthat methods that combine favorable expected gains shouldbe used in the evaluation of these progeniesThe gains expect-ed through indices for grain yield per se were larger thanthose obtained by direct and indirect monotrait selectionexcept for the flag leaf area (Tables 5 and 6) Mahdy [49]mentioned that selection-based index was predicted to be

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 2: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

2 International Journal of Agronomy

these traits to improve grain yield in diverse environments[8ndash10]

Selection-based index is another approach certainlycomplex but can avoid the limits of the single-trait selec-tion particularly the undesirable between-trait relationsthat present an additional nuisance in breedersrsquo work [11]Selection-based index approach targets the simultaneousimprovement of several traits at the same time includingthe grain yield [12 13] The indices allow the use of a singlevalue in the selection process since the analysis is carriedout by means of linear combinations of phenotypic dataof different traits of agronomic interest with the geneticproperties of a population [14] The objective is to guaranteethe improvement of the population genotypic values and con-sequently the efficiency of the selection process maximizingthe expected genetic gain In this purpose many selectionindices have been used as an effective selection criterion inplant breeding programs on different crops [15] Howeverthe conditions determining the usefulness of an appropriateselection index may vary with individual plant breeder Theobjective of this research paper was to evaluate the efficiencyand applicability of different selection criteria based on theestimation of genetic gain in F4 segregating populations ofbread wheat evaluated under semiarid conditions

2 Materials and Methods

21 Plant Material and Experimental Design This experi-ment had 609 genotypes comprising 20 F4-derived familiesand their 9 parents The history of these families is thatafter the initial crosses in 201011 [18] 30 F2 lines wereselected in each family by using pedigree method and wereevaluated during the 201213 cropping season [19] A totalof 600 lines were planted and harvested in bulk during twoconsecutive growing seasons 201314 and 201415 The F4lines along with their parents were planted in Setif ResearchUnit (36∘151015840N 5∘871015840E 1081 masl) of the Algerian NationalInstitute of Agronomic Research (INRAA) in a modifiedrandomized complete block design with three replicationsThe experimental unit consisted of a single row plot of 2mlength with rows spaced 02m apartThe plots were fertilizedwith 100 kg haminus1 of 46 superphosphate before sowing and75 kg haminus1 of 35 urea at tillering stage Weed control wasperformed chemically using 12 g haminus1 of the Granstar [tribe-nuron methyl] herbicide According to Chennafi et al [20]the climate of the region is a semiarid type continental withcold winter and hot and dry summer Total annual rainfallis around 350mm The soil used was classified as a silty claywith high CaCO3 content [20]

22Measurements The following traits weremeasured as perplot basis Chlorophyll content (CHL Spad) was determinedat heading stage using the SPAD-502 chlorophyll meter(Minolta Camera Co Osaka Japan) Flag leaf area (FLAcm2) was obtained using the formula described by Spagno-letti Zeuli and Qualset [21] FLA (cm2) = 119871 (cm) times 119897 (cm)where 119871 is the flag leaf length and 119897 refers to the flag leafwidth Heading date (HD days) was recorded as the number

Table 1 The skeleton of the analysis of variance

Source of variation df MS 119865-testBlock 119887 minus 1 M1 M1M6Genotype 119892 minus 1 M2 M2M6

Parents 119875 minus 1 M3 M3M6F4 119899 minus 1 M4 M4M6Parents vs F4 1 M5 M5M6

Residual (119892 minus 1)(119887 minus 1) M6 - -Total 119887119892 minus 1 - - - -

of calendar days from January first to the date when 50of the spikes were half way out from flag leaf Plant height(PH cm) was measured at maturity from the soil surface tothe top of the spike awns excluded Above ground biomass(BIO gmminus2) was estimated from a harvested area of 05mlong times 020m interrow spacing which served also to obtainthe grain yield (GY gmminus2) number of spikes (SN mminus2)and spikes weight (SW gmminus2) 1000-kernel weight (TKW g)was obtained from the count and weight of 250-kernel Thenumber of grains per spike (GN) was derived from estimatedvalues of grain yield number of spikes and 1000-kernelweight Harvest index (HI ) was estimated by the ratio be-tween grain yield and above ground biomass

23 Data Analysis Data collected were subjected to analysisof variance following the procedures described by McIntosh[22] the skeleton of the analysis of variance is shown inTable 1 The statistical model used considered the completerandomized block design as 119884119894119895 = 120583 + 119892119894 + 119887119894 + 120576119894119895 where 119884119894119895is the observation of the 119894th genotype evaluated in the 119895threplicate 120583 is the overall mean of the experiment 119892119894 is theeffect of the 119894th genotype 119887119895 is the effect of the 119895th block and120576119894119895 is the effect of the 119894119895th plot

In case 119865-test was significant standard error and criticaldifferences were calculated by using the least significantdifference test at 005 probability level (LSD005) accordingto Steel and Torrie [23] LSD005 = 119905005radic21205902119890 119887 where 119905005is the tabulated value of the 119905-test at 005 probability level for(119892 minus 1)(119887 minus 1) residual degrees of freedom 1205902119890 is the residualvariance and 119887 refers to the number of replications or blocks

The genotypic variance (1205902119892) and residual variance (1205902119890)were calculated and served to determine the following geneticand nongenetic parameters for each trait The coefficient ofexperimental variation (CV) was calculated by CV () =radic1205902119890 119883 where1205902119890 is the residual variance and119883 is the generalmean of the trait The coefficient of genetic variation (CV119892)was calculated by the following equation CV119892 () = radic1205902119892119883where1205902119892 is the genotypic variance and119883 is the generalmeanof the trait The variation index was determined as the ratioof CV119892CV119890 Broad-sense heritability of average progenies(ℎ2bs) was estimated by the expression [24] ℎ2bs () = 100 times[1205902119892(1205902119892+1205902119890 )]Narrow-sense heritability of individualswithinfamilies (ℎ2ns) was determined based on the parent-offspring

International Journal of Agronomy 3

regression To do so two methods were used the first wasby the linear regression of F4 on the parental F3 individualvalues [16] while the second approach was performed withstandardized data of offspring (F4) versus standardized of thecorresponding parent (F3) according to Frey andHorner [17]

The selection gains were estimated among families basedon three selection criteria direct selection indirect selectionand selection-based index considering the selection intensityof 5 of top families The expected gains by direct selectionfor each trait evaluated were estimated by the expression [25]Δ119866119894 = ℎ2119894times119878119894 = ℎ2119894times(119883119904119894minus1198830119894) whereΔ119866119894 is the gainwith thedirect selection carried for the 119894th trait ℎ2119894 is the heritability ofthe 119894th trait 119878119894 refers to the differential selection of the 119894th trait119883119904119894 is themean of the 119894th trait for the selected individuals and1198830119894 is the mean of the 119894th trait in the base population Theexpected gain of direct selection expressed as a percentage ofthe population mean is given by Δ119866119894119894 = (Δ119866119894 times 100)1198830119894

Gains from indirect response to selection were calculatedusing the following expression [25] GS119895(119894) = ℎ2119895 times (119883119894119895 minus1198830119895) = ℎ2119895 times DS119895(119894) where GS119895(119894) is the gain on the 119895th traitwith selection based on the 119894th trait 119883119904119895 is the mean of the119895th trait for the selected individuals based on the 119894th trait1198830119895 is the mean of the 119895th trait ℎ2119895 is the heritability ofthe 119895th trait and DS119895(119894) refers to the differential selection ofthe 119895th trait in which the selected lines presented the bestperformance for the 119894th trait The expected gain of indirectselection expressed as a percentage of the population meanis given by GS119895(119894) = (GS119895(119894) times 100)1198830119894

For the selection-based index the following methodolo-gieswere used for gains estimation the classic index proposedby Smith [26] and Hazel [27] the base index of Williams[28] the free weights and parameters index of Elston [29]the index of desired gains of Pesek and Baker [30] the multi-plicative index of Subandi et al [31] the sum of ranks index ofMulamba andMock [32] and the genotype-ideotype distanceindex proposed by Cruz [25] Each index displays certainparticularities in its calculations and as such applicationis generally laborious due to the need to assign adequateeconomic weights to each trait Based on the differentanalytical procedures of selection the best populations wereidentified and the gains from selection were calculated Allstatistical analyses were carried out using Genes software [33]and a Microsoft Excelcopy spreadsheet

3 Results and Discussion

31 Genetic Variability and Heritability The analysis of vari-ance revealed significant genotype effect for almost all thetraits under study except for flag leaf area and grain yieldwhich were not significant at 005 probability level (Table 2)This provides evidence of the presence of sufficient geneticvariability among parents and hybrids that can be exploitedin wheat breeding program through selection Partitioningthe genotype effect indicated significant differences betweenall parents for HD PH SN TKW SW GN BIO and HIand significant interpopulation differences for HD PH andGN The contrast ldquoParents versus F4rdquo was highly significant

for HD PH and SN while the F4 populations exhibitedsignificant differences for nearly all the observed quantitativephenotypic traits except for CHL FLA GN and HI whichwere not significant at the 5 probability level The resultsof this study corroborate those of Abd El-Shafi [34] whoreported significant and highly significant differences amonggenotypes (families + parents) and families for all studiedtraits across three segregating generations F2 F3 and F4Thisauthor also reported that greater response to selection canbe expected from selection in cross having greater phenotypicand genotypic variances

The coefficient of variation (CV) presented values be-tween 07 and 268 for heading date and grain yield respec-tively (Table 2) The CV above 20 is considered high indi-cating high dispersion of the experimental data which mayhave been caused by the genetic and phenotypic differencesbetween the studiedmaterials HighCV estimate obtained forgrain yield can be explained by the fact that it is quantitativetrait governed by several genes and highly influenced by theenvironment

The variances values coefficient of variation and geneticparameters estimates for wheat traits studied are presentedin Table 3 Broad-sense heritability is the proportion of totalphenotypic variation due to all genetic effects The knowl-edge of the genotypic determination coefficient (ℎ2bs) allowsestablishing an estimate of the genetic gain to be obtained anddefines the best strategy to be used in the plant breeding pro-gram [35] In this study the estimated broad-sense heritabil-ity varied from 000 to 9180The highest values were foundfor heading date (9180) followed by plant height (8082)1000-kernel weight (7290) biomass (6533) and numberof grains per spike (6164) indicating that these traits arehighly heritable among the genotypes evaluatedThese resultscan be confirmed with the values obtained by the CV119892CV119890ratio that were close to or greater than 1 for these traits sug-gesting satisfactory conditions for selection [36] Moderateestimates of ℎ2bs occurred for the number of chlorophyllcontents (4031) number of spikes (5200) spikes weight(5485) and harvest index (4067) On the other handthe lowest values of ℎ2bs were found for the flag leaf area(000) and grain yield (1939) These traits exhibited alsolow CV119892CV119890 ratio values indicating the dominant effect ofthe environment on crop

Generally literature indicates widely varying narrow-sense heritability estimates Mesele et al [37] reported highheritability values for days to heading days to maturityand 1000-kernel weight moderate estimates for grain fillingperiod spike length number of spikelets per plant grainsper spike and harvest index and low values for number oftillers per plant biomass yield and grain yield Evaluatingseven F2 populations derived through cross combinations offive parental varietieslines of bread wheat Saleem et al [38]found low to high broad-sense heritability values rangingfrom 475 to 926 depending on the trait and the crossThefindings of Yaqoob [39] showed that heritability estimateswere low for number of tillers per plant (20) grainsper spike (2681) days to maturity (3013) spike length(3666) and 1000-kernel weight (3868) moderate forplant height (4579) and high for heading date (8473)

