A bottleneck in lentil: widening its genetic base in South Asia

This paper introduces the concept of a bottleneck in lentil (Lens culinaris Medikus) in South Asia and then reviews its rupture. South Asia is the largest lentil growing region in the world and where indigenous lentils show a marked lack of variability. This results from its introduction from Afghanistan around 2000 B.C. and it limits breeding progress. Three approaches to widening the genetic base in the region have been tried, namely plant introduction, hybridization and mutation breeding. Introductions from West Asia flower as indigenous material matures. The asynchrony in flowering has isolated the local pilosae ecotype reproductively. However, the introduction of ILL 4605, an early, large-seeded line, has resulted in its release as ‘Manserha 89’ for wetter areas of Pakistan and its widespread use as a parent in breeding programs in the region. Hybridization between pilosae and exotic germplasm, primarily at International Center for Agricultural Research in the Dry Areas (ICARDA) followed by selection in the sub-continent has resulted in cultivars with improved disease resistance and yield in Bangladesh and Pakistan. Mutation breeding has given new morphological markers and several promising lines. These examples illustrate not only the widening of the genetic base of the lentil in South Asia, but also the evolution of a breeding program of an international center and national programs targeted toward specific adaptation. This revised version was published online in July 2006 with corrections to the Cover Date.     

Salinity Effects on the Growth and Ion Contents of Some Chickpea (Cicer arietinum L.) and Lentil (Lens culinaris Medic.) Varieties

The effect of salinity on seed germination, plant yield parameters, and plant Na, Cl and K concentrations of chickpea and lentil varieties was studied. Results showed that in both crops percentage emergence was significantly reduced by increasing NaCl levels (0–8dSm^?1). From the plant growth studies it was found that differences existed among chickpea and lentil varieties in their response to NaCl application. In chickpea, the variety Mariye showed the comparatively lowest germination percentage and the lowest seedling shoot dry weight in response to salinity and was also among the two varieties which had the lowest relative plant height, shoot and root dry weight and grain yield at maturity. Similarly, variety DZ-10-16-2, which was the second best in germination percentage and the highest in terms of seedling shoot dry weight, also had the highest relative plant height, shoot and root dry weights, and grain yield at maturity. In lentil, however, such relationships were less pronounced. Chloride concentration (mg g^?1) in the plant parts at salt levels other than the control was about 2–5 times that of Na. K concentration in the plants was significantly reduced by increasing NaCl levels. Chickpea was generally more sensitive to NaCl salinity than lentil. While no seeds were produced at salinity levels beyond 2dSm^?1 in chickpea (no seeds were produced at this salt level in the most sensitive variety, Mariye), most lentil varieties could produce some seeds up to the highest level of NaCl application. Overall, varieties R-186 (lentil) and Mariye (chickpea) were the most sensitive of all varieties. On the other hand, lentil variety NEL-2704 and chickpea variety DZ-10-16-2 gave comparatively higher mean relative shoot and root dry weights, and grain yield, thus showing some degree of superiority over the others. The observed variations among the varieties may be useful indications for screening varieties of both crops for salt tolerance.     

Linkage between three morphological markers in lentil

Thirteen monogenic morphological marker traits of lentil were analysed for their joint inheritance in 25 crosses. Only three traits showed linkage: Ert (for spreading-erect growth habit), Gs (for brown-green stem) and Bl (for brown-green leaf). The recombination frequency between Bl and Gs was 14%, between Bl and Ert 34%, and between Gs and Ert 38%. Thus, the gene Bl is located between Ert and Gs.     

The use of Rapd markers for lentil genetic mapping and the evaluation of distorted F2 segregation

To maximize the extent of polymorphism within a mapping population wide crosses are often made, frequently resulting in distorted segregation. Two parents used in the crosses in this study contained ca 50% from wild lentil genome (Lens culinaris ssp. Orientalis). We investigated the use of random amplified polymorphic DNA (RAPD) in the lentil (Lens culinaris Med.), for genetic mapping and testing for segregation distortion in F₂ populations. In cross 1, 83% of the RAPD markers showed segregation distortion, which was also observed for isozyme and morphological loci. By contrast, in cross 2, there was little (10%) segregation distortion. Out of 390 primers tested, 116 primers (29.7%) yielded 192 polymorphic fragments between parents of cross 2. This polymorphism was confirmed as reproducible. Seventy-eight segregating loci were analyzed for linkage, at a LOD score > 3.0, resulted in 28 RAPD, one RFLP, one morphological and three oligonucleotide markers, which were assigned to 9 linkage groups spanning 206 cM. Clearly, in lentil RAPD markers were valuable for genetic mapping and evaluation of segregation distortion. This revised version was published online in July 2006 with corrections to the Cover Date.     

