Mapping QTLs for salt tolerance with additive,epistatic and QTL × treatment interaction effects at seedling stage in wheat

Quantitative trait loci (QTLs) for salt tolerance with additive, epistatic and QTL × treatment interaction effects at seedling stage in wheat were identified. A set of 131 recombinant inbred lines derived from cross Chuan 35050 × Shannong 483 were evaluated under salt stress and normal conditions. Wide variation was found for all studied traits. A total of 18 additive and 16 epistatic QTLs were detected, among which five and 11 were with significant QTL × treatment effects. Ten QTL clusters were identified, and each may represent a single gene or closely linked genes. The locus controlling shoot K⁺/Na⁺ concentration ratio and shoot Na⁺ concentration on chromosome 5A may be identical to Nax2. The interval Xgwm6‐Xgwm538 on chromosome 4B for total dry weight was also identified in a previous study, both near the marker Xgwm6. The marker Xgwm6 may be useful for marker‐assisted selection. Six pairs of homoeologous QTLs were detected, showing synteny among the A, B and D genomes. These results facilitate understanding the mechanisms and the genetic basis of salt tolerance in wheat.     

Identification of microsatellite markers linked with yield components under drought stress at terminal growth stages in durum wheat

Grain yield and yield components are the main important traits involved in durum wheat (Triticum turgidum L.) improvement programs. The purpose of this research was to identify quantitative trait loci (QTL) associated with yield components such as 1000 grain weight (TGW), grain weight per spike (GWS), number of grains per spike (GNS), spike number per m² (SN), spike weight (SW), spike harvest index (SHI) and harvest index (HI) using microsatellite markers. Populations of F₃ and F₄ lines derived from 151 F₂ individuals developed from a cross between Oste-Gata, a drought tolerant, and Massara-1, a drought susceptible durum wheat genotypes, were used. The populations were evaluated under four environmental conditions including two irrigation regimes of drought stress at terminal growth stages and normal field conditions in two growing seasons. Two hundred microsatellite markers reported for A and B genomes of bread wheat were used for parental polymorphism analysis and 30 polymorphic markers were applied to genotype 151 F_2:3 families. QTL analysis was performed using genome-wide single marker regression analysis (SMA) and composite interval mapping (CIM). The results of SMA revealed that about 20% of the phenotypic variation of harvest index and TGW could be explained by Xcfd22-7B and Xcfa2114-6A markers in different environmental conditions. Similarly, Xgwm181-3B, Xwmc405-7B and Xgwm148-3B and marker Xwmc166-7B were found to be associated with SHI and GWS, respectively. A total of 20 minor and major QTL were detected; five for TGW, two for GWS, two for GNS, three for SN, five for HI, two for SHI and one for SW. The mapped QTL associated with ten markers. Moreover, some of these QTL were prominent and stable under drought stress and non drought stress environments and explained up to 49.5% of the phenotypic variation.     

Expression of barley yellow dwarf virus and Russian wheat aphid resistance genes in and fertility of spring wheat × triticale hybrids and backcross lines

Summary The Barley Yellow Dwarf Virus disease (BYDV) and the Russian wheat aphid (RWA) Diuraphis noxia (Mordvilko) have caused significant losses to wheat and barley in several areas of the world. Important sources of resistance to both BYDV and RWA have been found in Triticale. Different generations of interspecific wheat x Triticale crosses were produced and the progenies were screened for BYDV and RWA tolerance. Plants with equal chromosome numbers showed different levels of fertility. A significant correlation was observed between pollen fertility and seed set in primary florets (r=0.57). In generaL, pollen fertility, seed set and the number of euploid plants (2n=42) increased from one generation to the next. The expression of BYDV tolerance varied from population to population. Additive effects were predominant in F₁ and some backcross populations. A dominant effect of rye tolerance genes was also observed in few populations. A monogenic trait or a quantitative (polygenic) character would not agree with the observed segregation patterns. The heritability of this oligogenic tolerance was quite different between populations and in many populations the tolerance genes were only partially expressed. Some transgressive segregation for tolerance and sensitivity was demonstrated. The genes controlling tolerance to RWA in Triticale lines, Muskox 658 and Nord Kivu were not expressed in advanced lines resistant to BYDV. This indicates that tolerance genes for BYDV and RWA in these lines are located on different chromosomes.     

Genetic analysis of ratooning ability of rice (Oryza sativa L.)