4 International Journal of Agronomy

Table 2 Analysis of variance of different bread wheat traits studied

SV Bloc Genotypes Parents F4 Parents versus F4 Error CV ()df 2 28 8 19 1 56CHL 86 190lowastlowast 95ns 238lowastlowast 36ns 86 66HD 15 90lowastlowast 183lowastlowast 25lowastlowast 587lowastlowast 07 07FLA 894 111

ns69ns 66ns 01ns 149 204

PH 91 863lowastlowast 1030lowastlowast 800lowastlowast 726lowast 166 63SN 11831 111875lowastlowast 260924lowastlowast 26578ns 539945lowastlowast 53711 159TKW 335 282lowastlowast 394lowastlowast 127ns 2321lowastlowast 77 74SW 1170346 448736lowastlowast 744904lowastlowast 182488ns 3137463lowastlowast 202598 209GN 366 447lowastlowast 613lowastlowast 370lowast 410ns 171 168BIO 811895 1310774lowastlowast 2431421lowastlowast 436017ns 8965592lowastlowast 454408 185GY 341492 157279ns 268354ns 110965ns 562059lowast 126778 268HI 1222 623lowast 1366lowastlowast 361ns 214ns 37 168CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) HI harvest index () and nslowast and lowastlowast nonsignificant and significant effect at 005 and 001 probability

Table 3 Genetic and nongenetic parameters of different bread wheat traits studied

Traits 1205902119901 1205902119890 1205902119892 ℎ2bs ℎ2ns (SK) ℎ2ns (FH) CV119892 () CV119892CV119890Chl 336 200 135 4031 2855 3013 264 047HD 301 025 276 9180 1669 3432 131 193FLA 368 498 000 000 000 000 000 000PH 2878 552 2326 8082 2727 2727 743 119SN 372917 179037 193879 5200 000 000 959 060TKW 941 255 686 7290 3164 2599 698 095SW 1495791 675325 820467 5485 866 1070 1319 064GN 1467 563 904 6164 1670 714 1228 073BIO 4369245 1514693 2854552 6533 337 593 1448 079GY 524264 422592 101671 1939 1013 1231 758 028HI 2076 1231 845 4067 1410 1497 803 048CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY Grain yield (gmminus2) HI harvest index () ℎ2bsbroad-sense heritability ℎ2ns (SK) narrow-sense heritability according to Smith and Kinman [16] and ℎ2ns (FH) narrow-sense heritability according to Freyand Horner [17]

and grain yield (9983) His results also indicated that mostof these traits exhibited low heritability under drought stressconditions suggesting the presence of high genotype times envi-ronment interactions that affected the crop behavior

Narrow-sense heritability is the proportion of the totalphenotypic variation that is due to the additive effects ofgenes This component of variation is important because it isthe only variation that natural selection can act on Henceℎ2ns determines the resemblance of offspring to their parentsand the populationrsquos evolutionary response to selectionTherecan be considerable nonadditive genetic variance but thisdoes not contribute to the resemblance between parents andoffspring or the response to selection Low to moderatenarrow-sense heritability values were recorded in this studyChlorophyll content heading date plant height 1000-kernelweight and number of grains per spike recorded the highestestimates These traits were less influenced by the environ-mental factors and would respond positively to a selection

pressure in the current breeding programThe lowheritabilityvalues can be explained by the change occurring in thesegregating lines behavior from the precedent to the currentgeneration This change may be due to nonadditive geneaction andor high environmental factors effects

Means of the variables measured showed that the bestvalues varied depending on the cross and the trait and thefew populations had the best performances for several traitsat the same time (Table 4)The best grain yielding population(5575 g) was Ain Abid times El-Wifak which had also the highestaverage for the number of grains per spike (3147 grains)spikes weight (8573 g) above ground biomass (13503 g) andharvest index (413) Ain Abid timesHidhab cross combinationhad the longest vegetative cycle with an average of 1278days and presented the highest mean for the flag leaf area(210 cm2) Acsad1069 times El-Wifak had the highest average forthe chlorophyll content (540 Spad) Acsad1069 times Mahon-Demias was the tallest (7523 cm) while Acsad1135 times Rmada

International Journal of Agronomy 5

Table4Means

ofthem

easuredtraitsfor2

0F 4

breadwheatpo

pulatio

ns

Popu

latio

nCH

LHD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Acsad 901timesM

ahon

-Dem

ias

430

1260

201

737

4483

386

6340

220

11350

3792

334

Acsad 901timesR

mada

445

1252

199

608

4424

346

6336

281

10497

4255

405

Acsad 901timesH

idhab

425

1259

198

657

4267

334

6167

230

10983

3255

296

Acsad 901timesE

l-Wifa

k451

1249

204

704

4673

332

6843

304

11490

4604

401

Acsad 899timesM

ahon

-Dem

ias

437

1257

169

708

4760

364

6260

205

11517

3668

319

Acsad 899timesR

mada

434

1250

166

595

4797

378

6257

200

10327

3688

357

Acsad 899timesH

idhab

444

1269

182

566

4060

350

5980

228

9970

3360

337

Acsad 899timesE

l-Wifa

k454

1250

181

631

4147

355

5667

242

9657

3577

370

Acsad 1135timesM

ahon

-Dem

ias

416

1261

170

681

4567

398

6040

210

10737

3834

357

Acsad 1135timesR

mada

422

1252

168

615

5037

369

6853

240

11137

4404

396

Acsad 1135timesH

idhab

415

1270

175

688

4770

395

7447

232

12687

4509

355

Acsad 1135timesE

l-Wifa

k455

1252

170

646

3970

346

4893

221

8790

3099

353

Acsad 1069timesM

ahon

-Dem

ias

438

1258

182

752

4390

392

6233

217

11937

3713

311

Acsad 1069timesR

mada

442

1250

173

617

4147

348

5810

245

9617

3608

375

Acsad 1069timesH

idhab

467

1259

185

605

4113

350

6170

266

9767

3953

405

Acsad 1069timesE

l-Wifa

k540

1250

184

665

4233

346

6110

279

9940

4033

406

Ain

AbidtimesM

ahon

-Dem

ias

420

1277

200

727

4547

388

7473

272

12867

4818

375

Ain

AbidtimesR

mada

403

1263

205

658

4357

365

6993

282

11717

4581

391

Ain

AbidtimesH

idhab

432

1278

210

602

4050

368

6933

304

11307

4628

409

Ain

AbidtimesE

l-Wifa

k436

1269

203

651

4693

377

8573

315

13503

5575

413

ParentsR

ange

391

1250

87

530

2700

300

3200

975300

1850

141

522

1320

391

820

8000

490

12600

394

23300

8090

636

F 4Ra

nge

334

1240

76420

800

190

800

931500

430

126

538

1360

364

1000

8500

510

14800

568

24200

9210

700

LSD005

48

1463

67

1197

45

2352

68

3532

1851

99CH

Lchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)andHIharvestind

ex(

)

6 International Journal of Agronomy

exhibited the highest average for the number of spikes (5037spikes) For 1000-kernel weight Acsad1135 timesMahon-Demiaswas the best population with an average of 398 g

As the contrast ldquoParents versus F4rdquo was significant forPH SN TKW SW BIO and GY and compared with theleast significant difference (LSD005) significant differencesexisted between the parents and hybrids values for thesemea-sured traits These differences were 1 to 3 times higher thanthe LSD005 and were in favor of some F4 lines indicatingthat they perform better than the parents and suggestingthe possibilities of making significant changes through effec-tive selection These findings were in harmony with thoseobtained by Abd El-Shafi [34] Loffler and Busch [40] andAlexander et al [41]

32 Genetic Gain from Direct and Indirect Selection andSelection-Based Index The gains of direct and indirect selec-tion are shown inTable 5The results showed a big variation inthe percentages of gains among the measured traitsThe totalsum of gains per selection varied from minus3611 for headingdate to 3475 for flag leaf area Generally the direct selectionbased on chlorophyll content heading date spikes weightgrain yield and harvest index resulted in negative total gainsOn the other hand flag leaf area plant height number ofspikes 1000-kernel weight number of grains per spike andbiomass recorded positive total gains

The gains obtained by direct selection were higher thanthose of indirect selection But sometimes the indirect selec-tion may be more efficient especially if the secondary traitis highly correlated with yield and is easily measurable [42]The number of grains per spike (1586) followed by aboveground biomass (1230) plant height (1210) and numberof spikes (1084) exhibited the greatest gains from directselection Inversely heading date (minus113) spikes weight(minus499) and harvest index (minus381) showed negative gainvalues Negative gain for heading date is desired in the case ofthis study as the Algerian wheat breeding program designedfor semiarid regions precocity is a crucial criterion adoptedfor selection It is related to the ability of plants to shortentheir cycle so as to decrease their exposure to the late-seasonsirocco weathering

The direct selection for chlorophyll content resulted innegative indirect gains for nearly all the other traits exceptfor the flag leaf area number of grains per spike and harvestindex that showed positive responses For heading date theindirect selection gains were only positive for chlorophyllcontent number of spikes and harvest indexThe correlativeeffects of the selection based on the flag leaf area were desir-able for grain yield although the indirect gain for the remain-ing traits was practically negative

The direct selection for plant height resulted in positiveindirect gains for heading date number of spikes 1000-kernelweight and spikes weight In addition positive responsesof the selection based on yield and yield components wereobserved for grain yield itself heading date 1000-kernelweight and spikes weight However negative indirect gainswere exhibited for other traits including chlorophyll contentflag leaf area and number of grains per spike indicatingthat indirect selection for one variable for gain in another

is unfeasible because there will be a loss in the indirectlyselected variable In cases of negative gains the model is con-sidered inappropriate for selection in this plant material

The highest correlated responses for grain yield weregenerated though indirect selection on the base of theflag leaf area (804 ie 3387 gmminus2) followed by thespike fertility (365 ie 1539 gmminus2) 1000-kernel weight(363 ie 1531 gmminus2) and above ground biomass (274 ie1154 gmminus2)These results indicated that the indirect selectionwas to be more effective in improving the primary trait thanthe indirect selection based on other traits andor on thedirect selection based on the grain yield itself

Several authors have estimated the genetic gains of traitsinvolved in yield determination The results are often incon-sistent and scarce Our results were consistent with thoseof DePauw and Shebeski [43] and Inagaki et al [44] whomentioned that direct selection on the basis of yield is ineffec-tive in early generations Benmahammed et al [45] reportedthe same findings in barley crop Their results showed thatbiomass-based direct selection appearedmore discriminatingthan yield-based selection in their plant material Howeverthe results of this study do not corroborate findings reportedby Lalic et al [11] Mitchell et al [46] Lungu et al [47] andEl-Morshidy et al [48] and who observed the effectiveness ofthe direct selection on grain yieldThedifference in the resultsmay be attributed to differences of breeding material and togenotype times environments

The gains of selection-based index are shown in Table 6The total sum of gains from the selection-based index rangedfrom minus937 to 4286 Five out of seven used indicesshowed positive total gains The base index of Williamsrecorded the highest total gain (4286) followed by theclassic index of Smith and Hazel the index desired gains ofPesek and Baker (2746) and the genotype-ideotype dis-tance index of Cruz (2207) The sum of ranks index ofMulamba and Mock had a very low gain of 374 Subandirsquosand Elstonrsquos indices exhibited negative total gains

