Genetic analysis of drought tolerance in adapted × exotic crosses of maize inbred lines under managed stress conditions

Introduced maize (Zea mays L.) germplasm can serve as sources of favorable alleles to enhance performance in new maize varieties and hybrids under drought stress conditions. In the present study, the combining abilities of 12 exotic maize inbred lines from CIMMYT and 12 adapted maize inbred lines from IITA were studied for grain yield and other traits under controlled drought stress. The inbred lines from each institution were separated into groups using SSR-based genetic diversity and were intercrossed using a factorial mating scheme to generate 96 hybrids. These hybrids were evaluated under both controlled drought stress and well-watered conditions at Ikenne in Nigeria in 2010 and 2011. Average mean yields of hybrids under drought stress represented 23 % of the average yield of hybrids under full irrigation. General combining ability (GCA) effects accounted for 49–85 % of the observed variation for several traits recorded under both well-watered and drought stress conditions. Specific combining ability effects for grain yield, though positive in most hybrids, were not significant under drought stress conditions. All the twelve exotic and nine adapted lines had positive GCA effects (female, male, or both) for grain yield under either drought stress or full irrigation, or both environments. EXL03 and EXL15 that had positive and significant female and male GCA effects for grain yield under both environments can be used to improve their adapted counterparts for grain yield and drought tolerance. Normalized difference vegetation index had weak but significant correlation with grain yield.     

Hybrid chlorosis in wheat × rye crosses

Fourteen accessions of rye when crossed to Triticum aestivum cv. C 306 (Ne1ne2ch1Ch2) yielded chlorotic F1 hybrids and six accessions involved in hybrid combination with the same tester produced normal F1 hybrid plants. Two rye accessions, namely, EC 179188 and EC 143825 when crossed to the wheat lines HD 2329 (ne1Ne2ch1Ch2) and NI 5439 (ne1ne2ch1Ch2) also produced chlorosis. The hybrids between T. macha and two rye accessions produced normal plants. Variable degrees of chlorosis were observed among different wheat × rye F1 hybrids. It is suggested that the rye accessions producing chlorosis in combination with wheat cvs. C 306, HD 2329 and NI 5439 (all Ch2-carriers) carry one of the complementary genes for chlorosis. Gene symbol Chr1 is proposed for the chlorosis gene of rye.     

Quantitative trait locus mapping for Verticillium wilt resistance in a backcross inbred line population of cotton (Gossypium hirsutum × Gossypium barbadense) based on RGA-AFLP analysis

Verticillium wilt (VW), caused by Verticillium dahliae Kleb., is one of the most important diseases in cotton. The objective of this study was to map quantitative trait loci (QTLs) conferring VW resistance using resistance gene analog (RGA)-targeted amplified fragment length polymorphism (RGA-AFLP) markers in an interspecific backcross inbred line mapping population, consisting of 146 lines from a susceptible Sure-Grow 747 (Gossypium hirsutum L.) × resistant Pima S-7 (G. barbadense L.) cross. VW resistance was evaluated in replicated tests based on disease incidence in the field, and disease incidence and severity in the greenhouse. Of 160 polymorphic RGA-AFLP markers, 42 were significantly correlated with one or more VW traits and 41 were placed on a linkage map which covered 1,226 cM of the cotton genome and contained 251 other molecular markers. Three QTLs for VW resistance were detected, each of which explained 12.0–18.6 % of the phenotypic variation. Two of these QTLs for disease incidence and severity detected in the greenhouse inoculation tests using root wounding are located on chromosome c4. Both are closely linked to four RGA-AFLP markers and therefore considered as the same QTL for VW resistance. The other QTL detected in the field test was located on c19 and flanked by several RGA-AFLP markers. The desirable QTL allele on c4 for VW resistance detected in the greenhouse was from the VW susceptible Upland parent and absent from the resistant Pima parent which was more VW susceptible due to the disarmament of the first line of defense mechanism due to root wounding during inoculation. The other desirable VW resistance QTL allele, on c19, was from the resistant parent Pima S-7, consistent with the fact that Pima cotton was more resistant to VW when naturally infected in the field. The results should facilitate the development of more sequence specific markers and the transfer of VW resistance from Pima to Upland cotton through marker-assisted selection.     

