Inheritance of seed protein in two interspecific pigeonpea (Cajanus cajan) crosses

Pigeonpea is one of the primary sources of protein for most rural communities of south Asia and Africa. The enhancement of protein from the present level of 20–22% will help masses in their nutritional well-beings. Breeders have identified high-protein (28–30%) germplasm amongst the wild relatives of pigeonpea; and these can serve as donors in breeding high-yielding high-protein cultivars. To achieve this, information on the inheritance of seed protein will be useful in organizing interspecific matings, generation advancement, and selection schemes. In the present study, two interspecific crosses, involving two pigeonpea cultivars ('Pant A2' and 'ICP 6997') and a wild species (Cajanus albicans (W. & A.)), were made to study the inheritance of seed protein in F₁, F₂, and BC₁F₁ generations. The high-protein trait was found to be controlled by two independent dominant genes with complementary effects. These genes are tagged with symbols “HPt₁” and “HPt₂”. The authors believe that now opportunities exist for enhancing the protein levels in pigeonpea through breeding approaches.     

A mechanism for inhibiting cross-fertilization in pigeonpea (Cajanus cajan (L.) Millsp.)

Summary Natural out-crossing imposes considerable costs and inefficiencies in breeding, evaluation and commercialization of pigeonpea (Cajanus cajan (L.) Millsp.). This note reports identification of a modification of floral morphology which inhibits cross-fertilization. Floral morphology and possible mechanisms of action of this character are discussed.On leave from International Crops Research Institute for the Semi-Arid Tropics (ICRISAT). Hyderabad, India.     

First information on heterotic groups in pigeonpea [Cajanus cajan (L.) Millsp.]

To break the decades-old yield barrier in pigeonpea [Cajanus cajan (L.) Millsp.] a hybrid breeding technology was successfully developed and the first two hybrids were recently released in India. In order to produce heterotic hybrid combinations, the first logical step is the identification and selection of genetically diverse parents with favorable alleles. In this context, the concept of classifying hybrid parents into different heterotic groups was developed and successfully used in maize and later adopted in other crops. Since hybrid technology in pigeonpea is new, the authors have made the first attempt to identify heterotic groups using SCA effects of 102 crosses generated from line × tester mating and evaluated them at four locations. Based on the performance of hybrids in terms of SCA effects, seven heterotic groups were constituted. Besides this, a scheme to use this information in breeding high yielding hybrids with specific or wide adaptation is also discussed herein. Genetic diversity between lines and tester showed positive association with the heterotic pools generated on the basis of SCA.     

Intra species variation in Atylosia scarabaeoides (L.) Benth., a wild relative of pigeonpea (Cajanus cajan (L.) Millsp.)

Summary Atylosia scarabaeoides (L.) Benth., a wild relative of pigeonpea, possesses several useful genes which can be utilized for pigeonpea improvement. In the present study, 33 accessions of A. scarabaeoides were evaluated at ICRISAT Center during the 1987 rainy season for variation in some useful traits to identify parents for inter-generic hybridization. A large variation was observed for leaf components, seed size, pod length, seeds/pod, days to flowering, seed protein, sulphur amino acids, resistance to cyst nematode, phytophthora blight, sterility mosaic, fusarium wilt, pod borer, pod fly, and pod wasp. Only four accessions were found to have more than 28% protein content. Methionine and cystine contents were marginally higher than in pigeonpea but the variation was not large enough to utilize them in the breeding program. In A. scarabaeoides. accessions resistant to fusarium wilt, phytophthora blight, sterility mosaic, and cyst nematode were detected. Compared to pigeonpea, the A. scarabaeoides accessions were less susceptible to lepidopteran borer and were immune to pod fly damage. Accessions ICPW 89 and ICPW 111 in short- (100–120 days), and ICPW 94 and ICPW 118 in medium-duration (140–180 days) were identified as potential parents for use in inter-generic hybridization.ICRISAT Journal Article No. 967     