The base index of Williams yielded positive responses forall measured traits except for chlorophyll content flag leafarea and harvest index which showed negative gains Thisindex also achieved the highest grain yield response (627ie 2641 gmminus2) compared to the other indices employed inthis study The classic index of Smith and Hazel ranked sec-ond followed by the index of desired gains of Pesek and Bakerthe genotype-ideotype distance index proposed by Cruz andthe sum of ranks index of Mulamba andMock with selectionresponses of 3693 2746 2207 and 374 respectivelyThe single-trait responses varied from one index to anotherwith a more or less balanced distribution of positive andnegative estimates among the traits This shows that themonotrait selection was inadequate because it led to a higherfinal product when considering the grain yield and generatedunfavorable responses in other traits These results indicatedthat methods that combine favorable expected gains shouldbe used in the evaluation of these progeniesThe gains expect-ed through indices for grain yield per se were larger thanthose obtained by direct and indirect monotrait selectionexcept for the flag leaf area (Tables 5 and 6) Mahdy [49]mentioned that selection-based index was predicted to be

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 3: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

International Journal of Agronomy 3

regression To do so two methods were used the first wasby the linear regression of F4 on the parental F3 individualvalues [16] while the second approach was performed withstandardized data of offspring (F4) versus standardized of thecorresponding parent (F3) according to Frey andHorner [17]

The selection gains were estimated among families basedon three selection criteria direct selection indirect selectionand selection-based index considering the selection intensityof 5 of top families The expected gains by direct selectionfor each trait evaluated were estimated by the expression [25]Δ119866119894 = ℎ2119894times119878119894 = ℎ2119894times(119883119904119894minus1198830119894) whereΔ119866119894 is the gainwith thedirect selection carried for the 119894th trait ℎ2119894 is the heritability ofthe 119894th trait 119878119894 refers to the differential selection of the 119894th trait119883119904119894 is themean of the 119894th trait for the selected individuals and1198830119894 is the mean of the 119894th trait in the base population Theexpected gain of direct selection expressed as a percentage ofthe population mean is given by Δ119866119894119894 = (Δ119866119894 times 100)1198830119894

Gains from indirect response to selection were calculatedusing the following expression [25] GS119895(119894) = ℎ2119895 times (119883119894119895 minus1198830119895) = ℎ2119895 times DS119895(119894) where GS119895(119894) is the gain on the 119895th traitwith selection based on the 119894th trait 119883119904119895 is the mean of the119895th trait for the selected individuals based on the 119894th trait1198830119895 is the mean of the 119895th trait ℎ2119895 is the heritability ofthe 119895th trait and DS119895(119894) refers to the differential selection ofthe 119895th trait in which the selected lines presented the bestperformance for the 119894th trait The expected gain of indirectselection expressed as a percentage of the population meanis given by GS119895(119894) = (GS119895(119894) times 100)1198830119894

For the selection-based index the following methodolo-gieswere used for gains estimation the classic index proposedby Smith [26] and Hazel [27] the base index of Williams[28] the free weights and parameters index of Elston [29]the index of desired gains of Pesek and Baker [30] the multi-plicative index of Subandi et al [31] the sum of ranks index ofMulamba andMock [32] and the genotype-ideotype distanceindex proposed by Cruz [25] Each index displays certainparticularities in its calculations and as such applicationis generally laborious due to the need to assign adequateeconomic weights to each trait Based on the differentanalytical procedures of selection the best populations wereidentified and the gains from selection were calculated Allstatistical analyses were carried out using Genes software [33]and a Microsoft Excelcopy spreadsheet

3 Results and Discussion

31 Genetic Variability and Heritability The analysis of vari-ance revealed significant genotype effect for almost all thetraits under study except for flag leaf area and grain yieldwhich were not significant at 005 probability level (Table 2)This provides evidence of the presence of sufficient geneticvariability among parents and hybrids that can be exploitedin wheat breeding program through selection Partitioningthe genotype effect indicated significant differences betweenall parents for HD PH SN TKW SW GN BIO and HIand significant interpopulation differences for HD PH andGN The contrast ldquoParents versus F4rdquo was highly significant

for HD PH and SN while the F4 populations exhibitedsignificant differences for nearly all the observed quantitativephenotypic traits except for CHL FLA GN and HI whichwere not significant at the 5 probability level The resultsof this study corroborate those of Abd El-Shafi [34] whoreported significant and highly significant differences amonggenotypes (families + parents) and families for all studiedtraits across three segregating generations F2 F3 and F4Thisauthor also reported that greater response to selection canbe expected from selection in cross having greater phenotypicand genotypic variances

The coefficient of variation (CV) presented values be-tween 07 and 268 for heading date and grain yield respec-tively (Table 2) The CV above 20 is considered high indi-cating high dispersion of the experimental data which mayhave been caused by the genetic and phenotypic differencesbetween the studiedmaterials HighCV estimate obtained forgrain yield can be explained by the fact that it is quantitativetrait governed by several genes and highly influenced by theenvironment

The variances values coefficient of variation and geneticparameters estimates for wheat traits studied are presentedin Table 3 Broad-sense heritability is the proportion of totalphenotypic variation due to all genetic effects The knowl-edge of the genotypic determination coefficient (ℎ2bs) allowsestablishing an estimate of the genetic gain to be obtained anddefines the best strategy to be used in the plant breeding pro-gram [35] In this study the estimated broad-sense heritabil-ity varied from 000 to 9180The highest values were foundfor heading date (9180) followed by plant height (8082)1000-kernel weight (7290) biomass (6533) and numberof grains per spike (6164) indicating that these traits arehighly heritable among the genotypes evaluatedThese resultscan be confirmed with the values obtained by the CV119892CV119890ratio that were close to or greater than 1 for these traits sug-gesting satisfactory conditions for selection [36] Moderateestimates of ℎ2bs occurred for the number of chlorophyllcontents (4031) number of spikes (5200) spikes weight(5485) and harvest index (4067) On the other handthe lowest values of ℎ2bs were found for the flag leaf area(000) and grain yield (1939) These traits exhibited alsolow CV119892CV119890 ratio values indicating the dominant effect ofthe environment on crop

Generally literature indicates widely varying narrow-sense heritability estimates Mesele et al [37] reported highheritability values for days to heading days to maturityand 1000-kernel weight moderate estimates for grain fillingperiod spike length number of spikelets per plant grainsper spike and harvest index and low values for number oftillers per plant biomass yield and grain yield Evaluatingseven F2 populations derived through cross combinations offive parental varietieslines of bread wheat Saleem et al [38]found low to high broad-sense heritability values rangingfrom 475 to 926 depending on the trait and the crossThefindings of Yaqoob [39] showed that heritability estimateswere low for number of tillers per plant (20) grainsper spike (2681) days to maturity (3013) spike length(3666) and 1000-kernel weight (3868) moderate forplant height (4579) and high for heading date (8473)

4 International Journal of Agronomy

Table 2 Analysis of variance of different bread wheat traits studied

SV Bloc Genotypes Parents F4 Parents versus F4 Error CV ()df 2 28 8 19 1 56CHL 86 190lowastlowast 95ns 238lowastlowast 36ns 86 66HD 15 90lowastlowast 183lowastlowast 25lowastlowast 587lowastlowast 07 07FLA 894 111

ns69ns 66ns 01ns 149 204

PH 91 863lowastlowast 1030lowastlowast 800lowastlowast 726lowast 166 63SN 11831 111875lowastlowast 260924lowastlowast 26578ns 539945lowastlowast 53711 159TKW 335 282lowastlowast 394lowastlowast 127ns 2321lowastlowast 77 74SW 1170346 448736lowastlowast 744904lowastlowast 182488ns 3137463lowastlowast 202598 209GN 366 447lowastlowast 613lowastlowast 370lowast 410ns 171 168BIO 811895 1310774lowastlowast 2431421lowastlowast 436017ns 8965592lowastlowast 454408 185GY 341492 157279ns 268354ns 110965ns 562059lowast 126778 268HI 1222 623lowast 1366lowastlowast 361ns 214ns 37 168CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) HI harvest index () and nslowast and lowastlowast nonsignificant and significant effect at 005 and 001 probability

Table 3 Genetic and nongenetic parameters of different bread wheat traits studied

Traits 1205902119901 1205902119890 1205902119892 ℎ2bs ℎ2ns (SK) ℎ2ns (FH) CV119892 () CV119892CV119890Chl 336 200 135 4031 2855 3013 264 047HD 301 025 276 9180 1669 3432 131 193FLA 368 498 000 000 000 000 000 000PH 2878 552 2326 8082 2727 2727 743 119SN 372917 179037 193879 5200 000 000 959 060TKW 941 255 686 7290 3164 2599 698 095SW 1495791 675325 820467 5485 866 1070 1319 064GN 1467 563 904 6164 1670 714 1228 073BIO 4369245 1514693 2854552 6533 337 593 1448 079GY 524264 422592 101671 1939 1013 1231 758 028HI 2076 1231 845 4067 1410 1497 803 048CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY Grain yield (gmminus2) HI harvest index () ℎ2bsbroad-sense heritability ℎ2ns (SK) narrow-sense heritability according to Smith and Kinman [16] and ℎ2ns (FH) narrow-sense heritability according to Freyand Horner [17]

and grain yield (9983) His results also indicated that mostof these traits exhibited low heritability under drought stressconditions suggesting the presence of high genotype times envi-ronment interactions that affected the crop behavior

Narrow-sense heritability is the proportion of the totalphenotypic variation that is due to the additive effects ofgenes This component of variation is important because it isthe only variation that natural selection can act on Henceℎ2ns determines the resemblance of offspring to their parentsand the populationrsquos evolutionary response to selectionTherecan be considerable nonadditive genetic variance but thisdoes not contribute to the resemblance between parents andoffspring or the response to selection Low to moderatenarrow-sense heritability values were recorded in this studyChlorophyll content heading date plant height 1000-kernelweight and number of grains per spike recorded the highestestimates These traits were less influenced by the environ-mental factors and would respond positively to a selection

pressure in the current breeding programThe lowheritabilityvalues can be explained by the change occurring in thesegregating lines behavior from the precedent to the currentgeneration This change may be due to nonadditive geneaction andor high environmental factors effects

Means of the variables measured showed that the bestvalues varied depending on the cross and the trait and thefew populations had the best performances for several traitsat the same time (Table 4)The best grain yielding population(5575 g) was Ain Abid times El-Wifak which had also the highestaverage for the number of grains per spike (3147 grains)spikes weight (8573 g) above ground biomass (13503 g) andharvest index (413) Ain Abid timesHidhab cross combinationhad the longest vegetative cycle with an average of 1278days and presented the highest mean for the flag leaf area(210 cm2) Acsad1069 times El-Wifak had the highest average forthe chlorophyll content (540 Spad) Acsad1069 times Mahon-Demias was the tallest (7523 cm) while Acsad1135 times Rmada

