Are ordinal rating scales better than percent ratings? a statistical and “psychological” view

Cytoplasmic male sterility (CMS) is considered an efficient genetic tool in pearl millet hybrid breeding. Of the several CMS sources available in pearl millet, A₁ is the only CMS widely exploited to produce commercial hybrids in India. To explore the possibility of using alternate CMS sources, we studied the cytoplasmic effects of different CMS sources on agronomic characters in pearl millet. Five CMS (A) lines representing A₁, A₂, A₃, A₄ and A₅ cytoplasms, their respective maintainer (B) lines and eight restorer (R) lines were used to generate 40 A × R and B × R experimental crosses. The experimental material was evaluated at two different locations in India. Analysis of combining ability and heterosis revealed that A₄ and A₅ cytoplasms had desirable effects for earliness. The A₅ CMS was found to be particularly promising, as compared to other CMS sources for improving grain yield. The study also indicated that the cytoplasmic effects on general combining ability (GCA) for various agronomic characters were largely non-significant. However, cytoplasmic effects on specific combining ability and heterosis were found to be modulated by cytoplasmic-nuclear interactions and influenced by the environmental conditions. The study also demonstrated the advantage of utilizing diverse male-sterile and restorer combinations in maximizing the productivity as well as for genetic and cytoplasmic diversification of hybrids in pearl millet.     

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.     

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.     

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.     

A comparative assessment of the utility of PCR-based marker systems in pearl millet

A set of 22 pearl millet inbred lines including the parents of eleven mapping populations, was screened with 627 markers including 100 pearl millet genomic SSRs (gSSRs), 60 pearl millet EST-SSRs (eSSRs), 410 intron sequence haplotypes (ISHs), and 57 exon sequence haplotypes (ESHs). In all, 267 (59%) of the markers were informative for at least one of the 11 mapping populations, which segregate for traits like drought and salinity tolerance; host plant resistance to downy mildew, rust and blast; fertility restoration and sterility and maintenance of cytoplasmic male sterility etc. An average of 116 polymorphic markers was identified per mapping population. The average PIC values and number of profiles (P) per polymorphic marker were: gSSRs (PIC = 0.62, P = 6.1), ISHs (PIC = 0.39, P = 2.6), eSSRs (PIC = 0.36, P = 3.1) and ESHs (PIC = 0.35, P = 3.1). A high correlation (r > 0.97, P < 0.05) was observed between the patterns of diversity exposed by the different marker systems. The polymorphic markers identified are suitable for the de novo construction, or the supplementation of pearl millet linkage maps. The genetic relationships identified among the panel of inbred lines may be useful in designing strategies to improve the use of available genetic variation in the context of pearl millet breeding.     

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.     

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.     

A new cytoplasmic-nuclear male sterility system in pearl millet

Among the cytoplasmic-nuclear male sterility (CMS) systems reported in pearl millet, Pennisetum glaucum (L.) R. Br., the A_m= A₄ system produces the highest frequency of male-sterile hybrids. A CMS source identified in a large-seeded gene pool (LSGP) was compared with the A₄ system. Seven diverse restorer lines of the A4 system produced hybrids with 81A4 that were all fertile (pollen-shedding score 4 and 68–89% selfed seedset). In contrast, all the hybrids of these inbreds made with the isonuclear line with the LSGP cytoplasm were sterile (pollen-shedding score 1 and 0–3% selfed seedset). Topcross hybrids of four diverse composites made with 81A4 had 10–35% plants that had good fertility (> 50% selfed seedset). In comparison, no plant of any topcross hybrid with the isonuclear line having LSGP cytoplasm exceeded 20% selfed seedset, and it was rare for a plant to exceed even 10% selfed seedset. These differential fertility restoration patterns of hybrids indicate that the LSGP cytoplasm represents a CMS system that is different from the A4 and, by implication, from all those reported to date. This new CMS system is designated A₅.     

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.     

