Genetics of induced dwarf mutants in pearl millet,Pennisetum americanum (L.)Leeke

Summary Inheritance pattern of seven dwarf mutants revealed each of them to be controlled by a single recessive gene. Tests for allelism indicated existence of two groups of dwarfs, group 1 comprising six mutants and group 2 a single dwarf. The dwarfs of group 1 were found to be allelic to D₂ of the dwarf types already known in this crop and the dwarf of group 2 was allelic to D₁ of these types.Botany Department, R.V.V.N. College, Dharanikota, India.     

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.     

Confirmation of a new source of cytoplasmic-genic male-sterility in bulrush millet (Pennisetum americanum (L.) Leeke)

Summary Results of a previous study indicated that the male-sterility discovered in a population of ex-Bornu bulrush millet, an improved selection from northern Nigeria, may represent a new source of male-sterility in the species. The present experiment was conducted to test this hypothesis.Tifton 23B₁, Tifton 239B₂ and L67B₃ which are male-sterility maintainers for the three known sources of cytoplasmic-genic male-sterility in bulrush millet viz. A₁, A₂ and A₃, respectively, were crossed with male-sterile ex-Bornu. All three maintained male-sterility in ex-Bornu but not as well as gero-B, which is the selfed progeny of an ex-Bornu plant was an excellent maintainer for ex-Bornu. Gero-B maintained male-sterility in Tifton 23A₁ and L67A₃ sources but was a partial fertility restorer for Tifton 239A₂. Tifton 186 which was reported to be a maintainer for A₁, A₂ and A₃ sources restored male-fertility in ex-Bornu. These results confirmed the view that male-sterile ex-Bornu is a new source of cytoplasmic-genic male-sterility in bulrush millet.     

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.     

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.     

Evidence of a partial reproductive barrier between wild and cultivated pearl millets (Pennisetum glaucum)

Summary The occurrence of seed malformation in association with reduced thousand grain weight and germination ability has been observed in crosses between cultivated female plants and wild male plants. A survey of 16 cultivated accessions (P. glaucum subsp. glaucum) and 11 wild accessions (P. glaucum subsp. monodii) ranging over the whole species diversity showed this postzygotic incompatibility was general, but its intensity varied greatly with the cultivated female accession used and very little with the wild male parent origin. About 15% of the 123 cultivated x wild crosses observed gave normal seeds. Seed malformation has never been observed in crosses between cultivated accessions and appeared independent of genetic distances between the parents. The reciprocal crosses between wild female plants and cultivated male plants gave normal-looking seeds with good germination but consistently reduced thousand grain weight. Both seed malformation and seed small size are an expression of a genetic imbalance. These slight reproductive barriers seem to have been built during the domestication process.Abbreviation ICRISAT International Crop Research Institute for the Semi-Arid Tropics     

Impact of natural and human selection on the frequency of the F1 hybrid between cultivated and wild pearl millet (Pennisetum glaucum (L.) R. Br.)

In the Sahel, pearl millet yields are affected by the proportion of hybrid phenotype plants resulting from genetic mixing between domesticated and wild forms. Man counteracts this mixing by applying a production method, the efficiency of which is quantified in this study. Under experimental conditions, cultivated and wild pearl millet were hybridised in order to obtain cultivated pearl millet seeds including a known proportion of F1 hybrids tagged by two different allozymes. These seeds were sown in the field and the cultivation was conducted following practices common in the Sahel. The evolution of the survival rate of plants and the frequency of hybrids were followed over several stages during the season: sowing, germinating, emergence, thinning, flowering and maturing of the seeds. Owing to plant mortality in the experiment, the average tendency was a hybrid frequency that decreased steadily in the first part of the growing season from 42% during germination, to 37% at emergence. It then fell to 17% after the thinning of the plantlets by the farmer. At the end of the cycle, after thinning, only 11% of mature plants were hybrids. Thus, under the combined pressures of natural and human selection, the frequency of hybrids in the field declined drastically. In interaction with natural pressure, the farmer's practices of selection of seeds, sowing in pockets and thinning have the combined effect of heavily selecting the cultivated genotype and limiting without completely preventing the introgression of wild pearl millet genes into the cultivated genome. This revised version was published online in July 2006 with corrections to the Cover Date.     

Identification,characterization and geographic distribution of male-sterility restorer and maintainer lines from diverse pearl millet germplasm

Summary To determine the distribution and geographic specificity of sterility maintainers in pearl millet, Pennisetum americanum (L.) Leeke, 428 diverse pearl millet germplasm accessions representing variation from 12 countries were crossed with a male-sterile line 5141A. The F₁ hybrids were classified as male-fertile or male-sterile based on the seed set on bagged ear heads and an other morphology. Among these, 87 (20.3%) were classified as male-fertile, 32 (7.5%) as male-sterile, 282 (65.9%) as segregating for male-fertile/male-sterile and 27 (6.3%) behaved as male-fertile in the rainy and male-sterile in the postrainy season. Restorer lines were distributed in all the countries studied except Cameroon and USSR. Maintainer lines were observed from six countries but were concentrated in India. These maintainer lines differ from one another in several morphological and agronomic characters such as flowering, plant height, spike length and grain size. They may prove to be useful sources of material for generating new male-sterile lines. The restorers can be used to produce commercial hybrids.Submitted as J.A. No. 719 by the International Crops Research Institute for the Semi-Arid Topics (ICRISAT).     

