Ergot reaction of pearl millet hybrids affected by fertility restoration and genetic resistance of parental lines

Summary High ergot (Claviceps fusiformis Loveless) susceptibility of pearl millet (Pennisetum glaucum (L.) R. Br.) hybrids has often been associated with the A₁ cytoplasm of male-sterile lines (A-lines). To understand the underlying basis of this association and to examine the prospects of breeding ergot-resistant hybrids, we evaluated 56 hybrids and their 15 parental lines for ergot reaction and selfed seedset for 2 years in disease nurseries at ICRISAT Asia Center. Hybrids were made by crossing seven pollen parents (2 susceptible and 5 resistant) onto two resistant and two susceptible A-lines, and their four corresponding maintainer lines (B-lines). A-lines had no selfed seedset while B-lines had 32–75% selfed seedset. Hybrids of A-lines had significantly less selfed seedset than the hybrids of the corresponding B-lines. The reduced seedset of A-lines and their hybrids, however, was not always accompanied by significantly higher ergot susceptibility. Highly resistant hybrids were obtained where both A-lines and pollen parents were highly resistant, regardless of male fertility levels of the hybrids. Thus, although the A₁ cytoplasm, by its reduction of male fertility, had a large and significant effect in increasing ergot severity of hybrids, the contribution of nuclear genetic factors of female parents was about 1.8 times larger than that of the cytoplasm.Submitted as JA No. 1776 by the International Crops Research Institute for the Semi Arid Tropies.     

Smut severity of pearl millet hybrids with male sterile and fertile cytoplasm

Summary Effect of A₁ male sterile cytoplasm on smut severity in pearl millet (Pennisetum glaucum (L.) R.Br.) was studied by comparing 35 pairs of F₁ hybrids, each pair carrying male sterile and normal cytoplasm. Mean smut severity was not significantly different in the hybrids carrying male sterile or normal cytoplasm. This suggests that in pearl millet male sterile cytoplasm is not associated with higher smut susceptibility. Partitioning of variance into different components showed that pollinators, A/B line pairs and their interaction primarily influenced smut severity of hybrids. Smut susceptibility might be attributed to effects of cytoplasm × nuclear interactions.     

Stability of male-sterile sources and fertility restoration of their hybrids in pearl millet

Genetic background has a significant effect on the expression of cytoplasmic-nuclear male sterility (CMS) in pearl millet (Pennisetum glaucum (L.) R. Br.). Therefore, a reliable characterization of CMS sources requires the use of near-isonuclear lines and their hybrids. We used this approach to characterize five CMS sources (A₁, A₂, A₃, A_v, and A₄). Male sterility of 81A₄ was the most stable, followed by 81A₁ and 81A_v, indicating the relative merits of these CMS sources in breeding stable male-sterile lines. Male-sterile lines 81A₂ and 81A₃ were highly unstable, indicating their minimal value. Differential male fertility restoration patterns of hybrids made on 81A_V and 81A₄ suggest that the A_v and A₄ cytoplasms represent CMS systems that are different from each other and from A₁, A₂ and A₃. An evaluation of topcross hybrids of 15 diverse populations made on 81A₁ and 81A₄ showed that each population had restorers and maintainers of both cytoplasms and that the frequency of maintainers of A₄ was as high as, or higher than, that of the A₁ cytoplasm. Thus, use of the A₄ cytoplasm can substantially increase the probability of breeding stable male-sterile lines based on inbreds developed from diverse genetic backgrounds, and also provide the opportunity for breeding restorers from each of these diverse genetic sources.     

Commercial viability of alternative cytoplasmic-nuclear male-sterility systems in pearl millet