International Journal of Agronomy 5

Table4Means

ofthem

easuredtraitsfor2

0F 4

breadwheatpo

pulatio

ns

Popu

latio

nCH

LHD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Acsad 901timesM

ahon

-Dem

ias

430

1260

201

737

4483

386

6340

220

11350

3792

334

Acsad 901timesR

mada

445

1252

199

608

4424

346

6336

281

10497

4255

405

Acsad 901timesH

idhab

425

1259

198

657

4267

334

6167

230

10983

3255

296

Acsad 901timesE

l-Wifa

k451

1249

204

704

4673

332

6843

304

11490

4604

401

Acsad 899timesM

ahon

-Dem

ias

437

1257

169

708

4760

364

6260

205

11517

3668

319

Acsad 899timesR

mada

434

1250

166

595

4797

378

6257

200

10327

3688

357

Acsad 899timesH

idhab

444

1269

182

566

4060

350

5980

228

9970

3360

337

Acsad 899timesE

l-Wifa

k454

1250

181

631

4147

355

5667

242

9657

3577

370

Acsad 1135timesM

ahon

-Dem

ias

416

1261

170

681

4567

398

6040

210

10737

3834

357

Acsad 1135timesR

mada

422

1252

168

615

5037

369

6853

240

11137

4404

396

Acsad 1135timesH

idhab

415

1270

175

688

4770

395

7447

232

12687

4509

355

Acsad 1135timesE

l-Wifa

k455

1252

170

646

3970

346

4893

221

8790

3099

353

Acsad 1069timesM

ahon

-Dem

ias

438

1258

182

752

4390

392

6233

217

11937

3713

311

Acsad 1069timesR

mada

442

1250

173

617

4147

348

5810

245

9617

3608

375

Acsad 1069timesH

idhab

467

1259

185

605

4113

350

6170

266

9767

3953

405

Acsad 1069timesE

l-Wifa

k540

1250

184

665

4233

346

6110

279

9940

4033

406

Ain

AbidtimesM

ahon

-Dem

ias

420

1277

200

727

4547

388

7473

272

12867

4818

375

Ain

AbidtimesR

mada

403

1263

205

658

4357

365

6993

282

11717

4581

391

Ain

AbidtimesH

idhab

432

1278

210

602

4050

368

6933

304

11307

4628

409

Ain

AbidtimesE

l-Wifa

k436

1269

203

651

4693

377

8573

315

13503

5575

413

ParentsR

ange

391

1250

87

530

2700

300

3200

975300

1850

141

522

1320

391

820

8000

490

12600

394

23300

8090

636

F 4Ra

nge

334

1240

76420

800

190

800

931500

430

126

538

1360

364

1000

8500

510

14800

568

24200

9210

700

LSD005

48

1463

67

1197

45

2352

68

3532

1851

99CH

Lchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)andHIharvestind

ex(

)

6 International Journal of Agronomy

exhibited the highest average for the number of spikes (5037spikes) For 1000-kernel weight Acsad1135 timesMahon-Demiaswas the best population with an average of 398 g

As the contrast ldquoParents versus F4rdquo was significant forPH SN TKW SW BIO and GY and compared with theleast significant difference (LSD005) significant differencesexisted between the parents and hybrids values for thesemea-sured traits These differences were 1 to 3 times higher thanthe LSD005 and were in favor of some F4 lines indicatingthat they perform better than the parents and suggestingthe possibilities of making significant changes through effec-tive selection These findings were in harmony with thoseobtained by Abd El-Shafi [34] Loffler and Busch [40] andAlexander et al [41]

32 Genetic Gain from Direct and Indirect Selection andSelection-Based Index The gains of direct and indirect selec-tion are shown inTable 5The results showed a big variation inthe percentages of gains among the measured traitsThe totalsum of gains per selection varied from minus3611 for headingdate to 3475 for flag leaf area Generally the direct selectionbased on chlorophyll content heading date spikes weightgrain yield and harvest index resulted in negative total gainsOn the other hand flag leaf area plant height number ofspikes 1000-kernel weight number of grains per spike andbiomass recorded positive total gains

The gains obtained by direct selection were higher thanthose of indirect selection But sometimes the indirect selec-tion may be more efficient especially if the secondary traitis highly correlated with yield and is easily measurable [42]The number of grains per spike (1586) followed by aboveground biomass (1230) plant height (1210) and numberof spikes (1084) exhibited the greatest gains from directselection Inversely heading date (minus113) spikes weight(minus499) and harvest index (minus381) showed negative gainvalues Negative gain for heading date is desired in the case ofthis study as the Algerian wheat breeding program designedfor semiarid regions precocity is a crucial criterion adoptedfor selection It is related to the ability of plants to shortentheir cycle so as to decrease their exposure to the late-seasonsirocco weathering

The direct selection for chlorophyll content resulted innegative indirect gains for nearly all the other traits exceptfor the flag leaf area number of grains per spike and harvestindex that showed positive responses For heading date theindirect selection gains were only positive for chlorophyllcontent number of spikes and harvest indexThe correlativeeffects of the selection based on the flag leaf area were desir-able for grain yield although the indirect gain for the remain-ing traits was practically negative

The direct selection for plant height resulted in positiveindirect gains for heading date number of spikes 1000-kernelweight and spikes weight In addition positive responsesof the selection based on yield and yield components wereobserved for grain yield itself heading date 1000-kernelweight and spikes weight However negative indirect gainswere exhibited for other traits including chlorophyll contentflag leaf area and number of grains per spike indicatingthat indirect selection for one variable for gain in another

is unfeasible because there will be a loss in the indirectlyselected variable In cases of negative gains the model is con-sidered inappropriate for selection in this plant material

The highest correlated responses for grain yield weregenerated though indirect selection on the base of theflag leaf area (804 ie 3387 gmminus2) followed by thespike fertility (365 ie 1539 gmminus2) 1000-kernel weight(363 ie 1531 gmminus2) and above ground biomass (274 ie1154 gmminus2)These results indicated that the indirect selectionwas to be more effective in improving the primary trait thanthe indirect selection based on other traits andor on thedirect selection based on the grain yield itself

Several authors have estimated the genetic gains of traitsinvolved in yield determination The results are often incon-sistent and scarce Our results were consistent with thoseof DePauw and Shebeski [43] and Inagaki et al [44] whomentioned that direct selection on the basis of yield is ineffec-tive in early generations Benmahammed et al [45] reportedthe same findings in barley crop Their results showed thatbiomass-based direct selection appearedmore discriminatingthan yield-based selection in their plant material Howeverthe results of this study do not corroborate findings reportedby Lalic et al [11] Mitchell et al [46] Lungu et al [47] andEl-Morshidy et al [48] and who observed the effectiveness ofthe direct selection on grain yieldThedifference in the resultsmay be attributed to differences of breeding material and togenotype times environments

The gains of selection-based index are shown in Table 6The total sum of gains from the selection-based index rangedfrom minus937 to 4286 Five out of seven used indicesshowed positive total gains The base index of Williamsrecorded the highest total gain (4286) followed by theclassic index of Smith and Hazel the index desired gains ofPesek and Baker (2746) and the genotype-ideotype dis-tance index of Cruz (2207) The sum of ranks index ofMulamba and Mock had a very low gain of 374 Subandirsquosand Elstonrsquos indices exhibited negative total gains

The base index of Williams yielded positive responses forall measured traits except for chlorophyll content flag leafarea and harvest index which showed negative gains Thisindex also achieved the highest grain yield response (627ie 2641 gmminus2) compared to the other indices employed inthis study The classic index of Smith and Hazel ranked sec-ond followed by the index of desired gains of Pesek and Bakerthe genotype-ideotype distance index proposed by Cruz andthe sum of ranks index of Mulamba andMock with selectionresponses of 3693 2746 2207 and 374 respectivelyThe single-trait responses varied from one index to anotherwith a more or less balanced distribution of positive andnegative estimates among the traits This shows that themonotrait selection was inadequate because it led to a higherfinal product when considering the grain yield and generatedunfavorable responses in other traits These results indicatedthat methods that combine favorable expected gains shouldbe used in the evaluation of these progeniesThe gains expect-ed through indices for grain yield per se were larger thanthose obtained by direct and indirect monotrait selectionexcept for the flag leaf area (Tables 5 and 6) Mahdy [49]mentioned that selection-based index was predicted to be

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 4: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

4 International Journal of Agronomy

Table 2 Analysis of variance of different bread wheat traits studied

SV Bloc Genotypes Parents F4 Parents versus F4 Error CV ()df 2 28 8 19 1 56CHL 86 190lowastlowast 95ns 238lowastlowast 36ns 86 66HD 15 90lowastlowast 183lowastlowast 25lowastlowast 587lowastlowast 07 07FLA 894 111

ns69ns 66ns 01ns 149 204

PH 91 863lowastlowast 1030lowastlowast 800lowastlowast 726lowast 166 63SN 11831 111875lowastlowast 260924lowastlowast 26578ns 539945lowastlowast 53711 159TKW 335 282lowastlowast 394lowastlowast 127ns 2321lowastlowast 77 74SW 1170346 448736lowastlowast 744904lowastlowast 182488ns 3137463lowastlowast 202598 209GN 366 447lowastlowast 613lowastlowast 370lowast 410ns 171 168BIO 811895 1310774lowastlowast 2431421lowastlowast 436017ns 8965592lowastlowast 454408 185GY 341492 157279ns 268354ns 110965ns 562059lowast 126778 268HI 1222 623lowast 1366lowastlowast 361ns 214ns 37 168CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) HI harvest index () and nslowast and lowastlowast nonsignificant and significant effect at 005 and 001 probability

Table 3 Genetic and nongenetic parameters of different bread wheat traits studied

Traits 1205902119901 1205902119890 1205902119892 ℎ2bs ℎ2ns (SK) ℎ2ns (FH) CV119892 () CV119892CV119890Chl 336 200 135 4031 2855 3013 264 047HD 301 025 276 9180 1669 3432 131 193FLA 368 498 000 000 000 000 000 000PH 2878 552 2326 8082 2727 2727 743 119SN 372917 179037 193879 5200 000 000 959 060TKW 941 255 686 7290 3164 2599 698 095SW 1495791 675325 820467 5485 866 1070 1319 064GN 1467 563 904 6164 1670 714 1228 073BIO 4369245 1514693 2854552 6533 337 593 1448 079GY 524264 422592 101671 1939 1013 1231 758 028HI 2076 1231 845 4067 1410 1497 803 048CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY Grain yield (gmminus2) HI harvest index () ℎ2bsbroad-sense heritability ℎ2ns (SK) narrow-sense heritability according to Smith and Kinman [16] and ℎ2ns (FH) narrow-sense heritability according to Freyand Horner [17]

and grain yield (9983) His results also indicated that mostof these traits exhibited low heritability under drought stressconditions suggesting the presence of high genotype times envi-ronment interactions that affected the crop behavior

Narrow-sense heritability is the proportion of the totalphenotypic variation that is due to the additive effects ofgenes This component of variation is important because it isthe only variation that natural selection can act on Henceℎ2ns determines the resemblance of offspring to their parentsand the populationrsquos evolutionary response to selectionTherecan be considerable nonadditive genetic variance but thisdoes not contribute to the resemblance between parents andoffspring or the response to selection Low to moderatenarrow-sense heritability values were recorded in this studyChlorophyll content heading date plant height 1000-kernelweight and number of grains per spike recorded the highestestimates These traits were less influenced by the environ-mental factors and would respond positively to a selection

pressure in the current breeding programThe lowheritabilityvalues can be explained by the change occurring in thesegregating lines behavior from the precedent to the currentgeneration This change may be due to nonadditive geneaction andor high environmental factors effects

Means of the variables measured showed that the bestvalues varied depending on the cross and the trait and thefew populations had the best performances for several traitsat the same time (Table 4)The best grain yielding population(5575 g) was Ain Abid times El-Wifak which had also the highestaverage for the number of grains per spike (3147 grains)spikes weight (8573 g) above ground biomass (13503 g) andharvest index (413) Ain Abid timesHidhab cross combinationhad the longest vegetative cycle with an average of 1278days and presented the highest mean for the flag leaf area(210 cm2) Acsad1069 times El-Wifak had the highest average forthe chlorophyll content (540 Spad) Acsad1069 times Mahon-Demias was the tallest (7523 cm) while Acsad1135 times Rmada