Genetics of five hairy phenotypes and a linkage group of Pennisetum americanum

Summary In pearl millet hairy lamina, hairy sheath and hairy stem were inherited as monofactorial recessives while hairy leaf margin and hairy node were inherited as monogenic dominant traits. The gene for hairy lamina hl showed independent assortment from the gene for hairy node, Hn, and showed linkage with the genes hst (hairy stem), hs (hairy sheath) and Hm (hairy leaf margin). Furthermore, Hl was observed to have an epistatic effect on the expression of hs. The percentages of recombination between the gene pairs hl-hst, hl-hs and hl-Hm were 0.0, 8.30±0.44 and 19.81±0.98 respectively. Thus the genes hl-hst-hs Hm form one linkage group.     

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     

A synthetic hexaploid (2n = 42) oat from the cross of Avena strigosa (2n = 14) and domesticated A. magna (2n = 28)

A synthetic hexaploid oat was produced by chromosome doubling of a sterile triploid hybrid between cultivated Avena strigosa (2n = 14) cv. Saia and a domesticated form of A. magna (2n = 28). The synthetic hexaploid was intermediate between its parents in panicle shape and lemma color, similar to the tetraploid parent in spikelet structure, and to the diploid parent in having a single, albeit partially shriveled seed per spikelet, and low protein content. By the third generation, plants with yellowish lemmas, mostly two seeds per spikelet and better filled grains had been selected. Rust resistance of the diploid parent was retained in the synthetic hexaploid, but not tolerance to barley yellow dwarf virus disease (BYDV). Chromosome associations at meiosis in the triploid hybrid was low, with over 60% of them being univalents. Bivalent association was the rule in the synthetic hexaploid with an occasional one or two quadrivalents. Regular meiosis with 21 bivalents was observed in further generations. The preferential pairing of homologous chromosomes in the synthetic hexaploid was probably contributed by the A. strigosa genome which exhibits this tendency in artificial allopolyploid situations. Selection of yellow lemma color and two seeds per spikelet suggests that the genes controlling these traits are located on the chromosomes involved in quadrivalents in the synthetic hexaploid. The potential and limitations of utilizing the synthetic hexaploid in oat research and breeding are briefly discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

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.     

Diversity of penicillarian millets (Pennisetum glaucum and P. purpureum) as for the compatibility between their gynoecia and pollens from some other Poaceae

Summary The compatibility between gynoecia of different penicillarian millets (Pennisetum glaucum and P. purpureum) and pollens from some other Poaceae was evaluated in order to determine whether different penicillarian pistils showed the same pistil-pollen interactions with a same non-penicillarian pollen and to determine whether compatibility was dependent on the genetic divergence or the degree of sympatric evolution of the mating partners. Ten pearl millet lines (P. glaucum subsp. glaucum), six wild pearl millet accessions (P. glaucum subsp. monodii) and one P. purpureum accession were pollinated with five other Pennisetum species: P. pedicellatum, P. polystachyon, P. schweinfurthii, P. squamulatum, P. ramosum, and with three species from other genera: Cenchrus ciliaris, Panicum maximum and Zea mays. Each male species was represented by a unique accession. Pollen grain germination on the stigmata and pollen tube growth in the gynoecia were monitored by means of fluorescence microscopy after aniline blue staining. Significant compatibility differences were observed between a given pollen and gynoecia from different P. glaucum accessions. But there was no evidence of compatibility differences between the three female taxonomic groups. The eight pollen parents can be classified into three groups according to their mean degree of compatibility with the different penicillarian gynoecia. P. ramosum, P. schweinfurthii and P. squamulatum showed a high compatibility: more than 50% of penicillarian gynoecia had pollen tubes at the micropyle, six hours after pollination. Zea mays, Cenchrus ciliaris and Panicum maximum showed a low compatibility: pollen tubes were arrested in the style. P. pedicellatum and P. polystachyon showed nil compatibility: few pollen grains germinated on the stigmata and very rare tubes reached the entry of the style, where they stopped. It must be noted that such a strong gametic barrier has been observed only with pollinator species sympatric to pearl millet.     