Development of advanced fragrant rice lines from MR269 × Basmati 370 through marker-assisted backcrossing

Fragrance in rice is an appealing attribute to consumers. The increasing demand for fragrant rice highlights the need to develop fragrant rice variety that suit the preference of local consumers in addition to reduce fragrant rice imports. Marker-assisted backcrossing (MABC) was employed to develop advanced fragrant rice lines from the cross between MR269 and Basmati 370. MR269 is a Malaysian high-yielding rice variety but non-fragrant and was used as recurrent parent whereas Basmati 370 is a well-known fragrant traditional rice variety and was used as donor parent for the fragrance gene. Two generations of backcrosses and a generation of selfing were conducted to introgress the fragrance gene and restore the recurrent parent genome in the backcross progenies. As a result, 14 advanced fragrant rice lines were developed. These advanced fragrant rice lines carried homozygous alleles for the fragrance gene, similar to Basmati 370. The average recovery of recurrent parent genome was 88.4%. Besides being fragrant, the advanced fragrant rice lines also had most of the morphological and agronomical traits similar to MR269. Grain quality of the advanced fragrant rice lines in terms of gelatinization temperature, amylose content and gel consistency are also similar to both parents. Besides, the advanced fragrant rice lines had 2-acetyl-1-pyrroline content similar to Basmati 370. MABC approach applied in this study has successfully introgressed the fragrance gene and accelerated the recovery of recurrent parent genome in advanced fragrant rice lines, therefore these lines can be delivered to the farmers and consumers for use in due time.     

Analysis of endosperm development of allotriploid × diploid/tetraploid crosses in <Emphasis Type="Italic">Lilium</Emphasis>

Lilium are Fritillaria-type plants. Triploid lilies, regardless of their male sterility, can be used as female parents to cross with appropriate males, in contrast to other triploid Polygonum-type plants, which are usually seedless. Up to now, however, little attention has been paid to the reason. Here we made allotriploid × diploid/tetraploid crosses in Lilium. Endosperm and progenies of LAA × AA/AAAA crosses were analyzed for ploidy level and genome composition. The endosperm of some LAA × AA/AAAA ovules developed well but there was little or no development of LAA × LL/OO endosperm. The endosperm genome composition of LAA × AA, LAA × AAAA, LAA × LL, LAA × OO are theoretically derived as 5A + 2L, 6A + 2L, 4A + 3L, 4A + 2L + O, respectively. Genomic in situ hybridization showed that the progenies of LAA × AA/AAAA were aneuploid. Based on the experimental results and analyses, our hypothesis is that five same genomes of endosperms are essential for its development in triploid × diploid/tetraploid crosses of Lilium. The hypothesis can explain the success or failure of 3x × 2x/4x crosses in Lilium and is of importance for lily breeders who need to know the likelihood of success when producing new cultivars of this economically important horticultural crop.     

Epistasis in an Andean × Mesoamerican cross of common bean

The objective of this investigation was to check if epistasis is present in Andean × Mesoamerican beans crosses using triple test cross (TTC) method. The parents of the segregating population were Carioca–MG (Mesoamerican) and BRS Radiante (Andean). In July 2005, F₂ progenies (backcrossed with the parents and F₁ generation) were evaluated at two locations for three characters: number of pods plant⁻¹, number of grains plant⁻¹ and grain weight plant⁻¹. The presence of epistasis was detected for all yield components. In the partitioning of epistasis in additive x additive (i) and dominant x dominant (j) and dominant × additive (l) it was observed that, for the traits number of pods/plant and number of grains/plant, only epistasis of the type (j) + (l) were significant. For the trait grain mass/plant, all types of epistasis were significant.     

