Inheritance of resistance to Fusarium wilt and yield traits in pigeonpea

Fusarium wilt is the main pigeonpea production constraint in Malawi. The purpose of the study was to understand the nature and mechanism of inheritance of F. wilt resistance, yield and secondary traits in pigeonpea. 48 crosses were generated in a 12 lines × 4 testers mating scheme. Some F₁ plants were selfed for segregation analysis for inheritance pattern of resistance, while others were evaluated for resistance, yield and secondary traits. There were significant variations among F₁ plants for F. wilt, days to 50 % flowering, seed/pod, and number of secondary branches. Specific combining ability (SCA) effects were predominant for F. wilt, days to 50 % flowering and number of secondary branches. The general combining ability (GCA) effects, mainly due to maternal genotypes, were preponderant for yield and other secondary traits. The significance of GCA and SCA effects suggested that variations were due to additive gene action in both the testers and parental lines arising from their interactions, and the dominance effects due to interactions of the parental lines. The χ² analysis suggested dominant patterns of inheritance for wilt in most of the F₂ populations. The segregation ratios of 3:1, 15:1, and 9:7 suggested the involvement of single or two independent/complementary dominant genes in the test donors. Involvement of a few genes governing wilt resistance suggested the ease of breeding for this trait. Pedigree breeding method would be recommended for incorporating various traits in pigeonpea.     

Breeding high-protein pigeonpea genotypes and their agronomic and biological assessments

Proteins, inevitable for nutritional security of human beings and legumes, by far, are the cheapest source of this vital nutrient. The escalating prices and never halting population growth limit the per capita availability of protein-rich legumes. In view of limited land resource and need to grow other food crops, the greater protein harvests are possible only by increasing the protein levels of popularly grown legumes. In this context, attempts were made for raising the protein content in pigeonpea [Cajanus cajan (L.) Millsp.] through traditional plant breeding tools. For this, the high-protein trait was successfully transferred from wild relatives of pigeonpea to the cultivated types. In the derived inbred lines, the protein content was significantly enhanced from 20% - 22% to 28% - 30%. Two high-protein lines HPL 40 and HPL 8 also produced 2100 and 1660 kg/ha grain yield, respectively. This simply means that, in comparison with traditional cultivars, the cultivation of high-protein lines will provide additional 100 kg/ha of digestible protein to the farming family. This paper, besides describing the breeding procedures, also discusses the accomplishments of this breeding endeavour with respect to its various nutritional and biological properties.     

Interspecific and intergeneric hybridization and chromosomal engineering of Brassicaceae crops

In Brassicaceae crop breeding programs, wild relatives have been evaluated as genetic resources to develop new cultivars with biotic and abiotic stress resistance. This has become necessary because of the diversification of ecotypes of diseases and pests, changing food preferences, advances in production technology, the use of new approaches such as in vitro breeding programs, and the need for economical production of F₁ seed. To produce potential new cultivars, interspecific and intergeneric hybridizations have been performed between cultivated species and between cultivated species and their wild relatives. Furthermore, interspecific and intergeneric hybrids have been successfully produced using embryo rescue techniques. In this paper, we review the interspecific and intergeneric incompatibilities between Brassicaceae crops and their wild relatives, and the production, characterization, and improvement of synthetic amphidiploid lines, alien gene introgression lines, alloplasmic lines, monosomic alien chromosome addition lines, and monosomic alien chromosome substitution lines. The goal is to provide useful materials to support practical breeding strategies and to study the genetic effects of individual chromosomes on plant traits, the number of genes that control a trait, their linkage relationships, and genetic improvement in Brassicaceae crops.     

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.     

Introgression from wild Cicer reticulatum to cultivated chickpea for productivity and disease resistance

Interspecific hybridization is known to improve productivity and resistance to diseases in many crops. Therefore, an attempt was made to introgress productivity and disease resistance into chickpea from wild Cicer species. The true F₁ hybrids of cultivated chickpea genotypes ‘L550’ and ‘FGK45’ with C. reticulatum were backcrossed twice to their cultivated female parents to minimize the linkage drag of undesirable wild traits. The pedigree method was followed to advance the segregating populations from straight crosses (without backcross) and BC₁/BC₂ generations to F₅–F₇. The interspecific derivatives recorded up to a 16.9% increase over the check cultivars and a 25.2% increase over the female parent in a preliminary yield evaluation trial. Of the 22 interspecific derivatives thus derived, four desi and two kabuli lines were further evaluated for seed yield in replicated trials at three diverse locations. These lines possess a high degree of resistance to wilt, foot rot and root rot diseases, and recorded a 6.1–17.0% seed yield increase over the best check cultivars.     