Commercial viability of three cytoplasmic-nuclear male sterility (CMS) systems (A₄, A₅ and A_v) as potential alternatives to the most widely used A₁ system in pearl millet (Pennisetum glaucum (L.) R.Br.) was evaluated in terms of stability of complete male sterility of four isonuclear A-lines (81A₁, 81A₄, 81A₅ and 81A_v) and the level and stability of male fertility restoration of their 44 single-cross hybrids. Lines 81A₄ and 81A₅ had no pollen shedders (PS), and there were very low frequency of non-PS plants of these A-lines that had a maximum of 1–5% selfed seedset (SSS). In 81A₁ and 81A_v,there were, albeit low frequency (<1%) of PS plants, and relatively higher frequency of the non-PS plants in these two lines, the more so in 81A_v,had 1–5% and even greater SSS. Some hybrids made on each of the three A-lines (81A₁, 81A₄ and 81A_v) had high and stable male fertility, while others made on the same three A-lines displayed large variation in SSS across the environments, the more so in case of hybrids made on 81A_v. These results indicate that the A₄ CMS system provides a better alternative to the A₁ CMS system, while the A_v system does not. On the basis of highly stable male sterility and the highest frequency of pollinators behaving as maintainers, the A₅ CMS system appeared to be the best for A-line breeding. The commercial viability of this CMS system in breeding R-lines of grain hybrids, however, still remains to be ascertained as no hybrid on it was fully male fertile in any environment. This revised version was published online in August 2006 with corrections to the Cover Date.     

Cytoplasmic differentiation using near-isonuclear polycytoplasmic male sterile lines in pearl millet

Summary Five near-isonuclear polycytoplasmic versions of 81A₁ and two of Pb 402A₃ male sterile lines of pearl millet (Pennisetum typhoides), along with their corresponding maintainer lines (81B and Pb 402B) were studied for 14 agronomic and two disease traits, and for isozymes of peroxidase in anthers and leaves for assessing cytoplasmic differences. Significant differences among isonuclear polycytoplasmic lines of both 81A₁ and Pb 402A₃, having the same genome and variable cytoplasms, were observed for several agronomic traits. Banding pattern of peroxidase isozymes revealed clear cut differences among cytoplasms. The results provide evidence for the influence of cytoplasmic factors on the phenotypic expression of nuclear genes.     

Characterization of seed producing pearl millet x elephantgrass hexaploid hybrids

Summary Successful widespread use of a forage or biomass plant depends largely on its ease of establishment. Elephantgrass (Pennisetum purpureum Schum.) is used for both forage and biomass production, due to its high dry matter production, aggressiveness, perenniality, and forage quality. However, lack of high quality seed has limited the use of elephantgrass and its hybrids to pearl millet (P. glaucum L. R.Br.). Development of a seed-propagated pearl millet x elephantgrass hexaploid cultivar, which would combine desirable characteristics from both of these species, would be highly desirable. The objectives of this study were to characterize morphological and seed-related traits from the selfed progeny of seven hexaploid hybrids, and to determine whether seed size had any influence on these traits. Traits studied included number of tillers, height, leaf length and width, panicles per plant, days to flowering, panicle length, seed set, seed production, and weight of 100 seeds. Genetic differences were found among the progeny of the seven hybrids. Differences were also found between plants derived from large- and small-seed lots within families. Plants descended from larger seed had better growth and seed-related characteristics, which resulted in more desirable plants. These hexaploid hybrids showed potential for direct seeding into the field for biomass or forage production.     

Performance of pearl millet isonuclear hybrids involving different cytoplasmic male-sterility systems

The performance of pearl millet hybrids involving four sources of cytoplasmic male sterility (CMS), including the most-exploited A₁ source, was compared to evaluate the effects of cytoplasm on grain yield. The mean grain yield of hybrids possessing A₂, A₃ and A₄ cytoplasms was either similar to or significantly higher than that of their counterpart hybrids with A₁ cytoplasm. Hybrids based on A₃ and A₄ cytoplasms produced, on average, 8% more grain compared with those based on A₁ cytoplasm. This suggested that these CMS sources could be used as alternatives to A₁ cytoplasm to widen the cytoplasmic base of hybrids. The results indicated that most of the variation in hybrids was accounted for by pollinator and cytoplasm × pollinator interactions, suggesting the use of genetically diverse pollinators in pearl millet hybrid breeding.     

Stability of forage yield and quality in pearl millet hybrids heterogenous for rust resistance

Summary The susceptible pearl millet hybrid Tifleaf 1, the resistant hybrid Tifleaf 2, and two experimental 3-way hybrids with different proportions of resistant and susceptible plants were evaluated for rust resistance and forage yield and quality in 1990, 1991, and 1992. Different environmental conditions were obtained by varying planting date, planting density, and fungicide applications across three years of evaluation in the field. Rust severity of forage was negatively correlated with late season green yield, dry matter yield, in vitro dry matter digestibility, and digestible dry matter yield. The stability across environments of the 3-way hybrids was intermediate between resistant Tifleaf 2 and susceptible Tifleaf 1 for rust severity and the yield and quality measurements negatively correlated with rust severity. Mixtures of resistant and susceptible plants would probably provide greater control of rust than measured in these small-plot experiments. Although a mixture of resistant and susceptible plants appears to provide an alternative to monogenic control of rust in forage pearl millets, identification and utilization of additional sources of resistance would be beneficial to improve the performance of 3-way hybrids.     