Heritability estimates for winter hardiness in lentil under natural and controlled conditions

Heritability analysis for cold tolerance in lentil was conducted using parental, F₂ and F₃ populations at two locations in Balochistan, Pakistan and one controlled environment in Fort Collins, Colorado, USA. Populations of parental and F₂ families were grown over 2 years (1991‐92 and 1992‐93) at Quetta, Balochistan, Pakistan. In 1992‐93, parental and F₃ families were studied at Quetta, and Kalat, Balochistan, Pakistan. Evaluation for normality using non‐transformed and log‐transformed data failed. Attempts to use analysis of variance were then abandoned in favour of parent‐offspring regression for narrow‐sense heritability. Estimates of narrow‐sense heritability ranged from 0.31±0.06 to 0.71 ± 0.06 under field conditions. Under controlled conditions, the estimated heritability was maximized at 1.00 ± 0.17 using 6‐ to 8‐week‐old lentils. Significant transgressive segregants were found in five of the six populations in the F₃ generation. Transgressive segregants appeared in the controlled F₃ generation but were not observed in field environments. This indicates that cold tolerance is under additive gene control and is environmentally sensitive in gene expression.     

Seed Size and Water Potential Effects on Water Uptake, Germination and Growth of Lentil

Seed size is an important parameter for plant growth and yield. The effects of seed size and water potential on seed water uptake, germination and early growth of lentil ( Lens culinaris Medik. cvs. Jor-1 and Jor-2) were investigated. Rate of water uptake by seed size (small, medium and large) from solutions containing different water potentials (0, –0.5 and –1.0 MPa, as polyethylene gly-col-8000) was higher in large than in medium or small seeds of both cultivars, regardless of water potential. Rate of water taken into seeds was higher in Jor-2 than in Jor-1. Decreasing water potential (more stress) had adverse effects on rate of water uptake by seeds in both cultivars. In another experiment, with lentils grown in a greenhouse at different soil metric potentials (–0.03, –0.15 and –0.30 MPa), seed size or cultivar had no effects on germination percentage (GP), but GP was reduced as soil water potential decreased (more stress), in greenhouse soil, shoot dry matter (SDM), root dry matter (RDM), plant height, total root length (TRL) and number of primary branches per plant of 35-day-old plants from large seeds were larger than those of plants from medium and small seeds of both cultivars. Increasing soil water deficit progressively decreased each of these traits. Plants from large seeds had higher SDM, RDM and TRL than those from small seeds at intermediate soil water potential (–0.15 MPa) in comparison with the control (–0.03 MPa) or severe (–0.30 MPa) soil water potentials. Larger seeds produced larger plants than smaller seeds, and this appeared to be more pronounced under intermediate than well-watered or more severe water-stressed conditions. Faster early growth of plants from larger seeds may be advantageous in establishing plants under semiarid conditions.     

Attempts to overcome postfertilization barrier in interspecific crosses of the genus Lens

Wild relatives are a potential source of genetic diversity to lentil (Lens culinaris Medik). The objective of this research was to obtain viable interspecific hybrids between the domesticated lentil and its wild relatives. The paper details the results of a number of interspecific crosses among L. culinaris, L. orientalis, L. odemensis, L. ervoides and L. nigricans. Viable hybrids were produced between L. culinaris × L. orientalis, L. culinaris × L. nigricans, L. culinaris × L. ervoides and between L. culinaris × L. odemensis. Further viable hybrids were obtained between L. culinaris and L. ervoides, which have the potential to be a ‘bridge’ in hybridization to L. culinaris for specific L. nicrigans lines which proved recalcitrant in L. culinaris × L. nigricans crosses. This is the first time that four wild species of lentils have been used successfully in hybridization with cultivated lentils, and viable hybrids produced. This paper also suggests that the artificial supplement of GA₃, hormone is needed after fertilization for the normal growth of the hybrid embryo, possibly as the natural GA₃ production is restricted with alien pollinations in cultivated lentils in both F₁ and backcross hybrids.     

Restoration of meiotic regularity and fertility among derivatives of Lens culinaris x L. Nigricans hybrids

Summary The wild lentil, L. nigricans is cross compatible with the cultivated species. Chromosome pairing in their F₁ hybrids is highly irregular due to a difference of three translocations between the parental species. However, the hybrids are partially fertile and in the F₂ populations there is a marked improvement in fertility. About 19% of the F₂ plants were meiotically stable, fully fertile and their chromosome arrangement was similar to that of the cultivated species. All these are clear indications for the possibility of rapid introgression between the two species and hence systematic screening of L. nigricans for traits of economic value is fully justifiable.     