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.     

Maternal and Genetic Effects on Seed Weight of Tomato, and Effects of Seed Weight on Growth of Genotypes of Tomato (Lycopersicon esculentum Mill.)

A study was made on the inheritance of seed weight of tomato, and on the effect of seed weight on growth of tomato plants. Use was made of 15 parental genotypes and 105 F₁s produced by diallel crossing (F₁-seeds). From combining ability analysis of variance carried out for weight of F₁-seeds it appeared that the weight of seeds harvested from tomato plants was mainly determined by the maternal genotype and largely independent of the genotype of the male parent. A combining ability analysis of variance for weight of seeds produced by seilfing F₁'s (F₂-seeds) showed, however, that female and male parents contributed equally to the inheritance of seed weight and that the inheritance of seed weight is determined mainly by chromosomal additiveiy acting factors, but that also non-additive gene action occurred. Genotypes with large seeds produced heavier seedlings than genotypes with small seeds, In later growth stages, the correlation between seed weight and plant weight became smaller, presumably, due to a lower relative growth rate of the seedlings from large seeds. For this reason, it is doubtful whether breeding of large seeded tomato cultivars opens up prospects of improving growth and yield of tomato plants.     

Exploitation of wild annual Cicer species for widening the gene pool of chickpea cultivars

Introgression of unadapted genes from the wild Cicer species could contribute to the widening of genetic base of important traits such as yield, yield attributes and resistance to major biotic and abiotic stresses. An attempt was made successfully to intercross two wild annual Cicer species with three cultivated chickpea cultivars. Four interspecific cross‐combinations were made, and their true hybridity was ascertained through morphological and molecular markers. These cross‐combinations were also studied for some important quantitative traits under real field conditions. The range, mean and coefficient of variation of agro‐morphological traits were assessed in the parental lines, their F₁ and F₂ generations to determine the extent of variability generated in cultivated chickpea varieties. A high level of heterosis was recorded for number of pods/plant and seed yield/plant in F₁ generation. Three cross‐combinations of ‘Pusa 1103’ × ILWC 46, ‘Pusa 256’ × ILWC 46 and ‘Pusa 256’ × ILWC 239 exhibited substantially higher variability for important yield‐related traits. The present research findings indicate that these wild annual Cicer species can be easily exploited to broaden the genetic base of cultivated gene pool for improving seed yield as well as adaptation.     

An Inheritance Study of Domestication in Foxtail Millet Using an Interspecific Cross

In order better to understand domestication processes, the inheritance of eight morphological characters and isozymes was studied in the F₂ progeny of an interspecific cross between a wild and 2 cultivated foxtail millet (Setaria viridis and S. italica). Two linkage groups were determined, one involving seed shedding and the phenol colour reaction of the grain. These two characters, as well as the pericarp pigmentation of the seed, appear to he highly specific for the difference between collections of wild and cultivated plants and probably indicate a genotype particularly suitable for plant domestication.     

Maternal inheritance of male sterility in the progeny of a natural hybrid between Cajanus lineatus and C. cajan

Adoption of pigeonpea hybrids in central and southern India is showing high impact with on‐farm yield advantages of >30%. The hybrid pigeonpea technology, the first in any legume crop, is based on a cytoplasmic‐nuclear male‐sterility (CMS) system. For a long‐term sustainability of hybrid programme, it is imperative that both nuclear diversity and cytoplasmic diversity are maintained among hybrid parents. In this context, a continuous search for new CMS‐inducing cytoplasms is necessary. This paper reports detection of maternal inheritance of male sterility in the progeny derived from a natural hybrid between a wild relative [Cajanus lineatus (W. & A.) Maesen comb. nov.] of pigeonpea and an unknown pigeonpea [Cajanus cajan (L.) Millsp.] genotype. In the present study, the male sterility was maintained up to BC₇F₁ generation by an advanced breeding pigeonpea line ICPL 99044. This male sterility inducing cytoplasm of C. lineatus was tagged as A₆. In future, this CMS genetic stock can be used to develop a range of new pigeonpea hybrids with high yield and adaptation.     

