Ineffectiveness of wrapped flower in inhibiting cross-fertilization in pigeonpea

Summary The wrapped flower character has been reported to inhibit out-crossing in pigeonpea in Australia. However, one of the wrapped flower lines gave an average of about 16% out-crossing at ICRISAT Center near Hyderabad, India. This level of out-crossing was not different from that in cultivars having normal (unwrapped) flowers. This suggests that the wrapped flower trait is not a dependable character for reducing out-crossing in pigeonpea.ICRISAT Journal Article No. 578.     

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.     

RFLP analysis of cytoplasmic male-sterile lines of pigeonpea [Cajanus cajan (L.) Millsp.] developed by interspecific crosses

Total DNA from three putative cytoplasmic male sterile (CMS) progenies derived from crosses between the wild species Cajanus sericeus and the cultivated species Cajanus cajan, five C. cajan, one accession of C. sericeus and two genetic male sterile lines of pigeonpea were compared for their RFLP patterns using maize mitochondrial DNA (mtDNA) specific probes. Three putative cytoplasmic male sterile (CMS) progenies from the multiple cross genome transfer of pigeonpea lines (CMS 7–1, CMS 12–3, and CMS 33–1) showed hybridization patterns identical to that of C. sericeus when DNA was digested with EcoRI and HindIII and probed with maize mtDNA clones. The results suggested that these putative CMS progenies have the mitochondria of the female wild species parent. The hybridization patterns of the three male parental lines used in the development of the CMS progenies were similar in all the restriction enzyme-probe combinations except HindIII-atp6. The genetic male sterile lines, MS Prabhat and QMS 1 differed from each other in their hybridization pattern. The genomic DNA hybridization pattern of HindIII digested DNA from ICPL 87 differed from the other pigeonpea lines when probed with the maize mtDNA clones. The cluster analysis of the hybridization data suggested the occurrence of variation in the mitochondrial genome even among the cultivated species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Introgression of Cajanus platycarpus genome into cultivated pigeonpea, C. cajan

Summary Cajanus platycarpus, an incompatible wild species from the tertiary gene pool of pigeonpea (C. cajan (L.) Millspaugh), has many desirable characteristics for the improvement of cultivated varieties. To necessitate such transfers, embryo rescue techniques were used to obtain F₁ hybrids. The F₁ hybrids were treated with colchicine to obtain tetraploid hybrids, that were selfed to obtain F₂, F₃ and F₄ progenies. All of the hybrids and subsequent progenies had an intermediate morphology between the two parents. Backcrossing of the tetraploid hybrids with cultivated pigeonpea was not possible given embryo abortion, with smaller aborted embryos than those obtained in the F₀ parental cross.As a route of introgression, diploid F₁ hybrids were backcrossed with cultivated pigeonpea and BC₁ progeny obtained by in vitro culture of aborting embryos. BC₂ plants were obtained by normal, mature seed germination. Although embryo rescue techniques had to be used to obtain F₁ and BC₁ plants, it was possible to produce BC₂ and subsequent generations through direct mature seed. Every backcross to cultivated pigeonpea increased pollen fertility and the formation of mature seeds.Special project assistant till December, 2003.     

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.     

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.     

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.     

Role of partial cleistogamy in maintaining genetic purity of pigeonpea

Summary Natural out-crossing in pigeonpea (Cajanus cajan (L.) Millsp.) leads to genetic deterioration and in certain environments the out-crossing may be as high as 70%. Studies with partially cleistogamous lines derived from an inter-generic cross indicated that average natural out-crossing in these lines was much lower (2.5%) than that reported earlier for pigeonpea (36.5%) at ICRISAT Center. Based on seed yield and disease resistance, potential parents such as ICPL 87154 in short-duration determinate, ICPL 87018 in short-duration indeterminate, and ICPL 87159 in medium-duration indeterminate groups have been identified for introgression of the partially cleistogamous trait into pigeonpea.     

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.     

Variation for natural out-crossing in pigeonpea

Summary Several researchers have reported a considerable degree of natural out-crossing in pigeonpea from different environments. This paper reviews the subject with respect to the variation for natural out-crossability, pollinating vectors, extent of natural out-crossing, isolation specifications, and the possible utilization of natural out-crossing in pigeonpea improvement.ICRISAT Journal Article No. 831.     

Genetics of a high-selfing trait in pigeonpea

Summary A true-breeding line characterized by free filaments of anthers and modified keel petal was derived from F₂ population of an intergeneric cross between Cajanus cajan (L.) Millsp. and Atylosia lineata W. & A. This variant, designated as partial cleistogamy, favours a high level of self-fertilization. The inheritance of this trait was studied in the F₁, F₂, F₃, and BC₁F₁ generations of three crosses. The results suggest that the partial cleistogamy trait is governed by a single recessive gene, designated as pct.     