International Journal of Agronomy 5

Table4Means

ofthem

easuredtraitsfor2

0F 4

breadwheatpo

pulatio

ns

Popu

latio

nCH

LHD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Acsad 901timesM

ahon

-Dem

ias

430

1260

201

737

4483

386

6340

220

11350

3792

334

Acsad 901timesR

mada

445

1252

199

608

4424

346

6336

281

10497

4255

405

Acsad 901timesH

idhab

425

1259

198

657

4267

334

6167

230

10983

3255

296

Acsad 901timesE

l-Wifa

k451

1249

204

704

4673

332

6843

304

11490

4604

401

Acsad 899timesM

ahon

-Dem

ias

437

1257

169

708

4760

364

6260

205

11517

3668

319

Acsad 899timesR

mada

434

1250

166

595

4797

378

6257

200

10327

3688

357

Acsad 899timesH

idhab

444

1269

182

566

4060

350

5980

228

9970

3360

337

Acsad 899timesE

l-Wifa

k454

1250

181

631

4147

355

5667

242

9657

3577

370

Acsad 1135timesM

ahon

-Dem

ias

416

1261

170

681

4567

398

6040

210

10737

3834

357

Acsad 1135timesR

mada

422

1252

168

615

5037

369

6853

240

11137

4404

396

Acsad 1135timesH

idhab

415

1270

175

688

4770

395

7447

232

12687

4509

355

Acsad 1135timesE

l-Wifa

k455

1252

170

646

3970

346

4893

221

8790

3099

353

Acsad 1069timesM

ahon

-Dem

ias

438

1258

182

752

4390

392

6233

217

11937

3713

311

Acsad 1069timesR

mada

442

1250

173

617

4147

348

5810

245

9617

3608

375

Acsad 1069timesH

idhab

467

1259

185

605

4113

350

6170

266

9767

3953

405

Acsad 1069timesE

l-Wifa

k540

1250

184

665

4233

346

6110

279

9940

4033

406

Ain

AbidtimesM

ahon

-Dem

ias

420

1277

200

727

4547

388

7473

272

12867

4818

375

Ain

AbidtimesR

mada

403

1263

205

658

4357

365

6993

282

11717

4581

391

Ain

AbidtimesH

idhab

432

1278

210

602

4050

368

6933

304

11307

4628

409

Ain

AbidtimesE

l-Wifa

k436

1269

203

651

4693

377

8573

315

13503

5575

413

ParentsR

ange

391

1250

87

530

2700

300

3200

975300

1850

141

522

1320

391

820

8000

490

12600

394

23300

8090

636

F 4Ra

nge

334

1240

76420

800

190

800

931500

430

126

538

1360

364

1000

8500

510

14800

568

24200

9210

700

LSD005

48

1463

67

1197

45

2352

68

3532

1851

99CH

Lchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)andHIharvestind

ex(

)

6 International Journal of Agronomy

exhibited the highest average for the number of spikes (5037spikes) For 1000-kernel weight Acsad1135 timesMahon-Demiaswas the best population with an average of 398 g

As the contrast ldquoParents versus F4rdquo was significant forPH SN TKW SW BIO and GY and compared with theleast significant difference (LSD005) significant differencesexisted between the parents and hybrids values for thesemea-sured traits These differences were 1 to 3 times higher thanthe LSD005 and were in favor of some F4 lines indicatingthat they perform better than the parents and suggestingthe possibilities of making significant changes through effec-tive selection These findings were in harmony with thoseobtained by Abd El-Shafi [34] Loffler and Busch [40] andAlexander et al [41]

32 Genetic Gain from Direct and Indirect Selection andSelection-Based Index The gains of direct and indirect selec-tion are shown inTable 5The results showed a big variation inthe percentages of gains among the measured traitsThe totalsum of gains per selection varied from minus3611 for headingdate to 3475 for flag leaf area Generally the direct selectionbased on chlorophyll content heading date spikes weightgrain yield and harvest index resulted in negative total gainsOn the other hand flag leaf area plant height number ofspikes 1000-kernel weight number of grains per spike andbiomass recorded positive total gains

The gains obtained by direct selection were higher thanthose of indirect selection But sometimes the indirect selec-tion may be more efficient especially if the secondary traitis highly correlated with yield and is easily measurable [42]The number of grains per spike (1586) followed by aboveground biomass (1230) plant height (1210) and numberof spikes (1084) exhibited the greatest gains from directselection Inversely heading date (minus113) spikes weight(minus499) and harvest index (minus381) showed negative gainvalues Negative gain for heading date is desired in the case ofthis study as the Algerian wheat breeding program designedfor semiarid regions precocity is a crucial criterion adoptedfor selection It is related to the ability of plants to shortentheir cycle so as to decrease their exposure to the late-seasonsirocco weathering

The direct selection for chlorophyll content resulted innegative indirect gains for nearly all the other traits exceptfor the flag leaf area number of grains per spike and harvestindex that showed positive responses For heading date theindirect selection gains were only positive for chlorophyllcontent number of spikes and harvest indexThe correlativeeffects of the selection based on the flag leaf area were desir-able for grain yield although the indirect gain for the remain-ing traits was practically negative

The direct selection for plant height resulted in positiveindirect gains for heading date number of spikes 1000-kernelweight and spikes weight In addition positive responsesof the selection based on yield and yield components wereobserved for grain yield itself heading date 1000-kernelweight and spikes weight However negative indirect gainswere exhibited for other traits including chlorophyll contentflag leaf area and number of grains per spike indicatingthat indirect selection for one variable for gain in another

is unfeasible because there will be a loss in the indirectlyselected variable In cases of negative gains the model is con-sidered inappropriate for selection in this plant material

The highest correlated responses for grain yield weregenerated though indirect selection on the base of theflag leaf area (804 ie 3387 gmminus2) followed by thespike fertility (365 ie 1539 gmminus2) 1000-kernel weight(363 ie 1531 gmminus2) and above ground biomass (274 ie1154 gmminus2)These results indicated that the indirect selectionwas to be more effective in improving the primary trait thanthe indirect selection based on other traits andor on thedirect selection based on the grain yield itself

Several authors have estimated the genetic gains of traitsinvolved in yield determination The results are often incon-sistent and scarce Our results were consistent with thoseof DePauw and Shebeski [43] and Inagaki et al [44] whomentioned that direct selection on the basis of yield is ineffec-tive in early generations Benmahammed et al [45] reportedthe same findings in barley crop Their results showed thatbiomass-based direct selection appearedmore discriminatingthan yield-based selection in their plant material Howeverthe results of this study do not corroborate findings reportedby Lalic et al [11] Mitchell et al [46] Lungu et al [47] andEl-Morshidy et al [48] and who observed the effectiveness ofthe direct selection on grain yieldThedifference in the resultsmay be attributed to differences of breeding material and togenotype times environments

The gains of selection-based index are shown in Table 6The total sum of gains from the selection-based index rangedfrom minus937 to 4286 Five out of seven used indicesshowed positive total gains The base index of Williamsrecorded the highest total gain (4286) followed by theclassic index of Smith and Hazel the index desired gains ofPesek and Baker (2746) and the genotype-ideotype dis-tance index of Cruz (2207) The sum of ranks index ofMulamba and Mock had a very low gain of 374 Subandirsquosand Elstonrsquos indices exhibited negative total gains

The base index of Williams yielded positive responses forall measured traits except for chlorophyll content flag leafarea and harvest index which showed negative gains Thisindex also achieved the highest grain yield response (627ie 2641 gmminus2) compared to the other indices employed inthis study The classic index of Smith and Hazel ranked sec-ond followed by the index of desired gains of Pesek and Bakerthe genotype-ideotype distance index proposed by Cruz andthe sum of ranks index of Mulamba andMock with selectionresponses of 3693 2746 2207 and 374 respectivelyThe single-trait responses varied from one index to anotherwith a more or less balanced distribution of positive andnegative estimates among the traits This shows that themonotrait selection was inadequate because it led to a higherfinal product when considering the grain yield and generatedunfavorable responses in other traits These results indicatedthat methods that combine favorable expected gains shouldbe used in the evaluation of these progeniesThe gains expect-ed through indices for grain yield per se were larger thanthose obtained by direct and indirect monotrait selectionexcept for the flag leaf area (Tables 5 and 6) Mahdy [49]mentioned that selection-based index was predicted to be

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 5: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

International Journal of Agronomy 5

Table4Means

ofthem

easuredtraitsfor2

0F 4

breadwheatpo

pulatio

ns

Popu

latio

nCH

LHD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Acsad 901timesM

ahon

-Dem

ias

430

1260

201

737

4483

386

6340

220

11350

3792

334

Acsad 901timesR

mada

445

1252

199

608

4424

346

6336

281

10497

4255

405

Acsad 901timesH

idhab

425

1259

198

657

4267

334

6167

230

10983

3255

296

Acsad 901timesE

l-Wifa

k451

1249

204

704

4673

332

6843

304

11490

4604

401

Acsad 899timesM

ahon

-Dem

ias

437

1257

169

708

4760

364

6260

205

11517

3668

319

Acsad 899timesR

mada

434

1250

166

595

4797

378

6257

200

10327

3688

357

Acsad 899timesH

idhab

444

1269

182

566

4060

350

5980

228

9970

3360

337

Acsad 899timesE

l-Wifa

k454

1250

181

631

4147

355

5667

242

9657

3577

370

Acsad 1135timesM

ahon

-Dem

ias

416

1261

170

681

4567

398

6040

210

10737

3834

357

Acsad 1135timesR

mada

422

1252

168

615

5037

369

6853

240

11137

4404

396

Acsad 1135timesH

idhab

415

1270

175

688

4770

395

7447

232

12687

4509

355

Acsad 1135timesE

l-Wifa

k455

1252

170

646

3970

346

4893

221

8790

3099

353

Acsad 1069timesM

ahon

-Dem

ias

438

1258

182

752

4390

392

6233

217

11937

3713

311

Acsad 1069timesR

mada

442

1250

173

617

4147

348

5810

245

9617

3608

375

Acsad 1069timesH

idhab

467

1259

185

605

4113

350

6170

266

9767

3953

405

Acsad 1069timesE

l-Wifa

k540

1250

184

665

4233

346

6110

279

9940

4033

406

Ain

AbidtimesM

ahon

-Dem

ias

420

1277

200

727

4547

388

7473

272

12867

4818

375

Ain

AbidtimesR

mada

403

1263

205

658

4357

365

6993

282

11717

4581

391

Ain

AbidtimesH

idhab

432

1278

210

602

4050

368

6933

304

11307

4628

409

Ain

AbidtimesE

l-Wifa

k436

1269

203

651

4693

377

8573

315

13503

5575

413

ParentsR

ange

391

1250

87

530

2700

300

3200

975300

1850

141

522

1320

391

820

8000

490

12600

394

23300

8090

636

F 4Ra

nge

334

1240

76420

800

190

800

931500

430

126

538

1360

364

1000

8500

510

14800

568

24200

9210

700

LSD005

48

1463

67

1197

45

2352

68

3532

1851

99CH

Lchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)andHIharvestind

ex(

)

6 International Journal of Agronomy

exhibited the highest average for the number of spikes (5037spikes) For 1000-kernel weight Acsad1135 timesMahon-Demiaswas the best population with an average of 398 g

As the contrast ldquoParents versus F4rdquo was significant forPH SN TKW SW BIO and GY and compared with theleast significant difference (LSD005) significant differencesexisted between the parents and hybrids values for thesemea-sured traits These differences were 1 to 3 times higher thanthe LSD005 and were in favor of some F4 lines indicatingthat they perform better than the parents and suggestingthe possibilities of making significant changes through effec-tive selection These findings were in harmony with thoseobtained by Abd El-Shafi [34] Loffler and Busch [40] andAlexander et al [41]