Development of Yellow Seeded Brassica napus Through Interspecific Crosses

Yellow seeded Brassica napus was developed through interspecific crosses with the two mustard species, B. juncea and B. carinata. The objective of these two interspecific crosses was the introgression of genes for yellow seed colour from the A genome of B. juncea and C genome of B. carinata into the A and C genomes of B. napus, respectively. The interspecific F₁ generations were backcrossed to B. napus in an attempt to eliminate B genome chromosomes and to improve fertility. Backcross F₂ plants of the (B. napus×B. juncea) ×B. napus cross were then crossed with backcross F₂ plants of the (B. napus×B. carinata) ×B. napus cross. The objective of this intercrossing was to combine the A and C genome yellow seeded characteristics of the two backcross populations into one genotype. The F₂ generation of the backcross F₂ intercrosses was grown in the field, plants were individually harvested and visually rated for seed colour. Ninety-one yellow seeded plants were identified among the 4858 plants inspected. This result indicated that the interspecific crossing scheme was successful in developing yellow seeded B. napus.     

Inheritance of petal colour and its independent segregation from seed colour in Brassica rapa

The inheritance of petal (flower) colour and seed colour in Brassica rapa was investigated using two creamy‐white flowered, yellow‐seeded yellow sarson (an ecotype from Indian subcontinent) lines, two yellow‐flowered, partially yellow‐seeded Canadian cultivars and one yellow‐flowered, brown‐seeded rapid cycling accession, and their F₁, F₂, F₃ and backcross populations. A joint segregation of these two characters was examined in the F₂ population. Petal colour was found to be under monogenic control, where the yellow petal colour gene is dominant over the creamy‐white petal colour gene. The seed colour was found to be under digenic control and the yellow seed colour (due to a transparent coat) genes of yellow sarson are recessive to the brown/partially yellow seed colour genes of the Canadian B. rapa cvs.‘Candle’ and ‘Tobin’. The genes governing the petal colour and seed colour are inherited independently. A distorted segregation for petal colour was found in the backcross populations of yellow sarson × F₁ crosses, but not in the reciprocal backcrosses, i.e. F₁× yellow sarson. The possible reason is discussed in the light of genetic diversity of the parental genotypes.     

Interspecific hybrids in the genus Elaeis I. crossability,cytogenetics and fertility of F1 hybrids of E. guineensis x E. oleifera

Summary The taxonomy of the genus Elaeis is reviewed. E. guineensis and E. oleifera were found to hybridize. Interspecific barriers were only partially developed as was evident from reduced seed set in the interspecific cross and occasional incomplete pairing of chromosomes in the F₁ hybrids. However vigorous F₁ hybrids were obtained and the scope for interspecific breeding work is promising.     

Introgressing resistance to bacterial and viral diseases from the middle American to Andean common bean

The genetic base of cultivars within market classes of common bean (Phaseolus vulgaris L.) is narrow. Moreover, small- and medium-seeded Middle American cultivars often possess higher yield and resistance to abiotic and biotic stresses than their large-seeded Andean counterparts. Thus, for broadening the genetic base and breeding for higher yielding multiple stress resistant Andean cultivars use of inter-gene pool populations is essential. Our objective was to determine the feasibility of introgressing resistance to Been common mosaic virus (BCMV, a potyvirus), and the common [caused by Xanthomonas campestris pv. phaseoli (Xcp) and X. campestris pv. phaseoli var. fuscans (Xcpf)] and halo [caused by Pseudomonas syringae pv. phaseolicola (Psp)] bacterial blights from the Middle American to Andean bean, using gamete selection. Also, we investigated the relative importance of the use of a landrace cultivar versus elite breeding line as the last parent making maximum genetic contribution in multiple-parent inter-gene pool crosses for breeding for resistance to diseases. Two multiple-parent crosses, namely ZARA I = Wilkinson 2 /// ‘ICA Tundama’ / ‘Edmund’ // VAX 3 / PVA 773 and ZARA II = ‘Moradillo’ /// ICA Tundama / Edmund // VAX 3 / PVA 773 were made. From the F₁ to F₅ single plant selection was practiced for resistance to the common and halo bacterial blights in both populations at Valladolid, Spain. The parents and F₅-derived F₆ breeding lines were evaluated separately for BCMV, and common and halo bacterial blights in the greenhouse at Filer and Kimberly, Idaho in 2001. They were also evaluated for the two bacterial blights, growth habit, seed color and 100-seed weight at Valladolid in 2002. All 20 F₁ plants of ZARA I were resistant or intermediate to common and halo bacterial blights in the greenhouse, but their F₂ and subsequent families segregated for both bacterial blights. Segregation for resistant, intermediate, and susceptible plants for common bacterial blight occurred in the F₁ of ZARA II. Simple correlation coefficient for common bacterial blight between the F₁ and F₁-derived F₂ families was positive (r = 0.54 P < 0.05) for ZARA II. From the F₂ to F₅ the number of families resistant to both bacterial blights decreased in both populations. Only four of 20 F₁ plants in ZARA I resulted in seven F₆ breeding lines, and only one of 32 F₁ plants in ZARA II resulted in one F₆ breeding line resistant to the three diseases. None of the selected breeding lines had seed size as large as the largest Andean parent. The use of elite breeding line or cultivar as the last parent making maximum genetic contribution to the multiple-parent inter-gene pool crosses, relatively large population size in the F₁, and simultaneous selection for plant type, seed traits as well as resistance to diseases would be crucial for introgression and pyramiding of favorable alleles and quantitative trait loci (QTL) of interest between the Andean and Middle American beans.     