Possibility of genetic improvement of pigeonpea (Cajanus cajan (L.)Millsp.) utilizing wild gene sources

Summary Various wild relatives of pigeonpea,Cajanus cajan, namely some species ofAtylosia andRhynchosia, possess desirable characteristics that could be utilized for effecting genetic improvement of this crop. In total 73 cross combinations among two cultivars ofC. cajan and one accession each of eightAtylosia species and one ofRhynchosia were attempted. Twelve hybrids were obtained. Seven of these were analysed for F₁ fertility and their utility for agronomic improvement of theC. cajan. Fertility behaviour of the different F₁ hybrids varied and indicated that potential of gene transfer between the two genera,Atylosia andCajanus, was as good as within the genusAtylosia. From F₂ and F₃ families ofC. cajan × A. scarabaeoides andC. cajan × A. albicans, plants were selected with greater physiological efficiency and agronomic superiority. The prospects of transferring pod borer resistance and higher seed protein content from someAtylosia species to pigeonpea are discussed.     

Microsporogenesis and anther wall development in male-sterile and fertile lines of pigeon pea (Cajanus cajan (L.) Millsp.)

Summary Microsporogenesis and anther wall development of male-sterile and fertile lines in pigeon pea (Cajanus cajan (L.) Millsp.) was examined microscopically. Male-sterility was complete and was caused by breakdown at the young tetrad stage. Degeneration of the tapetum by vacuolation occurred during the first division of meiosis and appeared responsible for pollen mother cell breakdown. Sterile plants also differed from the fertile plants in the enlargement of the inner middle layer of the anther wall, and in the lack of development of the endothecium.     

Inheritance of compact dwarf plant type in pigeonpea (Cajanus cajan)

A compact dwarf plant type TDT2004-1 isolated from the progeny of a cross involving Cajanus cajan (cv. 'C-11') and C. cajanifolia was investigated for its inheritance. Pigeonpea genotype TDT2004-1 is semi-determinate in growth habit; short (approximately 37 cm) with reduced internodal length (1.2 cm), a compact canopy and a high harvest index (40%), while genotype TT44-4 is indeterminate in growth habit; tall (132 cm) with normal internodal length (2.3 cm), an open canopy and a low harvest index (28.9%). The inheritance study involving cross TT44-4 × TDT2004-1 showed that the compact dwarf plant type was governed by a single recessive gene. TDT2004-1 could be utilized for improving the harvest index and also for restructuring the plant canopy in pigeonpea.     

Genetics of fertility restoration in A2-based cytoplasmic genetic male sterility system in pigeonpea (Cajanus cajan)

The three short duration cytoplasmic genetic male sterility (CGMS) hybrids developed using A₂ (Cajanus scarabeoides) cytoplasm-based male sterile lines (CORG 990047A, CORG 990052A and CORG 7A) and the restorer inbred AK 261322 and their segregating populations (F₂ and BC₁F₁) were subjected to the study of inheritance of fertility restoration in pigeonpea. The fertility restoration was studied based on three different criteria, namely, anther colour, pollen grain fertility and pollen grain morphology and staining. The F₂ and BC₁F₁ populations of the three crosses, namely, CORG 990047A × AK 261322, CORG 990052A × AK 261322 and CORG 7A × AK 261322, segregated in the ratio of 3:1 and 1:1, for anther colour (yellow:pale yellow), pollen grain fertility (fertile:sterile) and for pollen grain morphology and staining. The above study confirmed that the trait fertility restoration was controlled by single dominant gene. This finding can be utilized for the identification of potential restorers, which can be further used in the development of CGMS-based hybrids in pigeonpea.     

Evaluation of Efficient Weed Management Systems in Pigeonpea (Cajanus cajan L.)