Summary Genetic analysis of generation means of F₁, F₂, F₃, F₄, and the parental populations of the cross IR10154-23-3-3A/IR15795-232-3-3-2 for ratooning ability was conducted. Good ratooning ability is a recessive trait. The parents differed by at least two pairs of major genes for ratooning ability. A large proportion of transgressive segregants in F₂ suggested that the expression of ratooning ability is influenced not only by major genes but also by modifiers. Broad-sense heritability estimates computed by different methods ranged from 0.66 to 0.88. Narrow-sense heritability by the F₄–F₃ regression method was 0.39. The heritability estimates were 0.42 and 0.33 by the variance component method. The prevalence of additive x additive type of gene effects along with prominent additive effects imply some scope for selection in the segregating generations. However, non-additive type of gene action also affects the expression of ratooning ability.     

Cytological and microsatellite mapping of mutant genes for spherical grain and compact spikes in durum wheat

Two mutants for sphaerococcoid seed (MA 16219) and compact spike (MA 17648) were isolated from M₃ progeny of durum wheat cultivar, Altaiskaya Niva, mutagenized with chemical mutagens. The chromosomal locations of the genes involved were determined by the use of a complete set of D-genome disomic substitutions in durum cultivar, Langdon. The gene for sphaerococcoid grain, s ¹⁶²¹⁹ , was allelic to S2, located in the centromeric region of chromosome 3B in hexaploid wheat. The gene for compact spike, C ¹⁷⁶⁴⁸ , was located on chromosome 5AL distal to the Q locus. Using microsatellite markers, C ¹⁷⁶⁴⁸ and awn inhibitor B1 were located in the F₂ of LD222 × MA17648. The gene order was Xbarc319—C ¹⁷⁶⁴⁸ —Xgwm179—Xgwm126—Xgwm291—B1.     

Inheritance of root traits and phosphorus uptake in common bean (Phaseolus vulgaris L.) under limited soil phosphorus supply

Heterosis is an important way to improve yield and quality for many crops. Hybrid rice and hybrid maize contributed to enhanced productivity which is essential to supply enough food for the increasing world population. The success of hybrid rice in China has led to a continuous interest in hybrid wheat, even when most research on hybrid wheat has been discontinued in other countries for various reasons including low heterosis and high seed production costs. The Timopheevii cytoplasmic male sterile system is ideal for producing hybrid wheat seeds when fertility restoration lines with strong fertility restoration ability are available. To develop PCR-based molecular markers for use in marker-assisted selection of fertility restorer lines, two F₂ populations derived from crosses R18/ND36 and R9034/ND36 were used to map fertility restoration genes in the two elite fertility restorer lines (R-lines) R18 and R9034. Over 678 SSR markers were analyzed, and markers closely linked to fertility restoration genes were identified. Using SSR markers, a major fertility restoration gene, Rf3, was located on the 1B chromosome in both populations. This gene was partially dominant in conferring fertility restoration in the two restorer lines. SSR markers Xbarc207, Xgwm131, and Xbarc61 are close to this gene. These markers may be useful in marker-assisted selection of new restorer lines with T. timopheevii cytoplasm. Two minor QTL conferring fertility restoration were also identified on chromosomes 5A (in R18) and 7D (in R9034) in two R-lines.     

Validation of a major QTL for scab resistance with SSR markers and use of marker-assisted selection in wheat

The objectives of this study were to validate the major quantitative trait locus (QTL) for scab resistance on the short arm of chromosome 3B in bread wheat and to isolate near‐isogenic lines for this QTL using marker‐assisted selection (MAS). Two resistant by susceptible populations, both using ‘Ning7840’ as the source of resistance, were developed to examine the effect of the 3BS QTL in different genetic backgrounds. Data for scab resistance and simple sequence repeat (SSR) markers linked to the resistance QTL were analyzed in the F_2:3 lines of one population and in the F_3:4 lines of the other. Markers linked to the major QTL on chromosome 3BS in the original mapping population (‘Ning7840’/‘Clark’) were closely associated with scab resistance in both validation populations. Marker‐assisted selection for the QTL with the SSR markers combined with phenotypic selection was more effective than selection based solely on phenotypic evaluation in early generations. Marker‐assisted selection of the major QTL during the seedling stage plus phenotypic selection after flowering effectively identified scab resistant lines in this experiment. Near‐isogenic lines for this 3BS QTL were isolated from the F₆ generation of the cross ‘Ning7840’/‘IL89‐7978’ based on two flanking SSR markers, Xgwm389 and Xbarc147. Based on these results, MAS for the major scab resistance QTL can improve selection efficiency and may facilitate stacking of scab resistance genes from different sources.     