Feasibility of breeding male-sterile populations for use in developing interpopulation hybrids of pearl millet

Inter‐population hybrids of pearl millet, Pennisetum glaucum (L.) R. Br., have a substantial grain yield advantage over open‐pollinated varieties that makes them an appropriate and economically viable proposition for many African agricultural situations, provided that stable male‐sterile populations can be developed for use as seed parents. The objective of this research was to examine the feasibility of breeding stable male‐sterile populations, using the d₂ dwarf version of Nigerian Composite NCD₂ and the A₄ cytoplasmic‐nuclear male sterility system as a test case. Results showed that two cycles of recurrent selection for sterility maintenance ability led to the development of a fully effective maintainer version of NCD₂. There was no significant difference between the original C₀ cycle bulk and the C₃ cycle bulk (developed from the third and final cycle of recurrent selection) for grain yield and other agronomic traits. The male‐sterile population at the third backcross stage, developed from the maintainer version of NCD₂, had as high a level of stable male sterility as the A¹ system commercial inbred male‐sterile line 841A₁. Thus, it is concluded that with the use of the A₄ cytoplasmic male‐sterile system, it would be possible rapidly to develop a maintainer version of any population without detrimental effects on grain yield and agronomic traits. Male sterility of populations developed from these maintainers will be highly stable, paving the way for their effective utilization as seed parents in breeding inter‐population hybrids.     

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.     

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).     

The Potential for Exploiting Variation in Salinity Tolerance in Pearl Millet (Pennisetum americanum (L.) Leeke)

Shoot and root growth and plant dry weight were determined for twenty four accessions of pearl millet, Pennisetum americanum (L.) Leeke, after two weeks growth in saline solution culture at EC 20 dS m^-1. The EC was achieved using NaCl + CaCl₂ 1:1 by weight in solution culture. Although salinity markedly inhibited growth of all accessions, three, 93613, KAT/PM-2 Kitui, and Kitui local produced significantly greater dry matter, and they and 93612 had longer shoots than the other accessions. Accessions WCA 78 and Bulk 7704 were highly sensitive to salinity in all 3 characters measured. There was considerable variation between the 24 accessions, suggesting that selection for increased tolerance to salinity in pearl millet should be possible. This was examined using normal seed of the cultivar Al/3. A first cycle of selection screened 16,000 seeds after 15 days growth in nutrient solution at EC 26 dS m^-1. 106 individuals were selected to form the S₁ selection line. Selected seedlings were grown to maturity and polycrossed. In a second cycle of selection, 30,000 S₁ polycross progeny were screened as previously, but at EC 30 dS m^-1. A selection intensity of 0.08% was achieved. The selected plants were again grown to maturity and again polycrossed. Efficiency of the selection procedure was assessed from comparison in sand culture of the performance of the selected line with the unselected Al/3 line and Kitui local which from the previous assessment of the 24 accessions was rated as relatively salinity tolerant. The selection line was superior to the other two lines at four salinity levels. The results of this study suggest that further improvement in salinity tolerance in pearl millet may be expected from further selection and breeding, using the methods described here.     

A Test of the 'Tall-Dwarf" Hypothesis in Pearl Millet, Pennisetum glaucum (L.) R. Br.

Positive height-grain yield relationships exist for many cereals, but cannot be fully used in breeding because of lodging and harvestability problems in tall cultivars. Law et al. (1978) proposed a “tall-dwarf” hypothesis for wheat, in which the positive effects of minor height genes could be exploited by selecting for them in a major dwarfing gene background. The applicability of this hypothesis to pearl millet was tested by crossing a set of dwarf S₁ progenies (from a single population) which varied in height onto two male-sterile lines. Mean (by S₁ pollinator) hybrid grain yield was closely related to mean hybrid height (r²= 0.60) over a range of mean yields of 3.0–3.9 t ha^?1 and a range of mean heights of 126–165 cm. The effect of height was expressed as an increase in grain number in one cross and as an increase in grain mass in the other, indicating the importance of background genetic effects on yield-height relationships in dwarf hybrids. The concept of “tall-dwarfs” appears to be applicable to pearl millet.     

Genetics of Five Seed Esterase Isozymes in Pearl Millet

Inheritance and linkage relationships of pearl millet seed esterase isozymes were studied using; polyacrylamide dise: gel electrophoresis and α-naphthyl acetate as substrate. The Zone of enzyme activity was resolved into five bands. The presence of a band showed complete dominance over its absence. Each one of the bands (1 to 4) was under the control of a single gene. Band five was found to be controlled by three independent loci with duplicate gent action Loci for Est₁, Est₃ and Est₄ Were found to be linked. Est₂ Was independent of this linkage group.     