Evaluation of landrace topcross hybrids of pearl millet for arid zone environments

Pearl millet (Pennisetum glaucum (L.) R. Br.) cultivars for marginal, arid environments need to combine the adaptation to stress conditions of indigenous landraces with an improved yield potential and disease resistance, to allow them to both perform well in farmers fields and to meet the requirements for cultivar release. This paper evaluates landrace-based topcross hybrids (adapted landraces crossed on high-yielding male-sterile lines), as a quick and efficient way of achieving this objective. Topcross hybrids showed a consistent increase in biomass production across all test environments, including the harsh arid zone environments. Depending upon the plant type of the male-sterile used to make the hybrid, this was expressed as increased grain yield only, or increased grain and fodder yields. The downy mildew (Sclerospora graminicola) reaction of the topcross hybrids was determined by the reaction of the male-sterile line used, with the resistant male-sterile producing resistant topcross hybrids and vice-versa. Topcrossing adapted landraces on high-yielding male-sterile lines thus provides an opportunity to improve disease resistance and grain and/or fodder yields, with no apparent loss of adaptation to the marginal environments in which the landraces have evolved.ICRISAT Journal Article no. 1575     

Influence of cytoplasmic-nuclear male sterility systems on microsporogenesis in pearl millet (Pennisetum glaucum (L.) R. Br.)

Influence of a range of cytoplasms on microsporogenesis and anther development in pearl millet was studied using six isonuclear A-lines having five cytoplasms (A₁, A₂, A₃, A₄ and A_v) and the nuclear genome of 81B. 81B was used as a male-fertile control. Microsporogenesis and anther development were normal in 81B. However, pollen mother cell (PMC)/microspore/pollen degeneration in the six A-lines occurred at different stages of anther development. Each cytoplasm had its unique influence on microsporogenesis and anther development as evidenced by different developmental paths followed by them leading to pollen abortion. The cause of pollen abortion differed from line to line, from floret to floret within a spikelet, from anther to anther within a floret, and in some cases even from locule to locule within an anther. Events that led to male sterility included anomalies in tapetum and callose behaviour, persistence of tapetum, endothecium thickness, and other unknown causes. The present study also indicated that anther/pollen development was more irregular in Pb 406A₃. In 81A₄ and 81A₁ > 95% of anther locules followed a definite developmental path to pollen abortion. In the other A-lines many developmental paths were observed within the line and pollen degeneration occurred at various stages. This could be one of the reasons for greater instability of male sterility in the A₂ and A₃ systems and greater stability of male sterility in the A₁ and A₄ systems. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effect of d2 dwarfing gene on grain yield and yield components in pearl millet near-isogenic lines

Summary A d₂ dwarfing gene in pearl millet [Pennisetum glaucum (L.) R. Br.] is currently being extensively used for the development of hybrid parents. Its effect on grain yield and yield components is poorly understood. Twelve pairs of tall and dwarf near-isogenic lines developed in the diverse genetic background of three composites were evaluated for grain yield and yield components for 2 years at two locations in southern India. The d₂ gene or the genes linked to it, on an average, reduced plant height by 42%, grain yield by 14%, and head girth by 8% but increased head length and number of tillers per plant by about 5–6%. Large variations were observed among pairs (genetic background) for the difference between tall and dwarf near-isogenic lines for all of the above yield components resulting in no significant difference in five pairs and 17–35% less yield in dwarfs as compared to their tall counterparts in six pairs. Days to 50% flowering and seed weight were least affected by the d₂ gene with the average difference between tall and dwarf groups of near-isogenic lines being of the order of 1–2%. These results indicate that the advantageous effects of d₂ dwarfing gene can be effectively exploited by manipulating the genetic background. The difference between the average grain yields of tall and dwarf groups of near-isogenic lines showed considerable variation across environments with the dwarfs yielding as much as tall group in one environment and up to 30% less than the tall group in the other, thus, indicating that the d₂ gene effect may be substantially modified by the environments.Submitted as JA No. 979 by the International Crop Research Institute for the Semi-Arid Tropics (ICRISAT).     