Effect of parental genotypes on haploid embryo and plantlet formation in wheat × maize crosses

Genotypic influence of both male and female parents on haploid production through interspecific crosses was studied using eight wheat and four maize genotypes. The average numbers of embryos and green haploid plantlets obtained per pollinated floret were 17.6% and 10.1%, respectively. Clear genotypic influence of the wheat genotype was detected, but heterozygosity of the wheat did not affect haploid production. Analogous response to anther culture and interspecific crossing was observed, still a wheat variety which did not respond to anther culture, produced 1.1 plantlets per pollinated spike upon maize pollination. This appears to be a major advantage of interspecific crossing compared to anther culture technique in wheat. Circumstantial evidence is presented for specific wheat × maize interaction on haploid plantlet formation. Rye chromatin enhanced haploid production but only in a complete 1B/1R substitution line. Ovaries with an embryo were found to be dispersed evenly all over the wheat spike, suggesting that within certain limits the developmental stage of ovaries and thus time of pollination within a spike are not as important as it was previously assumed. This revised version was published online in July 2006 with corrections to the Cover Date.     

A male-sterile line with ‘virescent’ marker character in Upland cotton

Summary Inheritance of 81A genetic male-sterility with virescent marker character and its cytology in Upland cotton (Gossypium hirsutum L.) are presented. The chromosome number of 81A is normal, 2n = 52, and pollen abortion occurs mainly in the late uninuclear stage. Male-sterility and virescence in 81A are controlled by one pair of recessive gene. Because male sterility and virescence showed no recombination, pleiotropy is assumed. However, very close linkage could not be ruled out. Allelic tests indicated that virescence observed in 81A is non-allelic to all virescence genes identified earlier in Upland cotton and its male-sterility is non-allelic to msc₁, msc₂ and msc₃ discovered in P.R. China. It is proposed that 81A male-sterile gene symbol be tentatively named to msc₇. The development of 81A genetic male-sterile line associated with virescence trait could raise considerably the efficiency of the hybrid seed production in cotton.     

Parental effects on the performance of cultivated × wild species hybrids in potato

Valuable genetic diversity in diploid wild Solanum species can be accessed through crosses to haploids (2n = 2×) of the tetraploid cultivated potato, Solanum tuberosum. Haploid-wild species hybrids segregate for the ability to tuberize in the field. In addition, they vary in male fertility, vine size, stolon length, and tuber size. In this study, three haploids were crossed with nine diploid wild Solanum species and 27 hybrid families were evaluated in the field for two years. The proportion of male fertile hybrid clones varied depending on the wild species parent. A large effect of the female parent was detected for vine size, stolon length, tuber size, percent tuberization, and percent plants selected for agronomic quality. An exceptional haploid (US-W4) was identified for the production of agronomically desirable haploid-wild species hybrids. In hybrids derived from US-W4, differences among wild species parents were observed for agronomic quality. Superior hybrids were produced by S. berthaultii and S. microdontum. Reciprocal crosses were evaluated for a subset of families. When the wild species was used as the female parent, male fertility was restored, but tuberization and tuber size were reduced. Careful selection of both haploid and wild species parents can result in a large proportion of fertile, agronomically desirable hybrid offspring.     

Inheritance of anthocyanin content in the ripe berries of a tetraploid × diploid grape cross population

Variation patterns and inheritance of anthocyanin content in the ripe berries of a tetraploid × diploid table grape cross population were investigated in two successive years. The population segregated for three different ploid levels: dipolids, triploids, and tetraploids. A total of 28 different anthocyanins were detected and quantified in the progeny population. Transgressive segregation for the total anthocyanin content was observed in all the three ploid progeny populations. The total anthocyanin content increased as the ploid level increased. The broad sense heritabilities (H²) of the total anthocyanin content were all relatively high, ranging from 0.53 to 0.98, 0.57 to 0.97 and 0.43 to 0.94 in the diploid, triploid and tetraploid population, respectively. Our results suggested that the total anthocyanin content followed an additive inheritance model in this polyploid segregation population. We also observed that the relative contribution of individual anthocyanins to the total anthocyanin content varied significantly among different ploid populations, suggesting that genetic background has important impact on the accumulation of the individual anthocyanin compounds. These results will help develop better breeding strategies in a polyploid table grape breeding program for improving the content of anthocyanins, an important class of polyphenolics possessing antioxidant activities and many other health-related benefits.     