Differential Competitive Ability and Growth Habit of Pearl Millet and Clusterbean Cultivars in a Mixed Cropping System in the Arid Zone of India

Dryland sustainable agriculture in the arid zone of India depends upon the choice of suitable cultivars for pure and mixed crop stands. Field experiments were conducted for two years to examine the response of two contrasting cultivars each of pearl millet (Pennisetum glaucum) and clusterbean (Cyamopsis tetragonoloba) in pure stands and in mixed pearl millet‐clusterbean stands. The differential response of cultivars of both crops to pure and mixed stands resulted in a significant genotype × cropping system interaction. Reduction in seed yield of both clusterbean cultivars was greater in mixed stands with tall and long duration pearl millet MH 179 than with medium statured and early maturing HHB 67. The degree of reduction was greater in Naveen, the branched clusterbean cultivar, than in RGC 197, the single stemmed cultivar. Mixing of pearl millet HHB 67 with medium duration clusterbean cultivar Naveen produced maximum pearl millet equivalent total yield. Higher land equivalent ratios (LERs) were also observed when clusterbean cultivars were mixed with early maturing and short statured pearl millet HHB 67.     

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.     

Effectiveness of Ethrel as a Male Gametocide in Pearl Millet and its Influence on Ergot

In field and greenhouse experiments Ethrel (2-chloroethyl phosphonic acid) was tested for its male garnetocidal effects on pearl millet (Pennisetum americanum) and its subsequent effects on ergot development. Application of Ethrel at 2000 ppm at late boot or early protogyny was the most effective for inducing male sterility in the hybrid, EJ 104. Female fertility in a male sterile line, however, was not affected by Ethrel treatment. Ethrel at 2030 ppm applied at ihe late boot stage resulted1 in partial paniele exsertion, and reduced plant height anc. panide length. In vitro Ethrel (2000 ppm) completely inhibned pollen germination but did not affect germination of conidia of Claviceps fusiformis, the causal agent of ergot of pearl millet. Ergot resistance or susceptibility in pearl millet lines was not affected., probably because Ethrel could not induce complete male sterility.     

Effects of Tillage and Cropping Systems on Soil Moisture Balance and Pearl Millet Yield

A 2-year study was conducted to determine the effects of tillage and cropping systems on soil moisture balance, growth and yield of pearl millet (Pennisetum glaucum (L.) R.Br.). Three tillage treatments, viz. minimum tillage (one harrowing), conventional tillage (two harrowing, cross) and deep tillage (ploughing followed by two har-rowings), and four cropping systems, viz. monoculture of pearl millet, pearl miliet-clusterbean (Cyamopsis tetra-gonoloba (L.) Taub.) rotation, monoculture of pearl millet with 5 t ha⁻¹ farm yard manure (FYM), and intercropping of pearl millet and clusterbean, were compared. Deep tillage improved the soil moisture storage, water use efficiency and grain yield of pearl millet while consumptive use of water was higher with minimum tillage. Total dry matter yield with deep tillage and conventional tillage was 23.2 and 10.2% higher than minimum tillage in the season 1, and the corresponding values for season 2 were 30.7 and 13.3%. The Pearl millct-clusterbean rotation and monoculture of pearl millet with the application of 5 t ha⁻¹ FYM gave 17.2 and 6.1% higher yield than monoculture of pearl millet, respectively. Maximum water use efficiency was observed in rotation followed by FYM application.     

抗丝黑穗病、抗败育高粱A2细胞质雄性不育系A2Sx3197的选育

A₁Tx3197曾经是我国广泛应用的高粱细胞质雄性不育系,20世纪70年代末,由于高粱丝黑穗病病菌生理小种分化,该不育系以及用其配制的杂交种逐渐失去了对高粱丝黑穗病菌的抗性。同时该不育系小花败育日渐严重,制种产量极低,甚至造成绝收。为了改良A₁Tx3197的抗病性及抗败育性,本研究利用A₂保持系在A₁位点含有A₁育性恢复基因MS₁MS₁和在A₁细胞质背景下表现恢复的特点,以不育系A₁Tx3197为轮回亲本,以含有抗丝黑穗病、抗败育基因的BV₄为供体,通过杂交和多代回交,得到含有双抗基因的A₂类型细胞质雄性不育的保持系BSx3197(MS₁ms₁ms₂ms₂),在该材料自交的同时,用其对A₂细胞质雄性不育系进行细胞核代换,经过多代回交和自交,最终育成了抗丝黑穗病、抗败育的A₂细胞质雄性不育系A₂Sx3197和保持系BSx3197(MS₁MS₁ms₂ms₂)。结果表明,新选育的不育系A₂Sx3197在A₁和A₅细胞质背景下表现恢复,在A₂、A₃、A₄、A₆和9E细胞质背景下表现不育,丝黑穗平均发病率为0~0.8%,败育率为0~8.4%,抗丝黑穗病性、抗败育性明显优于被改良不育系,接近或达到抗原供体BV₄水平;而在抽穗期、株高、穗长、穗宽、千粒重、穗粒重、粒色、壳色、穗形、穗型等主要性状方面与A₁Tx3197差异不显著。