Identification of Maintainer Genes in Brassica napus L. for the Male-Sterility-Inducing Cytoplasm of Diplotaxis muralis L.

Male fertility of F¹ interspecific hybrid plants derived from crosses between cytoplasmic male-sterile Brassica campestris in Diplotaxis muralis cytoplasm and 147 B. napus cultivars was Investigated. F¹, plants obtained, from crosses with the B. napus cultivars‘Mangum’and‘Hinchu’were male-sterile while F¹ plants derived from all other crosses were male-fertile. This indicated that these two cultivars carried maintainer genes far the male-sterility-inducing cytoplasm of D. muralis. Sterility was stable In plants derived from backcrosses of male-sterile F; plants with‘Mangun and‘Hinchu’but the seed set of backcross plants was low. With restorer genes readily available in B. napus, these findings could lead to the development of a new cytoplasmic male sterility system for the breeding of B. napus hybrid cultivars.     

Inter-subspecific maps of non-brittle rachis genes btr1/btr2 using occidental, oriental and wild barley lines

Brittle rachis of wild barley is controlled by two dominant complementary genes, Btr1 and Btr2, and mutation in either locus (btr1 or btr2) results in the non-brittle rachis of cultivated barley. In this study, a simple monogenic inheritance of non-brittle rachis was demonstrated, and moreover differentiation of multiple dominant alleles for either Btr1 or Btr2 among cultivated and wild barley lines was suggested. Two amplified-fragment-length polymorphism (AFLP) linkage maps of the genes were constructed using wild × btr1-type cultivar and wild × btr2-type cultivar F₂ populations. The order of AFLPs and the btr1/btr2 locus was constant between the wild × cultivar maps and a cultivar × cultivar map previously constructed. No suppression of recombination due to the inter-subspecific crosses was noticed in the interval studied. The btr1 locus and all AFLP loci were separated in the wild × btr1-type cultivar F₂ map, but the btr2 locus and eight AFLP loci did not recombine in the wild × btr2-type cultivar F₂ map, thus slightly different levels of affinity between parental cultivars with the wild line was suggested at the btr1/btr2 locus.     

Inheritance of Seed Dormancy in Lisianthus (Eustoma grandiflorum)

The genetics of chill-related seed dormancy, exhibited by a lisianthus (Eustoma grandiflorum) genotype native to North America, was investigated. Less than 1 % of the seeds germinated by direct hydration at 20°C, while full germination (98%) was attained by chilling imbibed seeds, for at least 11 days, at 3°C, before transferring them to 20°C. The inheritance of seed dormancy was analyzed in F₁, BC₁, and F₂, populations, derived from mating between a normal non-chill-requiring genotype (P₁) and the above chill-requiring genotype (P₂). F₁ seeds, of both reciprocals, and BC₁ (P₁) seeds were all non-dormant. The proportion of seed dormancy in F₂ and BC, (P₂) was 0.06 and 0.68, respectively. The inheritance model proposed includes six diallelic loci with cumulative effects, in which the presence of at least nine ‘dormancy-conferring’ alleles is necessary for inducing phenotypic seed dormancy. Possible implications of inherent seed dormancy on the utilization of native lisianthus germplasm for breeding are discussed.     