32 Genetic Gain from Direct and Indirect Selection andSelection-Based Index The gains of direct and indirect selec-tion are shown inTable 5The results showed a big variation inthe percentages of gains among the measured traitsThe totalsum of gains per selection varied from minus3611 for headingdate to 3475 for flag leaf area Generally the direct selectionbased on chlorophyll content heading date spikes weightgrain yield and harvest index resulted in negative total gainsOn the other hand flag leaf area plant height number ofspikes 1000-kernel weight number of grains per spike andbiomass recorded positive total gains

The gains obtained by direct selection were higher thanthose of indirect selection But sometimes the indirect selec-tion may be more efficient especially if the secondary traitis highly correlated with yield and is easily measurable [42]The number of grains per spike (1586) followed by aboveground biomass (1230) plant height (1210) and numberof spikes (1084) exhibited the greatest gains from directselection Inversely heading date (minus113) spikes weight(minus499) and harvest index (minus381) showed negative gainvalues Negative gain for heading date is desired in the case ofthis study as the Algerian wheat breeding program designedfor semiarid regions precocity is a crucial criterion adoptedfor selection It is related to the ability of plants to shortentheir cycle so as to decrease their exposure to the late-seasonsirocco weathering

The direct selection for chlorophyll content resulted innegative indirect gains for nearly all the other traits exceptfor the flag leaf area number of grains per spike and harvestindex that showed positive responses For heading date theindirect selection gains were only positive for chlorophyllcontent number of spikes and harvest indexThe correlativeeffects of the selection based on the flag leaf area were desir-able for grain yield although the indirect gain for the remain-ing traits was practically negative

The direct selection for plant height resulted in positiveindirect gains for heading date number of spikes 1000-kernelweight and spikes weight In addition positive responsesof the selection based on yield and yield components wereobserved for grain yield itself heading date 1000-kernelweight and spikes weight However negative indirect gainswere exhibited for other traits including chlorophyll contentflag leaf area and number of grains per spike indicatingthat indirect selection for one variable for gain in another

is unfeasible because there will be a loss in the indirectlyselected variable In cases of negative gains the model is con-sidered inappropriate for selection in this plant material

The highest correlated responses for grain yield weregenerated though indirect selection on the base of theflag leaf area (804 ie 3387 gmminus2) followed by thespike fertility (365 ie 1539 gmminus2) 1000-kernel weight(363 ie 1531 gmminus2) and above ground biomass (274 ie1154 gmminus2)These results indicated that the indirect selectionwas to be more effective in improving the primary trait thanthe indirect selection based on other traits andor on thedirect selection based on the grain yield itself

Several authors have estimated the genetic gains of traitsinvolved in yield determination The results are often incon-sistent and scarce Our results were consistent with thoseof DePauw and Shebeski [43] and Inagaki et al [44] whomentioned that direct selection on the basis of yield is ineffec-tive in early generations Benmahammed et al [45] reportedthe same findings in barley crop Their results showed thatbiomass-based direct selection appearedmore discriminatingthan yield-based selection in their plant material Howeverthe results of this study do not corroborate findings reportedby Lalic et al [11] Mitchell et al [46] Lungu et al [47] andEl-Morshidy et al [48] and who observed the effectiveness ofthe direct selection on grain yieldThedifference in the resultsmay be attributed to differences of breeding material and togenotype times environments

The gains of selection-based index are shown in Table 6The total sum of gains from the selection-based index rangedfrom minus937 to 4286 Five out of seven used indicesshowed positive total gains The base index of Williamsrecorded the highest total gain (4286) followed by theclassic index of Smith and Hazel the index desired gains ofPesek and Baker (2746) and the genotype-ideotype dis-tance index of Cruz (2207) The sum of ranks index ofMulamba and Mock had a very low gain of 374 Subandirsquosand Elstonrsquos indices exhibited negative total gains

The base index of Williams yielded positive responses forall measured traits except for chlorophyll content flag leafarea and harvest index which showed negative gains Thisindex also achieved the highest grain yield response (627ie 2641 gmminus2) compared to the other indices employed inthis study The classic index of Smith and Hazel ranked sec-ond followed by the index of desired gains of Pesek and Bakerthe genotype-ideotype distance index proposed by Cruz andthe sum of ranks index of Mulamba andMock with selectionresponses of 3693 2746 2207 and 374 respectivelyThe single-trait responses varied from one index to anotherwith a more or less balanced distribution of positive andnegative estimates among the traits This shows that themonotrait selection was inadequate because it led to a higherfinal product when considering the grain yield and generatedunfavorable responses in other traits These results indicatedthat methods that combine favorable expected gains shouldbe used in the evaluation of these progeniesThe gains expect-ed through indices for grain yield per se were larger thanthose obtained by direct and indirect monotrait selectionexcept for the flag leaf area (Tables 5 and 6) Mahdy [49]mentioned that selection-based index was predicted to be

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 6: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

6 International Journal of Agronomy

exhibited the highest average for the number of spikes (5037spikes) For 1000-kernel weight Acsad1135 timesMahon-Demiaswas the best population with an average of 398 g

As the contrast ldquoParents versus F4rdquo was significant forPH SN TKW SW BIO and GY and compared with theleast significant difference (LSD005) significant differencesexisted between the parents and hybrids values for thesemea-sured traits These differences were 1 to 3 times higher thanthe LSD005 and were in favor of some F4 lines indicatingthat they perform better than the parents and suggestingthe possibilities of making significant changes through effec-tive selection These findings were in harmony with thoseobtained by Abd El-Shafi [34] Loffler and Busch [40] andAlexander et al [41]

32 Genetic Gain from Direct and Indirect Selection andSelection-Based Index The gains of direct and indirect selec-tion are shown inTable 5The results showed a big variation inthe percentages of gains among the measured traitsThe totalsum of gains per selection varied from minus3611 for headingdate to 3475 for flag leaf area Generally the direct selectionbased on chlorophyll content heading date spikes weightgrain yield and harvest index resulted in negative total gainsOn the other hand flag leaf area plant height number ofspikes 1000-kernel weight number of grains per spike andbiomass recorded positive total gains

The gains obtained by direct selection were higher thanthose of indirect selection But sometimes the indirect selec-tion may be more efficient especially if the secondary traitis highly correlated with yield and is easily measurable [42]The number of grains per spike (1586) followed by aboveground biomass (1230) plant height (1210) and numberof spikes (1084) exhibited the greatest gains from directselection Inversely heading date (minus113) spikes weight(minus499) and harvest index (minus381) showed negative gainvalues Negative gain for heading date is desired in the case ofthis study as the Algerian wheat breeding program designedfor semiarid regions precocity is a crucial criterion adoptedfor selection It is related to the ability of plants to shortentheir cycle so as to decrease their exposure to the late-seasonsirocco weathering

The direct selection for chlorophyll content resulted innegative indirect gains for nearly all the other traits exceptfor the flag leaf area number of grains per spike and harvestindex that showed positive responses For heading date theindirect selection gains were only positive for chlorophyllcontent number of spikes and harvest indexThe correlativeeffects of the selection based on the flag leaf area were desir-able for grain yield although the indirect gain for the remain-ing traits was practically negative

The direct selection for plant height resulted in positiveindirect gains for heading date number of spikes 1000-kernelweight and spikes weight In addition positive responsesof the selection based on yield and yield components wereobserved for grain yield itself heading date 1000-kernelweight and spikes weight However negative indirect gainswere exhibited for other traits including chlorophyll contentflag leaf area and number of grains per spike indicatingthat indirect selection for one variable for gain in another

is unfeasible because there will be a loss in the indirectlyselected variable In cases of negative gains the model is con-sidered inappropriate for selection in this plant material

The highest correlated responses for grain yield weregenerated though indirect selection on the base of theflag leaf area (804 ie 3387 gmminus2) followed by thespike fertility (365 ie 1539 gmminus2) 1000-kernel weight(363 ie 1531 gmminus2) and above ground biomass (274 ie1154 gmminus2)These results indicated that the indirect selectionwas to be more effective in improving the primary trait thanthe indirect selection based on other traits andor on thedirect selection based on the grain yield itself

Several authors have estimated the genetic gains of traitsinvolved in yield determination The results are often incon-sistent and scarce Our results were consistent with thoseof DePauw and Shebeski [43] and Inagaki et al [44] whomentioned that direct selection on the basis of yield is ineffec-tive in early generations Benmahammed et al [45] reportedthe same findings in barley crop Their results showed thatbiomass-based direct selection appearedmore discriminatingthan yield-based selection in their plant material Howeverthe results of this study do not corroborate findings reportedby Lalic et al [11] Mitchell et al [46] Lungu et al [47] andEl-Morshidy et al [48] and who observed the effectiveness ofthe direct selection on grain yieldThedifference in the resultsmay be attributed to differences of breeding material and togenotype times environments

The gains of selection-based index are shown in Table 6The total sum of gains from the selection-based index rangedfrom minus937 to 4286 Five out of seven used indicesshowed positive total gains The base index of Williamsrecorded the highest total gain (4286) followed by theclassic index of Smith and Hazel the index desired gains ofPesek and Baker (2746) and the genotype-ideotype dis-tance index of Cruz (2207) The sum of ranks index ofMulamba and Mock had a very low gain of 374 Subandirsquosand Elstonrsquos indices exhibited negative total gains

The base index of Williams yielded positive responses forall measured traits except for chlorophyll content flag leafarea and harvest index which showed negative gains Thisindex also achieved the highest grain yield response (627ie 2641 gmminus2) compared to the other indices employed inthis study The classic index of Smith and Hazel ranked sec-ond followed by the index of desired gains of Pesek and Bakerthe genotype-ideotype distance index proposed by Cruz andthe sum of ranks index of Mulamba andMock with selectionresponses of 3693 2746 2207 and 374 respectivelyThe single-trait responses varied from one index to anotherwith a more or less balanced distribution of positive andnegative estimates among the traits This shows that themonotrait selection was inadequate because it led to a higherfinal product when considering the grain yield and generatedunfavorable responses in other traits These results indicatedthat methods that combine favorable expected gains shouldbe used in the evaluation of these progeniesThe gains expect-ed through indices for grain yield per se were larger thanthose obtained by direct and indirect monotrait selectionexcept for the flag leaf area (Tables 5 and 6) Mahdy [49]mentioned that selection-based index was predicted to be

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 7: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

International Journal of Agronomy 7

Table5Selectiongain

estim

ates

(GS

)obtainedfro

mdirect(diagonal)andindirect(ofd

iagonal)selectionin

different

breadwheattraits

Traits

119883 119900119883 119904

GS(

)To

tal

Chl

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

CHL

437

464

615

(269)

minus007

(minus009)

160

(030)

minus325

(minus211)

minus406

(minus1862)

minus049

(minus019)

minus412

(minus2831

)214

(052

)minus714

(minus8330)

minus110

(minus463)

577

(209)

minus458

HD

1265

1250

143

(063)

minus113

(minus14

3)minus774

(minus14

4)minus4

69

(minus305)

176

(809)

minus235

(minus088)minus1098

(minus7543)

minus495

(minus12

1)minus1233

(minus14386)minus3

76(minus1584)

863

(313

)minus3

611

FLA

186

194

minus151

(minus066)

054

(068)

43 0

(080)

minus065

(minus042)minus4

72

(minus2169)

minus071

(minus027)

876

(6015)

1400

(343)

640

(7463)

804

(3387)

031

(011)

3475

PH64

9728

minus130

(minus057)

036

(045)

minus457

(minus085)

1210

(786)

933

(4285)

714

(268)