Inheritance of threshing percentage in pearl millet

Summary The threshing percentage (TH%) has been suggested as a selection criterion to identify the pearl millet (Pennisetum glaucum (L.) R.Br.) lines with improved ability to fill and set grains under water limiting conditions. In this study, eight genetically diverse pearl millet inbreds and their 28 crosses produced by half diallel crossing design were used to examine range in TH%, to evaluate general combining ability (GCA) of parents and specific combining ability (SCA) effects of crosses. The results showed significant variation among parental lines for TH%. The inbreds differed for their GCA effects and crosses for their SCA effects. Parents with high TH% and positive GCA effects were identified. The results of the study revealed that both additive and dominance components, with the preponderance of later, were important in the inheritance of TH%. Heritability in narrow sense was moderate (55%) indicating that selection for high TH% might be effective.     

Genotypic differences in callus growth and organogenesis of eight pearl millet lines

Summary Calli were obtained from mesocotyl explants of eight varieties of pearl millet using MS medium supplemented with 2,4-D. Variable frequencies of callus initiation were observed among the varieties. Plantlets were regenerated from the calli by transferring on to the medium devoid of auxins. Incorporation of coconut milk, Kinetin, IAA or NAA did not enhance either the callus initiation or growth rate or plantlet regeneration. The eight genotypes differed among themselves in callus growth rate and frequency of shoot bud production.     

Utility of pearl millet landraces in breeding dual-purpose hybrids for arid zone environments of India

In pearl millet (Pennisetum glaucum (L.) R. Br.), single cross hybrids based largely on exotic germplasm, have found very limited adoption in drought-prone arid regions mainly due to their inadequate adaptation to prevalent extremely harsh agro-climatic conditions. This study tested the hypothesis that the use of restorer parents derived from adapted landrace germplasm could be an alternative strategy in developing suitable hybrids for arid regions. Forty inbred restorer lines developed from two phenotypically diverse landraces were used to develop hybrids evaluated in this study for their performance in five typical arid zone environments between 2003 and 2007. Inbred lines from both landrace populations differed significantly in their combining ability for biomass, harvest index and grain and stover yields. A larger proportion of experimental hybrids, based on selected inbred lines, significantly outperformed the commercial checks for grain and stover yields. Increase in grain and stover yields was achieved primarily due to their higher biomass productivity, with no decline in harvest index. Choice of landrace determined the relative grain and stover productivity of their hybrids. Phenotypic differences observed in parental landraces in tillering and panicle length were also visible in their respective hybrids, indicating that characteristics of landrace parental populations were transmitted in their hybrids.     

Seed parent breeding efficiency of three diverse cytoplasmic-nuclear male-sterility systems in pearl millet

Pearl millet (Pennisetum glaucum (L.) R. Br.) hybrids, grown widely in India and to some extent in the US, are all based on an A₁ CMS source, leaving the pearl millet hybrids vulnerable to potential disease or insect pest epidemics. A comparison of this CMS system with two additional CMS systems (A₄ and A₅) in the present study based on isonuclear A-lines (seed parents) and their isonuclear hybrids showed that A-lines with the A₄ cytoplasm had much fewer pollen shedders and much reduced selfed seed set in visually assessed non-shedding plants as compared to those with the A₁ cytoplasm. A-lines with the A₅ cytoplasm had neither any pollen shedders nor did they set any seed when selfed. This showed that the A₅ CMS system imparts complete and most stable male sterility, followed by the A₄ and A₁ CMS systems. The frequency of maintainers, averaged across a diverse range of 26 populations, was highest for the A₅ CMS system (98%), followed by the A₄ (59%) and the A₁ (34%) system indicating the greatest prospects for genetic diversification of A-lines lies with the A₅ cytoplasm, and the least with the A₁ cytoplasm. Mean grain yield of hybrids with the A₁ cytoplasm was 5% more than the A₄-system hybrids, while there was no difference between the mean grain yield of hybrids based on A₁ and A₅ CMS systems. Based on these results, it is suggested that seed parents breeding efficiency will be the greatest with the A₅ CMS system, followed by the A₄ CMS system, and least with the currently commercial A₁ CMS system.     