Identification and characterization of important sterile and maintainer lines from various genotypes for advanced breeding programmes of onion (Allium cepa)

Onion is one of the major vegetable crops in terms of production as well as consumption. In the current research, available onion genetic stock was evaluated to identify male-sterile lines and produce high-yielding F₁ hybrids for future breeding programmes. A mitochondrial DNA-based marker was mapped and correlated with phenotypic traits to isolate male-sterile plants. Based on the floral and pollen structure, nine putative male-sterile lines were identified. On the other hand, for nuclear marker identification at Ms locus, two sets of primers were used, one for Ms dominant allele and another for sterile and maintainer plants. Results revealed that 70% of open pollinated varieties (OPVs) possess plants with sterile cytoplasm coupled with genetic sterility at Ms locus, called sterile “A” line. Approximately 20% of plants in some genotypes were identified with normal (N) cytoplasm having recessive fertility gene at Ms locus, called maintainer “B” line. Based on the present findings, “A”, “B” and “R” (restorer line), future F₁ hybrid seed production systems in onion is discussed.     

Inheritance of the resistant reaction of the lettuce cultivar `Grand Rapids' to the southern root-knot nematode Meloidogyne incognita (Kofoid & White) Chitwood

Resistance to the southern root-knot nematode Meloidogyne incognita Chitwood would be an important attribute of lettuce Lactuca sativa L. cultivars adapted to both protected and field cultivation in tropical regions. `Grand Rapids' and a few other cultivars are reported to be resistant to this nematode. In this paper, we studied the inheritance of the resistant reaction of `Grand Rapids' (P₂) in a cross with a standard nematode-susceptible cultivar Regma-71 (P₁). F₁(Regina-71 × Grand Rapids) and F₂ seed were obtained, and inoculated along with the parental cultivars with different races of M. incognita to evaluate nematode resistance. Broad sense heritability estimates for the number of galls and of egg masses per root system, gall size and gall index were generally in the order of 0.5 or higher. Class distributions of these variables over generations P₁, P₂, F₁ and F₂ were in agreement with simulated theoretical distributions based on monogenic inheritance models. F₃ families were obtained from randomly sampled F₂ plants and tested for reaction to the nematode. The frequency ratio of homozygous resistant, segregating and homozygous susceptible F₃ families did not differ from the 1:2:1 ratio expected from monogenic inheritance. M. incognita resistance appears to be under control of a single gene locus. The Grand Rapids allele (for which the symbol Me is proposed) is responsible for the resistant reaction, and shows high (though incomplete) penetrance, variable expressivity and predominantly additive gene action. This revised version was published online in July 2006 with corrections to the Cover Date.     

Cytogenetic study of Helianthus laevigatus and its F1 and BC1F1 hybrids with cultivated sunflower, Helianthus annuus

Although the wild sunflower species Helianthus laevigatus has not been extensively studied it may be considered for sunflower breeding as a potential source of desirable genes for Sclerotinia stalk rot resistance and high contents of proteins and linoleic acid in the seed. A set of six H. laevigatus populations was crossed to cultivated sun~ower lines and produced nine F₁ (2-14 plants) and 66 BC₁F₁ hybrid combinations (1-13 plants). Male sterility occurred in F₁ and BC₁F₁ hybrid combinations and pollen viability was lower in the progenies than in the parents (51.6-77.2%in F₁ and in F₁ and 4.8-34.0% in BC₁F₁). Meiosis was normal in the H. laevigatus populations It was found that this tetraploid species also occurred in a hexaploid form Numerous irregularities were observed in the meiosis of the F₁ interspecific hybrids During diakinesis, quadrivalents and hexavalents were recorded in addition to bivalents Dislocated chromosomes and chromosome bridges were present in the other phases The chromosome number in F₁ was 68 (tetraploid). Irregularities in chromosome pairing were observed in the interspecific hybrids at BC₁F₁. There were many univalents, and trivalents quadrivalents and hexavalents were also present The chromosome number in the BC₁F₁ generation ranged from 34 to 60. The occurrence of meiotic irregularities in the F₁ and BC₁F₁ interspecific hybrids indicates that H. laevigatus and the cultivated sunflower differ in genome constitution.