Gene action and heritability for photosynthetic activity in two wheat crosses

Summary Gene action and heritability for photosynthetic activity were estimated from generation means in two wheat crosses during two stages (5 ^th leaf and flag leaf between 2 and 5 days after anthesis). Six generations were available for each cross: parents (P₁ and P₂), F₁, F₂ and backcrosses (BC₁ and BC₂).Correlations between some morphophysiological characters and photosynthetic activity of the flag leaf was also determined. The joint scaling test described by Mather & Jinks was used to determine the gene action. It showed that them; [d]; [h]; [i], [l] (mean, additivity, dominance, additive x additive interallelic interaction effects, dominance x dominance interallelic interaction effects) model fits the two crosses at both measurement times. All the model genetic components were significant for the flag leaf, however for the 5 ^th leaf only [h]; [i] and [l] were significant. The presence of additive and additive x additive effects suggested the possibility of selecting for this character using the flag leaf so as to obtain pure inbred lines. Dominance effects [h] were negative and dominance x dominance effects [l] were positive. Broad sense heritability values were medium to low. There were no correlations between the studied morphophysiological characters and the photosynthetic activity.     

Microsatellite Marker Identification of a Triticum Aestivum -Aegilops Umbellulata Substitution Line with Powdery Mildew Resistance

Summary Aegilops umbellulata acc. Y39 and Triticum carthlicum acc. PS5, immune to many powdery mildew isolates, were crossed to make an amphidiploid line Am9. The powdery mildew resistance of Am9 was transferred to common wheat cultivar Laizhou953 by crossing and backcrossing. In this study, the origin of powdery mildew resistance in a BC₃F_4:5 population derived from a cross of Am9 and Laizhou953 was identified. Microsatellite markers analysis showed that markers Xgwm257, Xgwm296, and Xgwm319, co-segregated with the powdery mildew resistance, whereas markers Xgwm210, Xgwm388/140, Xgwm388/170 and Xgwm526 were related to susceptibility and linked to resistance in repulsion. Of three markers related to resistance, Xgwm257 and Xgwm319 were codominant, whereas Xgwm296 was dominant. All three markers were Ae. umbellulata-specific indicating that resistance in the test population originated from Ae. umbellulata acc. Y39. The chromosome location and mapping of these linked microsatellite markers, the chromosome numbers of derived BC₃F_4:6 families, and chromosome pairing in F₁ plants from a cross of a homozygous resistant BC₃F_4:5 plant and Laizhou953, showed that wheat chromosome 2B was substituted by Ae. umbellulata chromosome 2U. This is the first gene conferring powdery mildew resistance transferred to wheat from Ae. umbellulata, and it should be a novel resistance gene to powdery mildew. It was temporarily designated PmY39.The first two authors made equal contributions     

QTL analysis of resistance to Fusarium head blight in wheat using a 'Wangshuibai'-derived population

Fusarium head blight (FHB) is a devastating disease that reduces the yield, quality and economic value of wheat. For quantitative trait loci (QTL) analysis of resistance to FHB, F₃ plants and F_3:5 lines, derived from a ‘Wangshuibai’ (resistant)/‘Seri82’(susceptible) cross, were spray inoculated during 2001 and 2002, respectively. Artificial inoculation was carried out under field conditions. Of 420 markers, 258 amplified fragment length polymorphism and 39 simple sequence repeat (SSR) markers were mapped and yielded 44 linkage groups covering a total genetic distance of 2554 cM. QTL analysis was based on the constructed linkage map and area under the disease progress curve. The analyses revealed a QTL in the map interval Xgwm533‐Xs18/m12 on chromosome 3BS accounting for up to 17% of the phenotypic variation. In addition, a QTL was detected in the map interval Xgwm539‐Xs15/m24 on chromosome 2DL explaining up to 11% of the phenotypic variation. The QTL alleles originated from ‘Wangshuibai’ and were tagged with SSR markers. Using these SSR markers would facilitate marker‐assisted selection to improve FHB resistance in wheat.     