563

(3869)minus1579

(minus386)

1238

(14436)minus123

(minus517)minus110

8(minus401)

1298

SN4592

5089minus0

88

(minus038)

010

(013

)minus8

17(minus15

2)458

(297)

1084

(4976)

422

(158)

552

(379

0)minus1215

(minus298)

815

(9501)

114

(480)

minus437

(158)

897

TKW

375

413

minus070

(minus030)

096

(122)

minus477

(minus089)

387

(252

)1119

(513

8)993

(373)

1084

(7442)minus1446

(minus354)

1404

(16376

)363

(1531)minus741

(minus268)

2714

SW6868

6525

minus126

(minus055)

040

(050)

minus336

(minus062)minus0

17(minus011)

minus718

(minus3295)

359

(135)

minus499

(minus34

25)

minus013

(minus003)minus4

07

(minus4744)

minus505

(minus2127)minus150

(minus054)minus2

371

GN

245

284

205

(089)

053

(067)

385

(072)

minus584

(minus379)minus1050

( minus4820)

minus099

(minus037)

070

(480)

1586

(388)

minus414

(minus4830)

365

(1539)

687

(249)

1204

BIO

11664

13099

minus122

(minus053)

063

(080)

minus358

(minus067)

793

(515

)74

4(3418)

615

(231

)73

6(5055)

minus894

(minus219)

1230

(143

50)

274

(115

4)minus744

(minus269)

2339

GY

4214

4222minus2

07

(minus090)

069

(087)

minus148

(minus027)minus0

95

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

HI

3 62

348

minus091

(minus040

)045

(057)

030

(006)

minus208

(minus13

5)minus5

74(minus2635

)272

(102)

060

(409)

minus080

(minus019)

014

(163)

minus354

(minus1490)minus3

81

(minus13

8)minus1267

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

pers

pikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GY

grainyield(gmminus2)HIharvestind

ex(

)1198830traitmeanvalueof

thebase

popu

latio

nand119883119904traitm

eanvalueof

theselected

popu

lationsvaluesb

etween

parenthesesc

orrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ield

andyield

-related

traits

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 8: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

8 International Journal of Agronomy

Table6Selectiongain

estim

ates

(GS

)obtainedfro

mselection-basedindexin

different

breadwheattraits

Indices

GS(

)To

tal

CHL

HD

FLA

PHSN

TKW

SWGN

BIO

GY

HI

Smith

ampHazel

195

(085)

097

(123)

minus093

(minus017)

874

(568)

428

(1965)

953

(357)

1020

(7002)

minus804

(minus19

7)1360

(15863)

394

(1659)

minus730

(minus264

)3693

Mulam

baampMock

minus271

(minus118)

003

(004)

minus002

(minus000)

405

(263)

minus160

(minus735)

208

(078)

minus235

(minus1611)

150

(037

)minus0

58

(minus677)

156

(657)

178

(065)

374

Williams

minus043

(minus019)

156

(198)

minus037

(minus007)

571

(371)

085

(391)

844

(317

)1243

(8535)

000

(000)

1478

(17243)

627

(2641)

minus640

(minus232)

4286

Suband

iminus137

(minus060)

078

(099)

006

(001)

327

(212

)minus9

31(minus4275)

462

(173)

minus318

(minus2185)

253

(062)

minus110

(minus1284)

minus308

(minus1299)

minus258

(minus093)

minus937

Elsto

nminus2

07

(minus090)

069

(087)

minus148

(minus027)

minus095

(minus062)minus6

49

(minus2982)

340

(127)

minus138

(minus94

5)437

(107)

minus247

(minus2884)

020

(085)

210

(076

)minus4

08

PesekampBa

ker

223

(097)

078

(099)

438

(081)

minus511

(minus332)

033

(151)

588

(221)

872

(598

9)minus0

31(minus008)

791

(9223)

542

(2283)

minus276

(minus10

0)2746

Cruz

minus146

(minus064

)095

(120)

224

(042)

015

(010

)minus799

(minus3669)

382

(143)

363

(2495)

1049

(257)

159

(1850)

576

(2427)

288

(104)

2207

CHLchloroph

yllcon

tent

(Spad)H

Dheading

date(days)FLA

flag

leafarea

(cm2)PH

plant

height

(cm)SN

num

bero

fspikesTK

W100

0-kernelweight(g)SWspikesw

eight(gmminus2)GNnum

bero

fgrains

perspikeB

IOabo

vegrou

ndbiom

ass(gmminus2)GYgrainyield(gmminus2)andHIharvestind

ex(

)andvaluesbetweenparenthesescorrespon

dto

theg

eneticgain

interm

sofp

ercent

meanfory

ieldandyield-related

traits

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 9: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

International Journal of Agronomy 9

Table 7 Differences of the F4 selected lines relatively to the standard check Hidhab for the measured traits

Traits F4 selected lines mean Hidhab Difference Difference in of Hidhab LSD005Chl 4226 4330 minus105 minus241 480HD 12657 12967 minus310lowast minus239 140FLA 1915 1940 minus024 minus126 631PH 6867 6233 633 1016 667SN 61567 46333 15233lowast 3288 11967TKW 3997 3833 163 426 453SW 114767 82333 32433lowast 3939 23517GN 3157 2483 675lowast 2718 675BIO 183367 137667 45700lowast 3320 35315GY 75813 44133 31680lowast 7178 18510HI 4178 3223 955 2963 994CHL chlorophyll content (Spad) HD heading date (days) FLA flag leaf area (cm2) PH plant height (cm) SN number of spikes TKW 1000-kernel weight(g) SW spikes weight (gmminus2) GN number of grains per spike BIO above ground biomass (gmminus2) GY grain yield (gmminus2) and HI harvest index ()lowastSignificant effect at 005 probability

superior for yield improvement as compared to themonotraitselection method

The first three axes of the Principal Component Analysis(PCA) explainedmore than 7793of the total variation avail-able in the data subjected to analysis Heading date (0386lowast)flag leaf area (0308lowast) spikes weight (0923lowast) biomass(0853lowast) and grain yield (0866lowast) correlated significantly toPCA1 PCA2 was mainly related to the chlorophyll content(0276lowast) 1000-kernel weight (0440lowast) number of grains perspike (0655lowast) and harvest index (0547lowast) (Figure 1) Thenumber of spikes (0333lowast) was related to PCA3 (Figure 2)ThePCA biplots showed that the populations Acsad899 timesHidhab(P7) Acsad899 times El-Wifak (P8) Acsad1135 times Hidhab (P11)Acsad1135 times El-Wifak (P12) Acsad1069 times Rmada (P14) AinAbid timesMahon-Demias (P17) Ain Abid times Rmada (P18) andAinAbidtimesEl-Wifak (P20) werewell represented on the planeformed by the first axis PCA1 (Figure 1)

Ain Abid times Mahon-Demias (P17) Ain Abid times Rmada(P18) and Ain Abid times El-Wifak (P20) had positive coordi-nates with this axis They were characterized by a long vege-tative cycle and high biomass spikes weight and grain yieldvaluesThe last two populations Ain Abid timesRmada (P18) andAin Abid times El-Wifak (P20) were also distinguished by highvalues of chlorophyll content flag leaf area spikes fertilityand harvest index relatively to PCA2 but were associatedwith low number of ears relatively to PCA3 (Figures 1 and2) On the other hand Ain Abid timesMahon-Demias (P17) wasdistinguished relatively to PCA2 by high estimates of the1000-kernel weight and plant height and associatedwith highnumber of spikes relatively to PCA3 (Figures 1 and 2) Fromthese results it could be concluded that effective selectionof superior individuals within this population certainly con-tributes to the improvement of yield and yield componentsin a semi-late genetic background

33 Selection of Superior Genotypes for Grain Yield Geno-types were first ranked according to grain yield then the 5highest yielding lines were selected and theirmean yield esti-matedThe 30 lines thus selected are derived from 15 out of 20F4 populations studied Half part of these lines was equitably

P1

P2

P3

P4

P5P6

P7P8

P9

P10

P11

P12

P13

P14

P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH SN

TKW

SW

GN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4minus3minus2minus1

012345

PCA

2 (2

429

)

Figure 1 Principal Component Analysis (PCA) projections on axes1 and 2 accounting for 6356 of total variation for 20 F4 popula-tions of bread wheat

derived from Acsad1135 times Hidhab (P11) Ain Abid times Hidhab(P19) and Ain Abid times El-Wifak (P20) crosses The popula-tion Ain Abid times Mahon-Demias (P17) previously identifiedamong the promising populations contributed with fourlines Acsad1069 times Rmada (P14) and Ain Abid times Rmada (P18)participated with two lines each Seven other populationscontributed by one line each Relative to the standard checkHidhab which is the most cultivated variety in Algeriathe top 30 lines selected were characterized by significantimprovements in yield components including the numberof spikes (15233 spikes mminus2) spikes weight (32433 gmminus2)spikes fertility (675 grains spikeminus1) above ground biomass(45700 gmminus2) and grain yield (31680 gmminus2) (Table 7)Besides they were distinguished by significant reduction ofthe duration in the vegetative phase with 310 days (Table 7)

4 Conclusion

The results of this study indicated appreciable genetic varia-bility among the evaluated populations Selection based-indexwas more efficient to improve grain yield compared to direct

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 10: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

10 International Journal of Agronomy

P1

P2

P3

P4

P5

P6

P7

P8

P9

P10

P11P12

P13

P14 P15

P16

P17

P18

P19

P20

CHL

HD

FLA

PH

NE

TKW

SWGN

BIO

GY

HI

minus5 minus4 minus3 minus2 minus1 0 1 2 3 4 5 6minus6PCA1 (3927)

minus4

minus3

minus2

minus1

0

1

2

3

4

5

PCA

3 (1

438

)

Figure 2 Principal Component Analysis (PCA) projections on axes1 and 3 accounting for 5365 of total variation for 20 F4 popula-tions of bread wheat

and indirect single-trait selection The analytical proceduresof the different selection methods showed possibilities ofapplications in advanced generations of breeding beingsuperior when compared with direct and indirect selectionWilliamsrsquos index was predicted to be more effective than theother selection indices for improving multiple traits at timeIt brought the highest total genetic gain and the best yieldgain per se associated with positive correlated responses formost of yield components Compared to the check cultivarHidhab the 30 F4 selected lines at 5 selection intensity werecharacterized by significant increase in grain yield and yieldrelated traits

Conflicts of Interest

The authors declare that there are no conflicts of interest re-garding the publication of this article

References

[1] ABenbelkacem ldquoAdaptation of cereal cultivars to extreme agro-ecologic environments of North Africardquo Field Crops Researchvol 45 no 1-3 pp 49ndash55 1996

[2] A Mekhlouf F Dehbi H Bouzerzour A Hannchi A Benma-hammed and A Adjabi ldquoRelationships between cold tolerancegrain yield performance and stability of durumwheat (TriticumdurumDesf) genotypes grown at high elevation area of EasternAlgeriardquo Asian Journal of Plant Sciences vol 5 no 4 pp 700ndash708 2006

[3] G A Slafer and FH Andrade ldquoChanges in physiological attrib-utes of the dry matter economy of bread wheat (Triticumaestivum) through genetic improvement of grain yield potentialat different regions of the world - A reviewrdquo Euphytica vol 58no 1 pp 37ndash49 1991

[4] S P Loss and K H M Siddique ldquoMorphological and Phys-iological Traits Associated with Wheat Yield Increases inMediterranean Environmentsrdquo Advances in Agronomy vol 52no C pp 229ndash276 1994