Field experiments were conducted at Agricultural College and Research Institute, Tamil Nadu Agricultural University, Coimbatore to study the efficacy of different methods of weed management systems namely manual, chemical and biological in Pigeonpea during kharif and rabi 1983–84. The treatments consisted of three chemicals (fluchloralin, pendimethalin and oxadiozon alone and supplemented with one hand hoeing, manual (two hoeings at 20 & 40 DAS and two intercropping systems (Pigeonpea + sorghum and pigeonpea + moong and intercropping combined with one initial hoeing) along with unweeded check, accounting twelve treatments replicated four times in randomized block design. The results revealed that application of pendimethalin 0.50 kg ai ha⁻¹ supplemented with hand hoeing 40 DAS is found to be effective in controlling the weeds in sole pigeonpea under irrigated conditions.     

Pollen mother cell and anther wall development in a photoperiod-insensitive male-sterile mutant of pigeon pea (Cajanus cajan (L.) Millsp.)

Summary The development of the pollen mother cells (PMC's) and anther wall of a male-sterile mutant was microscopically compared with that of fertile sib plants in pigeon pea (Cajanus cajan). Male-sterility was complete and caused by breakdown of microsporogenesis at prophase I. Delayed and incomplete development of the anther wall appeared to be responsible for PMC degeneration. The sterile mutant also differed from the fertile plants in the size and number of the PMC's.On leave from International Crops Research Institute for the Semi-Arid Tropics (ICRISAT), Hyderabad, India.     

Highly variable ‘Arhar’ simple sequence repeat markers for molecular diversity and phylogenetic studies in pigeonpea [Cajanus cajan (L.) Millisp.]

Pigeonpea is an important tropical grain legume widely cultivated in South and South‐East Asia for versatile end usage as food, feed, fodder and fuel. Recent publication of draft genome sequence of pigeonpea (‘Arhar’) has allowed mining of large numbers of genomic simple sequence repeat (SSR) markers most of which are either not validated or show very little polymorphism in simple agarose gel–based assays. Here, we describe a special category of 370 validated highly variable ‘Arhar’ genomic SSR (HASSR) markers, which provide much higher level of polymorphism than a random set of SSR markers. These markers were validated for consistent amplification in a set of eight pigeonpea varieties and 152 of these (41%) showed allelic polymorphism on agarose gel electrophoresis. Twenty‐four highest polymorphic HASSR markers were used on a larger set of forty pigeonpea varieties and eight wild Cajanus species for the analysis of genetic diversity and phylogenetic relationship. Genomic HASSR markers described here are highly suitable for genetic diversity and phylogenic studies in pigeonpea.     

Random amplified polymorphic DNA analyses of cytoplasmic male sterile and male fertile Pigeonpea (Cajanus cajan (L.) Millsp.)

Randomly amplified polymorphic DNA (RAPD) markers were used for the identification of two pigeonpea cytoplasmic genic male sterile (CMS) lines derived from crosses between the wild (Cajanus scarabaeoides & C. sericeus) and the cultivated species of Cajanus cajan. The male sterile (A) line and its maintainer (B) line could be easily differentiated with certain random primers. The two male sterile (288 A and 67 A)systems are based on C. scarabaeoidesand the other is based on C. sericeus could also be differentiated. Amplification product of 600bp amplified by primer OPC-11 was observed in both the cytoplasmic male sterile lines (288 A and 67A), which was absent in the maintainer lines (288 B and 67 B) and the putative R-line (TRR 5 and TRR 6). CMS lines, putative R lines, other cultivars and wild species under the study could be easily distinguished with the help of different primers. Dendrogram constructed based on the similarity index showed that considerable genetic variation exist sbetween CMS lines, two putative R line and wild species studied. This revised version was published online in July 2006 with corrections to the Cover Date.     

Inheritance of resistance to two isolates of sterility mosaic pathogen in pigeonpea (Cajanus cajan (L.) Millsp.)