Phenotypic segregation and relationships of agronomic traits in Monastrell × Syrah wine grape progeny

The aim of this work was to study the phenotypic segregation of different agronomic and fruit quality traits, and their relationships, in Monastrell × Syrah wine grape progeny. Twenty-two agronomic traits were evaluated and compared for three consecutive years in this progeny. The results show the phenotypic diversity existing in a cross between two different wine grape cultivars. Most of the phenological, productive, morphological, and enological parameters evaluated displayed continuous variation within the progeny, suggesting a polygenic inheritance. Some correlations between traits were detected by the Spearman correlation test, although high coefficients were not found for most of them. Cluster analysis of the progeny grouped the hybrids based on criteria with significance for wine grape breeding. Also, we investigated the relationship between the skin color and total content of anthocyanins with the VvmybA genotype, using the CAPS (Cleaved Amplified Polymorphic Sequence) marker 20D18CB9. The results show that hybrids with two copies of the functional color allele tend to have increased anthocyanins content. Based on this study, 14 genotypes were pre-selected from the breeding population for additional quality studies.     

Screening for grain dormancy in segregating generations of dormant × non-dormant crosses in white-grained wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Pre-harvest sprouting (PHS) in wheat (Triticum aestivum L.) is a significant problem. Introgression of genes controlling grain dormancy into white-grained bread wheat is one means of improving resistance to PHS. In this study seven dormant (containing the SW95-50213 and AUS1408 sources) × non-dormant crosses were produced to investigate the effectiveness of selection for grain dormancy in early segregating generations. Each generation (F₁–F₄) was grown in a temperature controlled glasshouse with an extended photoperiod (i.e. continuous light). F₂ and F₃ generations were subject to selection. Five hundred harvest-ripe grains were tested for germination over a 14 day period, and the 100 most dormant grains were retained and grown-on to produce the next generation within each cross. The response to selection was assessed through analysis of the time to 50% germination (G₅₀) in the F₂, F₃ and F₄ generations. In addition, changes in marker class frequencies for two SSR markers (barc170 and gpw2279) flanking a known quantitative trait locus (QTL) for grain dormancy on chromosome 4A were assessed in DNA from F₂ plants selected from early germinating (non-dormant) and late germinating (dormant) phenotypic extremes within each cross. Selection for grain dormancy in the F₂ and F₃ generations effectively recovered the dormant phenotype in all seven crosses, i.e. the F₄ generation was not significantly different from the dormant parent. Further, selection based on individual F₂ grains changed marker class frequencies for the 4A dormancy QTL; in most cases eliminating the marker class homozygous for the non-dormant alleles. Application of this screening method will enable breeders to better select for grain dormancy and may lead to development of new cultivars offering effective resistance to PHS in the near future.     

Embryo lethality in wheat × rye hybrids—mode of inheritance and the identification of a complementary gene in wheat

Crosses between hexaploid wheat and rye can only succeed when pre- and post-zygotic barriers have been overcome. A rye gene determining embryo lethality (Eml-R1), which is involved in post-zygotic isolation, has been mapped to chromosome 6R. In the present paper the mode of inheritance of Eml-R1 was studied by employing a wheat/rye chromosome 6R addition line. We show that Eml-R1 exists in at least two alternative forms, with the mutant allele Eml-R1b being dominant with respect to wild-type Eml-R1a. Furthermore, we have exploited nulli-tetrasomic lines of wheat to detect a complementary wheat gene present on chromosome 6A. This gene has been designated Eml-A1.     