Mapping antixenosis genes on chromosome 6A of wheat to greenbug and to a new biotype of Russian wheat aphid

Greenbug and Russian wheat aphid (RWA) are two devastating pests of wheat. The first has a long history of new biotype emergence and recently. RWA resistance has just started to break down. Thus, it is necessary to find new sources of resistance that will broaden the genetic base against these pests in wheat. Seventy‐five doubled haploid recombinant (DHR) lines for chromosome 6A from the F₁ of the cross between “Chinese Spring’ and the “Chinese Spring (Synthetic 6A) (Triticum dicoccoides × Aegilops tauschii)” substitution line were used as a mapping population for testing resistance to greenbug biotype C and to a new strain of RWA that appeared in Argentina in 2003. A quantitative trait locus (QTL) (br antixenosis to greenbug was significantly associated with the marker loci Xgwm1009 and Xgwm1185 located in the centromere region of chromosome 6A. Another QTL which accounted for most of the antixenosis against RWA was associated with the marker loci Xgwm1291 and Xiinni1150. both located on the long arm of chromosome 6A. This is the first report of greenbug and RWA resistance genes located on chromosome 6A. It is also the first report of antixenosis against the new strain of RWA. As most of the RWA resistance genes present in released cultivars have been located in [he D‐ genome, it is highly desirable to find new sources in other genomes to combine the existing resistance genes with new sources.     

Combining ability analysis of resistance to Helminthosporium leaf blight in spring wheat

Wheat breeders in South Asia are attempting to develop wheat (Triticum aestivum L.) cultivars resistant to Helminthosporium leaf blight (HLB), which occurs mainly as a complex of spot blotch caused by Cochliobolus sativus (Ito & Kuribayashi) Drechs. ex Dastur, and tan spot caused by Pyrenophora tritici-repentis (Died.) Drechs. Information on the combining ability for HLB resistance in wheat cultivars of South Asia is not available. This study was undertaken to examine the resistance to HLB in nine genetically diverse wheat parents, and to evaluate their general combining ability (GCA) and specific combining ability (SCA) effects toward determining the genetic basis of disease resistance. Nine parents were crossed in a half-diallel mating design to produce 36 populations. The F₁ and F₂ progenies, and the parents were evaluated in replicated field tests at Rampur, Nepal. Multiple disease scores were recorded, and area under the disease progress curve (AUDPC) was calculated to measure disease severity over time. The combining ability analysis was performed using Griffing's Method 2, Model 1. The parents chosen showed wide variation for resistance to HLB. They and the F₁ and F₂ progenies differed significantly for AUDPC. GCA and SCA effects were significant in both generations suggesting that additive as well as non-additive genetic mechanisms were involved in the expression of resistance in these parents. Wheat genotypes 'SW89-5422', 'G 162', 'NL 781'and 'Chirya 7' had significantly negative GCA effects for AUDPC in both F₁ and F₂ generations, suggesting their prime suitability for use in wheat breeding programs to improve resistance to HLB. The estimate of narrow-sense heritability was 0.77 in both generations suggesting that selection for HLB resistance should be effective in these crosses. The results indicate a predominance of additive gene action in the inheritance of HLB resistance in spring wheat.     

Inheritance of β-carotene concentration in durum wheat (Triticum turgidum L. ssp. durum)

Despite being one of the important characteristics in determining pasta quality in durum wheat (Triticum turgidum ssp. durum), there is no direct report on inheritance of β-carotene concentration. The objectives of this study were to determine the inheritance of β-carotene concentration and the number of genes involved in six crosses of durum. For the cross PDW-233 (P₁) × Bhalegaon-4 (P₂), F₁, F₂, BCP₁ and BCP₂ populations were developed. For all other crosses, only the F₁ and F₂ populations were developed. β-carotene concentration was determined for all populations and parents of each cross grown at Hol, Maharastra, India. The cross PDW-233 × Bhalegaon-4 was also evaluated at Dharwad, Karnataka, India. Low β-carotene concentration was partially dominant in most of the crosses. Broad sense heritability was 67 and 91% at Dharwad and Hol, respectively, for the cross PDW-233 × Bhalegaon-4 and varied from 74 to 93% for the other five crosses indicating the presence of additive gene effects. The frequency distributions of the trait in the F₂ populations were not normal and were skewed towards the lower parent. Segregation of β-carotene concentration in the six F₂ populations indicated that at least two major genes and two or three minor genes with modifying effects govern the trait. Analysis of variance indicated that environment had comparatively little influence on the trait and this should allow for easy selection. The joint scaling test revealed additive × additive, additive × dominance and dominance × dominance epistatic interactions in the cross PDW-233 × Bhalegaon-4. These authors contributed equally.     