The resistant reaction to tomato spotted wilt virus (TSWV) was found to be determined by a single dominant gene in three Capsicum chinense Jacq. accessions (‘PI 152225’, ‘PI 159236’, ‘7204’). Allelism studies indicated that all C. chinense lines bear the same allele located at the Tsw locus. All the inoculated plants in the allelism tests displayed a resistant hypersensitive phenotype characterized by necrotic local lesions followed by abscission of the inoculated organ. However, a small proportion of them showed late systemic infection. Nine TSWV isolates obtained from these individual plants with systemic symptoms were backinoculated to the three resistant parents. All isolates were able to infect systemically all the resistant accessions without inducing local necrotic lesions. Serological analysis confirmed that these nine viral isolates belong to the TSWV species (serogroup I). Consequently, the susceptible plants in the allelism tests could not be interpreted as possessing a recombinant genotype because of the virulence change in the viral strain. Hobbs et al. (1994) already reported the existence of TSWV pathotypes overcoming the resistance of C. chinense resistant accessions. Practical consequences for pepper breeding associated with the emergence of these resistance-breaking isolates are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Nitrogen Requirements at Different Growth Stages of Short-duration Pigeonpea (Cajanus cajan L. Millsp)1

The response to N fertilization of a short-duration pigeonpea genotype, ICPL 87, was studied in the field to assess the scope for genetically improving symbiotic N₂ fixation by pigeonpea. The field study was undertaken during 1985, 1986 and 1987 growing seasons on Vertisol and Alfisol at ICRISAT Center (peninsular India), Inceptisol at Gwalior (central India) and Entisol at Hisar (northern India) in as non-limiting environmental conditions as possible. Nitrogen fertilizer was applied to the soil at various growth stages to determine when N becomes most limiting. There was a significant response in grain yield to fertilizer N applied at flowering in Vertisol but not in Alfisol, Inceptisol or Entisol. This suggests that biological N₂ fixation by short-duration pigeonpea was not adequate to meet N requirements of the crop grown in Vertisol but that it was probably adequate in the other three soil types. These results are discussed in relation to the nodulation and acetylene reductase activity of pigeonpea and also N mineralization potential of different soils. It can be concluded that there is a need for genetic improvement of N₂ fixing ability of short-duration pigeonpea grown on heavy textured soils such as Vertisols.     

Chloroplast-DNA variation in the wild and cultivated olives (Olea europaea L.) of Morocco

The male sterile plants that segregated in a BC₅F₂ of `C. sericeus × C. cajan var. TT-5' population were maintained by sib mating. The male sterile plants were crossed with ICPL-85012.Approximately 50% of the F₁ plants were sterile. F₂ plants derived from the fertile F₁ plants did not segregate for male sterility. The reciprocal hybrid i.e. ICPL-85012 × Fertile derivatives from C. sericeus × TT-5, did not express male sterility. However, among the 12 F₂ plant to row progenies, two segregated 25% male sterile plants and remaining 10 did not segregate. The segregation pattern in subsequent progenies revealed that the sterility was under control of a single recessive allele. Studies on the backcross and their BC₁F₂ and BC₁F₃progenies revealed another sterility gene which was found to be dominant in inheritance. This paper shows that what was thought to be cytoplasmic male sterility from C. sericeus cytoplasm is actually a single dominant gene possibly acting in concert with a single recessive gene to mimic cytoplasmic male sterility. This revised version was published online in July 2006 with corrections to the Cover Date.     

A possible gene source for early,day-length neutral pigeon peas,Cajanus cajan (L.) Millsp.

Summary Atylosia scarabaeoides Benth. and A. platycarpa Benth., close relatives of the species Cajanus cajan (L.) Millsp., were screened for photoperiodic response. Four photoperiods ranging from 12 h 45 min to 19 h were studied in three environments. A. scarabaeoides flowered freely only in the first photoperiod. A. platycarpa not only flowered early (39 to 63 days after planting) in all four photoperiods, but also exhibited a relatively constant vegetative phase up to 16 h of illumination. Cajanus cajan, in which most cultivars-if not all-exhibit a response to photoperiod, was successfully hybridized with the two Atylosia species. It is suggested that genes for earliness and insensitivity to day-length could be transferred from A. platycarpa to C. cajan.     