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.     

Natural out-crossing in dwarf pigeonpea

Summary Natural out-crossing rate in pigeonpea was studied at ICRISAT Center using plant stature (tall plants in dwarf progenies) as the genetic marker. The data indicated natural out-crossing rates of 9.7% to 24.1% with a pooled value of 13.1% in the six populations studied. These data were comparable to earlier studies at the same site using stem colour and growth habit as genetic markers in tall pigeonpea cultivars thus suggesting that foraging of insect pollination vectors is not influenced by plant type. The implications of natural out-crossing on breeding and maintenance of genetic purity of cultivars is discussed.     

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.     

Physico-chemical traits of Cajanus cajan (L.) Millsp. pod wall affecting Melanagromyza obtusa (Malloch) damage

Pigeonpea [Cajanus cajan (L.) Millsp.] is an important legume crop in the semi-arid tropics, and pod fly [Melanagromyza obtusa (Malloch)] is an important emerging constraint to increase the production and productivity of this crop under subsistence farming conditions. Host plant resistance can be used as an important tool for the management of this pest. Therefore, a set of ten pigeonpea genotypes from a diverse array of plant growth types and maturity groups including two appropriate commercial checks, was evaluated for resistance to pod fly under field conditions, and characterized for physico-chemical pod traits. The non-determinate type GP 75 (extra early maturing) and GP 118 (early maturing), and determinate type GP 233 (extra early maturing) and GP 253 (early maturing) genotypes had significantly lower pod and seed damage as compared to determinate (Prabhat) and non-determinate (Manak) early maturing checks, suggesting that resistance to pod fly is not linked to plant growth type and maturity period of the genotype in pigeonpea. Pod wall thickness, trichome density, reducing and non-reducing sugars, total phenols, tannins, and crude fiber were found to be negatively associated (r = −0.83** to −0.97**), while total protein positively associated (r = 0.88** to 0.97**) with pod fly infestation. Therefore, these traits particularly total phenols, tannins, crude fiber, trichome density, and pod wall thickness, can be used as physico-chemical markers to identify pigeonpea genotypes with resistance to M. obtusa, and use in pod fly resistant breeding program 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.     

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.     

Production of hybrids between Cajanus acutifolius and C. cajan

There are many wild species of pigeonpea which are endemic to Australia. These wild species are cross incompatible with cultivated species of Indian origin. Cajanus acutifolius is one such species which does not easily cross with cultivated pigeonpea. Interspecific pollinations lead to hybrid seeds which were semi-shrivelled. Very few seeds germinated to give rise to F1 plants. Backcrossing the hybrid plants to C. cajan, the male parent, gave rise to aborting seeds which did not germinate in vivo hence BC1 plants are obtained after saving the aborting embryos in vitro. BC₁ plants showed normal meiotic pairing, but had low pollen fertility. The reasons for embryo abortion and low pollen fertility in spite of normal meiosis could be due to the effect of wild species cytoplasm. This revised version was published online in August 2006 with corrections to the Cover Date.     

A cytoplasmic-nuclear male-sterility system derived from a cross between <Emphasis Type="Italic">Cajanus cajanifolius</Emphasis> and <Emphasis Type="Italic">Cajanus cajan</Emphasis>

Cytoplasmic-nuclear male-sterility is an important biological tool, which has been used by plant breeders to increase yields in cross-pollinated cereals and vegetables by commercial exploitation of the phenomenon of hybrid vigor. In legumes, no such example exists due to the absence of an economic way of mass pollen transfer from male to female parent. Pigeonpea [Cajanus cajan (L.) Millsp.], however, is a different legume where a moderate level of insect-aided natural out-crossing (25–70%) exists and it can be used to produce commercial hybrid cultivars, if an efficient and stable cytoplasmic-nuclear male-sterility (CMS) system is available. This paper reports the development of a stable CMS system (ICP 2039A), derived from an inter-specific hybrid of Cajanus cajanifolius, a wild relative of pigeonpea, with a cultivar ICP 11501. Using this genetic material, designated as the A₄ cytoplasm, a number of fertility restorers and maintainers have been developed. The best short-duration experimental pigeonpea hybrid ICPH 2470 produced 3205 kg ha⁻¹ grain yield in 125 days, exhibiting 77.5% advantage over the control cultivar UPAS 120. At present, all the important biological systems necessary for a successful commercial hybrid breeding program are available in pigeonpea and the package of this technology has been adopted by private seed sector in India for the production and marketing of hybrid varieties.