Generation and characterisation of colchicine-induced polyploid Lavandula × intermedia

Double podding in cultivated chickpeas (Cicer arietinum L.) can increase yield and yield stability. In the present study, we performed reciprocal crosses of ‘kabuli’ (double podded) and ‘desi’ (single podded) chickpeas to determine (i) the expressivity and penetrance of double podding, (ii) the correlations of yield and yield components, and (iii) the heritability of double podding, flower color, and stem pigmentation in F₂ plants. Reciprocal crosses were performed with two genotypes, AC 2969 (kabuli) and ICC 4969 (desi), to generate F₁ and F₂ plants. The results indicated hybrid vigor (heterosis) for yield in F₁ plants and better performance of F₂ plants. Yield and yield components of some lines in F₂ were superior to the best parent, indicative of transgressive segregation. In particular, the presence of double podding (‘s’ allele) significantly increased yield in some of the transgressive segregants. Expressivity and penetrance of the ‘s’ allele depends on the background of the female parent. Some of the double podding progeny had greater seed yields than those of the single podding progeny and greater seed yields than the best parents. Double podding, stem pigmentation, and pink flowers each appears to be governed by a single recessive gene. Stem pigmentation and pink flowers appear to be linked traits that depend on the genetic background of the crossed chickpeas. Taken together, our studies of reciprocal crosses of kabuli and desi chickpeas clearly showed that yield could be improved by selection for transgressive phenotypes that have double podding.     

Genetic analysis and genotype × environment (G × E) for grey leaf spot disease resistance in elite African maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) germplasm

Maize grey leaf spot (GLS) disease remains an important foliar disease in sub-Saharan Africa accounting for more than 25% yield losses in maize. Information on inheritance of GLS resistance of germplasm adapted to African environments is required in new sources being identified. Therefore, hybrids generated from a 10 × 10 half-diallel mating of tropical advanced maize inbred lines were evaluated in six environments to determine combining ability, genotype × environment interaction (G × E) and the impact of GLS disease on grain yield. General combining ability effects were highly significant and accounted for 72 and 68% of the variation for GLS resistance and grain yield, respectively. Significant specific combining ability effects associated with reduced disease levels were observed in some hybrids when one parent was resistant, and these may be exploited in developing single cross maize hybrids. Regression analysis showed a 260–320 kg ha^?1 decrease in maize grain yield per each increase in GLS disease severity score, and significant associations (r = ?0.31 to ?0.60) were observed between grain yield and GLS severity scores. This showed the potential of GLS disease to reduce yield in susceptible varieties grown under favourable disease conditions, without control measures. Genotype and genotype × environment biplots and correlation analysis indicated that the significant G × E observed was not due to changes in hybrid ranking, implying absence of a significant crossover interaction. Therefore, predominance of additive gene effects imply that breeding progress for GLS disease resistance would be made through selection and this could be achieved at a few hot-spot sites, such as Baynesfield and Cedara locations in South Africa, and still deploy the resistant germplasm to other environments in which they are adapted.     

Inheritance of sugar and acid contents in the ripe berries of a tetraploid × diploid grape cross population

Pearl millet [Pennisetum glaucum (L.) R. Br.] is a major warm-season cereal, grown primarily for grain production in the arid and semi-arid tropical regions of Asia and Africa. Iron (Fe) and zinc (Zn) deficiencies have been reported to be a food-related primary health problem affecting nearly two billion people worldwide. Improving Fe and Zn densities of staple crops by breeding offers a cost-effective and sustainable solution to reducing micronutrient malnutrition in resource poor communities. An understanding of the genetics of these micronutrients can help to accelerate the breeding process, but little is known about the genetics and heterosis pattern of Fe and Zn densities in pearl millet. In the present study, ten inbred lines and their full diallel crosses were used to study the nature of gene action and heterosis for these micronutrients. The general combining ability (GCA) effects of parents and specific combining ability (SCA) effects of hybrids showed significant differences for both of the micronutrients. However, the predictability ratio (2σ²gca/(2σ²gca + σ²sca)) was around unity both for Fe and Zn densities, implying preponderance of additive gene action. Further, highly significant positive correlation between mid-parent values and hybrid performance, and no correlation between mid-parent values and mid-parent heterosis confirmed again the predominant role of additive gene action for these micronutrients. Barring a few exceptions with one parent, hybrids did not outperform the parents having high Fe and Zn levels. This showed that there would be little opportunity, if any, to exploit heterosis for these mineral micronutrients in pearl millet. In general, high Fe and Zn levels in both of the parental lines would be required to increase the probability of breeding high Fe and Zn hybrids.