Cytological and microsatellite mapping of the genes determining liguleless phenotype in durum wheat

Liguleless phenotypes of wheat lack ligule and auricle structures on all leaves of the plant. Two recessive genes principally control the liguleless character in tetraploid wheat. The F₂ progenies of k17769 (liguleless mutant)/Triticum dicoccoides and k17769/T. dicoccum segregated in a 15:1 ratio, whereas the F₂ progenies of k17769/T. durum and k17769/T. turgidum segregated in a 3:1 ratio. A new gene, lg₃, was found on chromosome 2A. Segregation of F₂ progenies between k17769 and chromosome substitution lines for homoeologous group 2 chromosomes suggested that the liguleless genotype had occurred by mutation at the lg₃ locus on chromosome 2A, and then by mutation at the lg₁ locus on chromosome 2B, in the process of domestication of tetraploid wheat. The gene (lg₁) was linked to Tc₂ (11.9 cM), which determines phenol colour reaction of kernels, on the long arm of chromosome 2B. The distance of lg₁ to the centromere was found to be 60.4 cM, and microsatellite mapping established the gene order, centromere – Xgwm382 – Xgwm619 – Tc₂ – lg₁ on the long arm of chromosome 2B.     

Diallel analysis of sooty stripe resistance in sorghum

Sooty stripe [Ramulispora sorghi (Ellis and Everhart) Olive and Lefebre] is a widespread foliar disease of sorghum [Sorghum bicolor (L.) Moench] in West Africa, responsible for grain yield losses up to 46%. We studied the inheritance of sooty stripe resistance in a 9 × 9 sorghum F₂-population diallel grown together with parent lines and checks in1996 under natural disease pressure at two locations in Mali. The percentage of infected leaf area was determined twice over a two-week interval during the season. At the second evaluation, the mean sooty stripe severity amounted to 13% infected leaf area at Samanko and 12% at Cinzana. The frequency distribution of the entries was approximately normal for the mean disease severity, averaged across assessment dates and locations, pointing to the involvement of multiple genes. With the data combined across the two locations, genetic differences among lines and among F₂ populations were highly significant. Genotype × location interaction variances were also significant but much smaller than the genetic variances. Broad-sense heritability estimates were 0.92 for lines and 0.94 for the F₂ populations, for the mean percentage infected leaf area across the two assessment dates. General combining ability effects (GCA) determined most of the differences among the F₂ populations. Specific combining ability effects (SCA), and the interactions of GCA or SCA with locations were also significant but less important. Line performance per se was highly correlated with GCA. Because of the high heritability and predominance of additive effects, prospects are good for the genetic improvement of resistance to sooty stripe in sorghum in Mali, using simple pedigree or recurrent selection procedures. This revised version was published online in August 2006 with corrections to the Cover Date.     

The inheritance of resistance to head blight caused by Fusarium culmorum in winter wheat

Summary Crosses were made among ten winter wheat genotypes representing different levels of resistance to Fusarium head blight to obtain F₁ and F₂ generations. Parents, F₁ and F₂ were inoculated with one strain of Fusarium culmorum. Data on incidence of head blight 21 days after first inoculation were analyzed. Broad-sense heritabilities averaged 0.39 and ranged from 0.05 to 0.89 in the individual F₂ families. The joint-scaling test indicated that the inheritance of Fusarium head blight resistance was adequately described by the additive-dominance model, with additive gene action being the most important factor of resistance. With respect to the non-additive effects, dominance of resistance predominated over recessiveness. The number of segregating genes governing resistance in the studied populations was estimated to vary between one and six. It was demonstrated that resistance genes differed between parents and affected resistance differently.     

Estimation of genetic parameters for in vitro traits in wheat immature embryo cultures involving high X low regeneration capacity genotypes

Summary Estimates of gene actions were obtained for five in vitro traits of immature wheat (Triticum aestivum L.) embryo cultures from a cross of two wheat cultivars and the resulting reciprocal, F₁, F₂ and backcross populations. The contribution of additive gene effects to in vitro traits was not as important as the dominance gene effects. Epistatic gene effects were relatively more important than either additive or dominance gene effects. Of the individual types of digenic epistatic effects, the dominance x dominance estimates were relatively larger in magnitude for all in vitro culture traits measured. The maternal effect played a minor role in the inheritance of the in vitro studied traits since the difference among the reciprocal values was not significant. It is shown from the generation mean method that epistasis played a major role in the inheritance of most of the traits under study. The negative values of additive and dominance genetic variance were estimates of zero. Heritability estimates, in broad sense, were relatively high for the in vitro studied traits. In some cases, heritability estimates in broad and narrow senses are almost equal since the estimation of dominance genetic variance led to negative values. According to the results of the gene effects, dominance and epistasis were important for the shoot formation trait. Selection would be effective among the isolated genotypes on individual basis.     