Male fertility and cytology of triploid hybrids between tetraploidSolanum commersonii (2n=4x=48, 2EBN) and Phureja-Tuberosum haploid hybrids (2n=2x=24, 2EBN)

Summary Solanum commersonii Dun. is a diploid (2n=2x=24, 1EBN) wild species of potential value for potato breeding. It is a reproductively isolated species and cannot be crossed with Tuberosum haploids (2n=2x=24, 2EBN) or other diploid 2EBNSolanum species. In order to overcome the EBN barriers, triploid hybrids were produced between Phureja-Tuberosum haploid hybrids, which form 2n pollen grains by parallel spindles, and tetraploidS. commersonii. Microsporogenesis analysis of the triploids indicated a trend towards low values of chromosome distribution at Anaphase I; lagging chromosomes were often observed as well. Despite these abnormalities, the percentage of stainable pollen was very high, ranging from 5.0% to 74.3%. A high variation in pollen grain diameter was also evident. Parallel and tripolar orientation of spindles at Metaphase II of microsporogenesis was a common feature of all the triploids analyzed, but dyads and triads were observed at a very low frequency. Therefore, also the frequency of 2n pollen was very low; the different size of stainable pollen appears to represent the ploidy levels which are possible according to the distribution of chromosomes in Anaphase I. The results obtained also suggest thatS. commersonii could have minor genes acting at the end of meiosis in such a way that, despite the presence of parallel/tripolar spindles, dyads/triads are not formed.Contribution no. 124 from the Research Center for Vegetable Breeding.     

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).     

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.     

Analysis of reproductive isolation between pearl millet (Pennisetum glaucum (L.) R.Br.) and P. ramosum, P. schweinfurthii, P. squamulatum, Cenchrus ciliaris

Crosses between pearl millet lines and Pennisetum ramosum, P. schweinfurthii, P. squamulatum or Cenchrus ciliaris were observed for the frequency and development of zygotes, the possibility of embryo rescue, and the fertility of F1 hybrids obtained. Eight per cent of the ovules from diploid millet × P. ramosum crosses showed small embryos which could not be rescued. However, 59% of the ovules from tetraploid millet × P. ramosum crosses showed well-developed embryos that were easy to rescue 14 days after pollination. F1 hybrids were male sterile but female fertile when pollinated by diploid millet. Both diploid and tetraploid millet ovules showed the presence of hybrid zygotes after pollination with P. schweinfurthii at rates ranging from 25% to 45%. The diploid millet× P. schweinfurthii hybrid zygotes often developed almost normal seeds giving, without embryo rescue, totally sterile plants. The tetraploid millet × P. schweinfurthii hybrid embryos were normal but the endosperm was severely defective. A hybrid obtained by embryo rescue was totally sterile. A diploid millet-P. schweinfurthii amphidiploid was obtained by somatic embryogenesis associated with colchicine treatment during callogenesis. This amphiploid plant was male sterile, but gave many seeds when pollinated by a tetraploid millet and few seeds when pollinated by a diploid millet. P. squamulatum pollinating diploid millets produced proembryos with large undifferentiated endosperms in 73% of the ovules. A normal seed set was observed on tetraploid millets pollinated by P. squamulatum and the resulting F1 hybrids were partially male and female fertile. Backcrosses of these hybrids were much more fertile when pollination was from a tetraploid millet rather than from a diploid millet. C. ciliaris pollinating a diploid millet showed, in 60% of the ovules, proembryos and endosperms similar to those observed with P. squamulatum and no hybrid could be rescued. Crosses with a tetraploid millet could not be attempted due to the pistil-pollen incompatibility of tetraploid millets available with C. ciliaris. Ploidy levels of mating partners do not seem to influence pistil-pollen compatibility, but play a major role in post-zygotic abortion. With adequate ploidy levels of parents, and embryo rescue, it seems that the pearl millet gene pool can be considerably enlarged by germplasm from many other species.     

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

Summary Pearl millet (Pennisetum glaucum (L.) R. Br.) hybrids based on the A₁ cytoplasmic-nuclear male-sterile (CMS) lines are more susceptible to smut (Tolyposporium penicillariae Bref.) than open-pollinated varieties. Seventy eight pairs of hybrids, made onto male-sterile (A) lines and their counterpart maintainer (B) lines, were evaluated to examine the effects of male sterility and genetic resistance of parental lines on the smut severity of hybrids. The A-line hybrids had higher smut severity and lower selfed seedset than the counterpart B-line hybrids, indicating that it is the CMS-mediated male sterility rather than the A₁ cytoplasm per se that caused greater smut severity of A-line hybrids. However, with the use of resistant parental lines even male-sterile hybrids of A-lines, in several cases, were as resistant as some of the highly resistant male-fertile hybrids of B-lines. It would be possible to produce smut resistant hybrids (< 10% severity) on A-lines, albeit in low frequency, even if only one parent of a hybrid were resistant. However, the probability of producing such hybrids would be higher when both parents were resistant to smut. Thus, improvement in smut resistance of parental lines and fertility restoration ability of pollinators would provide the most effective genetic approach to smut disease management in hybrids.Submitted as JA No 1737 by the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT).     

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.     

The introgression of characters of the wild oat Avena magna (2n=4x=28) into the cultivated oat A. sativa (2n=6x=42)

Summary A study of derivatives from the hybrid Avena sativa × A. magna backcrossed to A. sativa clearly indicate that it is possible to introduce characters of the wild oat into the cultivated oat. Progeny with improved grain characters and crude protein content of the groat have been isolated and their use as breeding material is discussed.