[5] RM TrethowanMVanGinkel K Ammar et al ldquoAssociationsamong twenty years of international bread wheat yield evalua-tion environmentsrdquo Crop Science vol 43 no 5 pp 1698ndash17112003

[6] H Bouzerzour A Djekoun A Benmahammed and K LHassous ldquoContribution de la biomasse aerienne de lrsquoindicede recolte et de la precocite a lrsquoepiaison au rendement grainde lrsquoorge (Hordeum vulgare L) en zone drsquoaltituderdquo Cahiers delrsquoAgriculture vol 8 pp 133ndash137 1998

[7] A Benmahammed H Nouar L Haddad Z Laala A Oulmiand H Bouzerzour ldquoAnalyse de la stabilite des performances derendement du ble dur (Triticum durum Desf) sous conditionssemi-aridesrdquo Biotechnologie Agronomie Societe et Environ-nement vol 14 no 1 pp 177ndash186 2010

[8] B RWhan R Knight and A J Rathjen ldquoResponse to selectionfor grain yield and harvest index in F2 F3 and F4 derived linesof two wheat crossesrdquo Euphytica vol 31 no 1 pp 139ndash150 1982

[9] M P Reynolds R P Singh A Ibrahim O A A Ageeb ALarque-Saavedra and J S Quick ldquoEvaluating physiologicaltraits to complement empirical selection for wheat in warmenvironmentsrdquo Euphytica vol 100 no 1-3 pp 85ndash94 1998

[10] M Balota A J Green C A Griffey R Pitman andWThoma-son ldquoGenetic gains for physiological traits associated with yieldin soft red winter wheat in the Eastern United States from 1919to 2009rdquo European Journal of Agronomy vol 84 pp 76ndash83 2017

[11] A Lalic D Novoselovic J Kovacevic et al ldquoGenetic gain andselection criteria effects on yield and yield components in barley(Hordeum vulgare L)rdquo Periodicum biologorum vol 112 no 3pp 311ndash316 2010

[12] MA BabarMVanGinkelM P Reynolds B Prasad andA RKlatt ldquoHeritability correlated response and indirect selectioninvolving spectral reflectance indices and grain yield in wheatrdquoAustralian Journal of Agricultural Research vol 58 no 5 pp432ndash442 2007

[13] MMCosta AODiMauro S HUneda-Trevisoli et al ldquoAnal-ysis of direct and indirect selection and indices in soybean seg-regating populationsrdquoCrop Breeding and Applied Biotechnologyvol 8 no 1 pp 47ndash55 2008

[14] J M S Viana V R Faria F F e Silva and M D V de ResendeldquoCombined selection of progeny in crop breeding using bestlinear unbiased predictionrdquo Canadian Journal of Plant Sciencevol 92 no 3 pp 553ndash562 2012

[15] C Y Lin ldquoIndex selection for genetic improvement of quanti-tative charactersrdquo Theoretical and Applied Genetics vol 52 no2 pp 49ndash56 1978

[16] J D Smith and M L Kinman ldquoThe Use of Parent-OffspringRegression as an Estimator of Heritability1rdquo Crop Science vol5 no 6 p 595 1965

[17] K J Frey and T Horner ldquoHeritability in standard unitsrdquo Agro-nomy Journal vol 49 no 2 pp 59ndash62 1957

[18] Z Fellahi A Hannachi H Bouzerzour and A BoutekrabtldquoLine times Tester Mating Design Analysis for Grain Yield andYield Related Traits in Bread Wheat (Triticum aestivumL)rdquoInternational Journal of Agronomy vol 2013 pp 1ndash9 2013

[19] Z Fellahi A Hannachi H Bouzerzour S Dreisigacker AYahyaoui and D Sehgal ldquoGenetic analysis of morpho-physio-logical traits and yield components in F2 partial diallel crossesof bread wheat (Triticum aestivum L)rdquo Revista FacultadNacional de Agronomia vol 70 no 3 pp 8237ndash8250 2017

[20] H Chennafi A Aıdaoui H Bouzerzour and A Saci ldquoYieldresponse of durumwheat (Triticum durumDesf) cultivarWahato deficit irrigation under semi arid growth conditionsrdquo AsianJournal of Plant Sciences vol 5 no 5 pp 854ndash860 2006

[21] P L Spagnoletti Zeuli and C O Qualset ldquoFlag Leaf Variationand the Analysis of Diversity in DurumWheatrdquo Plant Breedingvol 105 no 3 pp 189ndash202 1990

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 11: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

International Journal of Agronomy 11

[22] M SMcIntosh ldquoAnalysis of combined experimentsrdquoAgronomyJournal vol 75 no 1 pp 153ndash155 1983

[23] R G D Steel and J H Torrie Principles and procedures ofstatistics McGraw-Hill Books New York NY USA 1982

[24] GAcquaahPrinciples of plant genetics and breeding JohnWileyand Sons 2009

[25] C D Cruz Programa GENES Biometria Editora UFV VicosaBrazil 1st edition 2006

[26] H F Smith ldquoA discriminates function for plant selectionrdquo An-nals of Eugenics vol 7 pp 240ndash250 1936

[27] L N Hazel ldquoThe genetic basis for constructing selection in-dexesrdquo Genetics vol 28 no 6 pp 476ndash490 1943

[28] J S Williams ldquoThe evaluation of a selection indexrdquo Biometricsvol 18 no 3 pp 375ndash393 1962

[29] R C Elston ldquoA Weight-Free Index for the Purpose of Rankingor SelectionwithRespect to Several Traits at a TimerdquoBiometricsvol 19 no 1 p 85 1963

[30] J Pesek and R J Baker ldquoComparison of predicted and observedresponses to selection for yield in wheatrdquo Canadian Journal ofPlant Science vol 51 no 3 pp 187ndash192 1971

[31] W Subandi A Compton and L T Empig ldquoComparison of theefficiencies of selection indices for three traits in two varietycrosses of cornrdquo Crop Science vol 13 no 2 pp 184ndash186 1973

[32] N N Mulamba and J J Mock ldquoImprovement of yield potentialof the ETO blanco maize (Zea mays L) population by breedingfor plant traitsrdquoThe Egyptian Journal of Genetics and Cytology vol 7 no 1 pp 40ndash51 1978

[33] C D Cruz ldquoGENES - A software package for analysis in exper-imental statistics and quantitative geneticsrdquo Acta Scientiarum -Agronomy vol 35 no 3 pp 271ndash276 2013

[34] M A Abd El-Shafi ldquoEstimates of Genetic Variability and Effi-ciency of Selection for Grain Yield and Its Components in TwoWheat Crosses (Triticum aestivum L)rdquo nternational Journal ofAgriculture and Crop Sciences vol 7 no 2 pp 83ndash90 2014

[35] R W Allard Principles of Plant Breeding John Wily and SonsInc New York NY USA 2nd edition 1960

[36] C D Cruz A J Regazi and P C S Carneiro Modelos bio-metricos aplicados ao melhoramento genetico UFV Vicosa Bra-zil 4th edition 2012

[37] A Mesele W Mohammed and T Dessalegn ldquoEstimation ofHeritability and Genetic Advance of Yield and Yield RelatedTraits in Bread Wheat (Triticum aestivum L) Genotypes atOfla District Northern Ethiopiardquo International Journal of PlantBreeding and Genetics vol 10 no 1 pp 31ndash37 2015

[38] B Saleem A KhanM Shahzad and F Ijaz ldquoEstimation of her-itability and genetic advance for various metric traits in sevenF2 populations of bread wheat (Triticum aestivum L)rdquo Journalof Agricultural Sciences Belgrade vol 61 no 1 pp 1ndash9 2016

[39] M Yaqoob ldquoEstimation of genetic variability heritability andgenetic advance for yield and yield related traits in wheat underrainfed conditionsrdquo Journal of Agricultural Research vol 54 no1 pp 1ndash14 2016

[40] C M Loffler and R H Busch ldquoSelection for Grain ProteinGrain Yield and Nitrogen Partitioning Efficiency in Hard RedSpring Wheatrdquo Crop Science vol 22 no 3 p 591 1982

[41] W L Alexander E L Smith and C Dhanasobhan ldquoA compar-ison of yield and yield component selection in winter wheatrdquoEuphytica vol 33 no 3 pp 953ndash961 1984

[42] J Kumar and P N Bahl ldquoDirect and indirect selection for yieldin chickpeardquo Euphytica vol 60 no 3 pp 197ndash199 1992

[43] R M DePauw and L H Shebeski ldquoAn evaluation of an earlygeneration yield testing procedure in Triticum aestivumrdquo Cana-dian Journal of Plant Science vol 53 no 3 pp 465ndash470 1973

[44] M N Inagaki G Varughese S RajaramM Van Ginkel and AMujeeb-Kazi ldquoComparison of bread wheat lines selected bydoubled haploid single-seed descent and pedigree selectionmethodsrdquo Theoretical and Applied Genetics vol 97 no 4 pp550ndash556 1998

[45] A Benmahammed H Bouzerzour A Djekou and K HassousldquoEfficacite de la selection precoce de la biomasse chez lrsquoorge(Hordeum vulgare L) en zone semi-ariderdquo Sciences and Tech-nologie vol C no 22 pp 80ndash85 2004

[46] J W Mitchell R Baker and D R Knott ldquoEvaluation of Honey-comb Selection for Single Plant Yield in Durum Wheatrdquo CropScience vol 22 no 4 pp 840ndash843 1982

[47] DM Lungu P J Kaltsikes and EN Larter ldquoHoneycomb selec-tion for yield in early generations of spring wheatrdquo Euphyticavol 36 no 3 pp 831ndash839 1987

[48] M A El-Morshidy K A Kheiralla M A Ali and A A S Ah-med ldquoEfficiency of pedigree selection for earliness and grainyield in two wheat populations under water stress conditionsrdquoAssiut Journal of Agricultural Sciences vol 37 pp 77ndash94 2010

[49] E EMahdy ldquoSingle andMultiple Traits Selection in a Segregat-ing Population of Wheat Triticum aestivum Lrdquo Plant Breedingvol 101 no 3 pp 245ndash249 1988

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom

Page 12: Analysis of Direct and Indirect Selection and Indices …downloads.hindawi.com/journals/ija/2018/8312857.pdflowCV/CV ratiovalues,indicatingthedominanteectof theenvironmentoncrop. Generally,

Nutrition and Metabolism

Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

Food ScienceInternational Journal of

Hindawiwwwhindawicom Volume 2018

International Journal of

Microbiology

Hindawi Publishing Corporation httpwwwhindawicom Volume 2013Hindawiwwwhindawicom

The Scientific World Journal

Volume 2018

Hindawiwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2018

AgricultureAdvances in

Hindawiwwwhindawicom Volume 2018

PsycheHindawiwwwhindawicom Volume 2018

BiodiversityInternational Journal of

Hindawiwwwhindawicom Volume 2018

ScienticaHindawiwwwhindawicom Volume 2018

GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Plant GenomicsInternational Journal of

Hindawiwwwhindawicom Volume 2018

Biotechnology Research International

Hindawiwwwhindawicom Volume 2018

Forestry ResearchInternational Journal of

Hindawiwwwhindawicom Volume 2018

BotanyJournal of

Hindawiwwwhindawicom Volume 2018

EcologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Veterinary Medicine International

Hindawiwwwhindawicom Volume 2018

Cell BiologyInternational Journal of

Hindawiwwwhindawicom Volume 2018

Hindawiwwwhindawicom Volume 2018

BioMed Research International

Agronomy

Hindawiwwwhindawicom Volume 2018

International Journal of

Submit your manuscripts atwwwhindawicom


Recommended