A comparison of marker-assisted and phenotypic selection for high grain protein content in spring wheat

Wheat grain protein content (GPC) is a primary end-use quality determinant for hard spring wheat (Triticum aestivum L.), and marker-assisted selection (MAS) could help plant breeders to develop high GPC cultivars. Two experiments were conducted using two populations developed by crossing low GPC cultivars (Ember) and (McVey) with (Glupro), which contains a high GPC QTL from Triticum dicoccoides (DIC). In one experiment, MAS and phenotypic selection (PS) were employed to select high GPC genotypes, and the selected genotypes were grown in six North Dakota (ND), USA environments. In a second experiment, molecular markers were used to select BC₂F₂ plants from each marker class for the DIC allele from each population. These plants were twice self-pollinated to produce BC₂F₄ plants, which were grown in single ND and Minnesota (MN) environments. Mean GPC was highest among lines using PS at two environments and not significantly different between MAS and PS in the other four environments. Lines presumably homozygous for DIC alleles had significantly higher GPC than their respective low GPC parents. The phenotypic GPC variation explained by the markers (r ²) was 30% at the ND and 15% at the MN environment. The use of PS was as effective as MAS in selecting for high GPC genotypes and more effective in some environments. This likely can be attributed to PS enabling selection for both the major QTL and other genes contributing to GPC. The use of molecular markers might be more advantageous for transferring the high GPC DIC QTL in a backcrossing program during parent development.     

Molecular characterization of a novel powdery mildew resistance gene Pm30 in wheat originating from wild emmer

Powdery mildew caused by Erysiphe graminis f. sp. tritici is one of the most important wheat diseases in many regions of theworld. A powdery mildew resistance gene, originating from wild emmerwheat (Triticum dicoccoides) accession `C20', from Rosh Pinna, Israel,was successfully transferred to hexaploid wheat through crossing andbackcrossing. Genetic analysis indicated that a single dominant genecontrols the powdery mildew resistance at the seedling stage. SegregatingBC₁F₂ progenies of the cross 87-1/C20//2*8866 wereused for bulked segregant analysis (BSA). The PCR approach was used togenerate polymorphic DNA fragments between the resistant and susceptibleDNA pools by use of 10-mer random primers, STS primers, and wheatmicrosatellite primers. Three markers, Xgwm159/430,Xgwm159/460, and Xgwm159/500, were found to be linked tothe resistance gene. After evaluating the polymorphic markers in twosegregating populations, the distance between the markers and the mildewresistance gene was estimated to be 5–6 cM. By means of ChineseSpring nullisomic-tetrasomics and ditelosomics, the polymorphic markersand the resistance gene were assigned to chromosome arm 5BS and werephysically mapped on the gene rich regions of fragment length (FL) 0.41–0.43 by Chinese Spring deletion lines. As no powdery mildew resistancegene has been reported on chromosome arm 5BS, the mildew resistancegene originating from C20 should be a new gene and is designated Pm30.     

Variation of harvest index in several wheat crosses

Summary Harvest index was studied in F₁ and F₂ generations of eight wheat (Triticum aestivum L.) crosses and their reciprocals. The parental varieties involved in the crosses represented a fairly wide range of character expression for plant height, tillering potential, grain yield per plant and harvest index. Differences between reciprocal crosses were not evident for the expression of harvest index of the crosses under study. Means and degrees of dominance of F₁ and F₂ populations suggested partial dominance of high harvest index over low harvest index. The pattern of variation among F₂ segregates was quantitative and the distribution was normal. The gene action governing the expression of harvest index was largely additive. Evidence was obtained for non-additive gene action in some crosses. There was complete absence of high parent heterosis for harvest index in the F₁'s but midparent heterosis was found to be present in all crosses. Estimates of heritability and genetic advance were moderate to high. Usefulness of selecting for harvest index as a measure of yield efficiency particularly in early generations is discussed.