Inheritance of genetic malesterility in Pennisetum americanum (L.) Leeke

Summary A spontaneously occurring male sterile condition was located in an inbred line Vg 272 of Pennisetum americanum. Male sterility was inherited as a monogenic recessive condition and showed independent assortment with hairy leaf surface phenotype. Transfer of male sterile allele to inbreds Vg 212 and IP 482 revealed no cytoplasmic effects on the expression of male sterility. Stigmas of the male sterile plants were found to possess the same amount of receptivity upto three days with normal seed set, upto six days with reduced seed set and almost with nil receptivity on the seventh day.     

Effect of water availability pattern on yield of pearl millet in semi-arid tropical environments

Summary Throughout much of the semi-arid tropics, fluctuations in grain yield can largely be attributed to differences in timing and intensity of drought stress. Since seasonal rainfall in these environments is often poorly related to grain yield, the aim of this paper was to establish a relationship between water availability and grain yield for pearl millet (Pennisetum glaucum (L.) R. Br.), grown across 24 semi-arid tropical environments in India. We used a simple soil water budget to calculate a water satisfaction index (WSI) throughout the season. The cumulative WSI at maturity explained 76% of the variance in grain yield. This was three times as much as explained by actual rainfall, because WSI accounted for differences in water losses and pan evaporation. A classification of environments into four groups of water availability patterns explained 75% of the environmental sum of squares for grain yield. For a subset of 13 environments, environmental differences in grain number could also be explained by water availability patterns, whereas differences in grain mass were related to both water availability and temperature. Our results indicate that cumulative WSI, which is an integrated measure of plant-available water, can provide an adequate estimation of the environmental potential for yield in environments where grain yield is mainly limited by variable availability of water.ICRISAT Journal Article 1637     

Heterosis in landrace-based topcross hybrids of pearl millet across arid environments

This study quantified the magnitude of heterosis in pearl millet (Pennisetum glaucum) topcross hybrids produced by crossing 16 diverse landraces and three high yielding open-pollinating varieties on two homozygous male-sterile lines. Hybrids and pollinators were grown in 12 year ×;location combinations in India that were grouped into three zones. Genetic components of variance quantifying the differences among these hybrids were estimated. The hybrids showed a conspicuous heterosis for grain yield, earliness and biomass yield but not for straw yield. The level and direction of heterosis for time to flowering depended strongly on the earliness of the male-sterile line. In the terminal drought stress zone hybrids made on the early maturing male-sterile line 843A had the highest level of heterosis for grain yield (88%). This was partly due to escape from terminal stress. In the other two zones the heterosis for grain yield was on average 30%. Heterosis for biomass yield and biomass yield per day was on average also positive in all three zones. For all traits, except time to flowering and biomass yield per day, pollinator effects were the only significant source of variation. Differences between hybrids were mostly caused by additive genetic effects. Significant amount of heterosis observed in landrace-based topcross hybrids for grain yield and other productivity-related traits suggested that substantial improvement in pearl millet productivity in and environments can be obtained by topcrossing locally adapted landraces on suitable male-sterile lines. This revised version was published online in July 2006 with corrections to the Cover Date.     

Structure of variation among morphological and physiological traits in three pearl millet composites

Summary The plant breeder's task of improving and stabilizing many plant traits simultaneously is complicated by interrelationships that occur among the traits. Factor analyses were conducted on three phenotypically diverse pearl millet [Pennisetum glaucum (L.) R. Br.] composites to describe the structure of relationships among yield, morphological, and physiological traits. Approximately 1000 S₀ spaced-plants from each composite were evaluated for 20 traits, and random samples of 289 S₁ progenies from each composite were evaluated for 18 of these traits. Factors extracted in S₀ and S₁ populations identified unique sets of traits that were interrelated along axes of (a) biological yield, (b) panicle size, (c) dry matter partitioning and (d) compensation between number and size of seeds. Several plant traits had large loading coefficients on the Biological Yield and also, but with opposite signs, on the Dry Matter Partitioning factor. The traits having large loading on these two factors differed between space-planted and normal-density stands, showing that environmental conditions contributed to the associations observed among traits. Correlations of S₁ with parental S₀ factor scores for the Biological Yield, Panicle Size and Seed Paramters factors produced significant correlation coefficients, indicating that these trait complexes had a genetic basis. The implications of these results for millet breeding are discussed.