Inheritance of stem termination in pigeon pea (Cajanus cajan (L.) Mill sp.)

Summary Inheritance studies on the stem termination in pigeon pea using F₁, F₂ and F₃ generations of two crosses between determinate and indeterminate lines suggested that two dominant genes with epistatic (inhibitory) interaction of one of them control the interminate growth habit. The gene symbols D. idid and ddIdId have been designated to the parental plants with determinate and indeterminate growth habits, respectively. The gene IdId was epistatic (inhibitory) to the gene D giving a ratio of 13 indeterminate: 3 determinate inthe F₂'s observed. F₃ segregation supported the proposed model on the mode of inheritance.     

Role of Arbuscular Mycorrhizae in the Alleviation of Ionic, Osmotic and Oxidative Stresses Induced by Salinity in Cajanus cajan (L.) Millsp. (pigeonpea)

Salinity stress causes ion toxicity and osmotic imbalances, leading to oxidative stress in plants. Arbuscular mycorrhizae (AM) are considered bio‐ameliorators of saline soils and could develop salinity tolerance in crop plants. Pigeonpea exhibits strong mycorrhizal development and has a high mycorrhizal dependency. The role of AM in enhancing salt tolerance of pigeonpea in terms of shoot and root dry weights, phosphorus and nitrogen contents, K⁺ : Na⁺, Ca²⁺ : Na⁺ ratios, lipid peroxidation, compatible solutes (proline and glycine betaine) and antioxidant enzyme activities was examined. Plants were grown and maintained at three levels of salt (4, 6 and 8 dSm^?1). Stress impeded the growth of plants, led to weight gain reductions in shoots as well as roots and hindered phosphorus and nitrogen uptake. However, salt‐stressed mycorrhizal plants produced greater root and shoot biomass, had higher phosphorus and nitrogen content than the corresponding uninoculated stressed plants. Salt stress resulted in higher lipid peroxidation and membrane stability was reduced in non‐AM plants. The presence of fungal endophyte significantly reduced lipid peroxidation and membrane damage caused by salt stress. AM plants maintained higher K⁺ : Na⁺ and Ca²⁺ : Na⁺ ratios than non‐AM plants under stressed and unstressed conditions. Salinity induced the accumulation of both proline and glycine betaine in AM and non‐AM plants. The quantum of increase in synthesis and accumulation of osmolytes was higher in mycorrhizal plants. Antioxidant enzyme activities increased significantly with salinity in both mycorrhizal and non‐mycorrhizal plants. In conclusion, pigeonpea plants responded to an increased ion influx in their cells by increasing the osmolyte synthesis and accumulation under salt stress, which further increased with AM inoculation and helped in maintaining the osmotic balance. Increase in the antioxidant enzyme activities in AM plants under salt stress could be involved in the beneficial effects of mycorrhizal colonization.     

An overview of male-sterility systems in pigeonpea [<Emphasis Type="Italic">Cajanus cajan</Emphasis> (L.) Millsp.]

For commercial development of hybrids the four pre-requisites are; availability of perfect male-sterility system, efficient mass pollen transfer mechanism, hybrid vigor, and the large scale seed production of hybrids for commercialization. The type of male-sterility governs the acceptance of hybrids by farmers. Genetic male-sterility (GMS) system was not accepted by farmers due to the economics of large scale seed production. The major drawback was rouging of fertile counterpart from the female plot, which was time consuming and labor intensive. Cytoplasmic-genic male-sterility (CMS) system usually was a better option for large scale seed production. Hybrid vigor has been utilized in some cereal and vegetable crops. Pigeonpea (Cajanus cajan) displays considerable natural out-crossing and now CMS lines are available with different cytoplasmic backgrounds. This mini-review reports the research on development of CMS lines and CMS-based hybrids in pigeonpea.