Diversity analysis and species identification in Lens using PCR generated markers

Using random amplified polymorphic DNA (RAPD) analysis we assessed the genetic relationships between 16 accessions and cultivars of lentil (Lens culinaris ssp. culinaris) in the Australian lentil breeding program. All lines exhibited polymorphism with a maximum dissimilarity value of 0.36. This indicated a limited degree of genetic variation. Polymerase chain reaction (PCR) with primers based on the flanking regions of the 5S rRNA gene from Pisum sativum amplified the non-translated spacer (NTS) region from within the 5S rRNA gene of Lens. Three distinct amplification banding patterns differentiated between restricted genomic DNA of Lens spp. L. culinaris ssp. culinaris and L. culinaris ssp. orientalis shared similar markers of two distinctly different NTS sizes. L. nigricans and L. odemensis shared the same amplification pattern of a single sized NTS region. However, L. ervoides contained two separate sizes of NTS, distinct from other Lens species. In an effort to widen the genetic base of cultivated lentil, these species-specific molecular markers may be used to follow potential introgression between species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effect of Sowing Date on the Yield of Lentil (Lens culinaris Medik.)

A sowing date experiment was conducted over 6 years (1978–1983) to determine the optimum sowing date for lentil on vertisols in Debre Zeit area, Ethiopia. The results showed that sowing between the last week of June and the second week of July maximizes lentil yield. Later planting after mid-July, reduced both yield and time to maturity.     

Relationships among economic characters in lentil

Summary There are several reports of the association among lentil characters based on a few, relatively similar lines grown in single environments. Their results have often been inconsistent because of inadequate sampling of genotypes and environments. This study examined associations between economic characters in lentil over a wide range of genetic material and environments in West Asia. The first part of the study concerned correlations within large samples from the world lentil collection grown in two seasons, and the second part covered a smaller sample of genetic material from the collection sown over a wider range of conditions.The results showed similar phenotypic correlations over two seasons, contrasting in rainfall, in the world lentil collection. Similar genetic and phenotypic correlations were also shown by the smaller sample of germplasm over 10 environments. These results indicated low importance of covariance due to environmental and genotype-environment interaction effects. Seed yield was positively correlated with straw yield, indicating selection for either character will increase the other trait. The correlation between seed protein content and seed yield was small and negative, while the correlation of protein with straw yield was positive in sign. This suggests that the current ICARDA goals of selection for high seed and straw yield will not have a major correlated effect on seed protein content. There was a strong positive correlation between seed size and cooking time (r=0.96), therefore seed size can be used to predict cooking time.     

Variation between and within lentil landraces from Yemen Arab Republic

Summary The variation in 31 landraces of lentil collected from Yemen Arab Republic was partitioned to estimate the variation between populations and the variation between families within populations by studying a total of 568 progenies of randomly selected plants. There was significant variation between populations and between families within populations in seedling pigmentation, time to flower, time to maturity and plant height, but the predominant source of variation was between populations. This variation was unrelated to altitude of collection. The landraces were equally variable across characters. The implications of this analysis of variation on lentil breeding and germplasm collection are discussed.     

Genetics of Resistance to Four Races of Xanthomonas campestris pv. oryzae in Some Rice Cultivars

Inheritance of resistance to four Philippine races of bacterial Might caused by Xanthomonas campestris pv. oryzae was investigated in four cultivars of rice, Oryza sativa L. Resistance to three races in ‘Benamuri’ and ‘Aus 192’ is governed by xa-5. In ‘Tepal Boro’ and ‘Bazail 975′, resistance to races 1, 2. and 3 is conferred by xa-5, but another recessive gene confers resistance to race 4. This recessive gene is closely linked to xa-5 and may be allelic to xa-13. Rice cultivars with xa-13 are resistant to prevalent races of bacteria] blight in the Indian subcontinent and should thus prove useful as donors for resistance to bacterial blight in rice breeding programs.     

Geographic distribution of variation in quantitative traits in a world lentil collection

Summary In a world lentil collection the distribution of variation amongst accessions from 13 major lentil-producing countries was examined on the basis of nine quantitative morphological characters by discriminant analysis and canonical analysis. Stepwise discriminant analysis revealed major differences between accessions from different countries. Three major regional groups were apparent: 1) a levantine group (Egypt, Jordan, Lebanon and Syria, 2) a more northern group composed of Greece, Iran, Turkey, and USSR, and 3) accessions from India and Ethiopia with strikingly similar quantitative morphological characters. Misclassifications of individuals within groups were frequent. Characters useful in discriminating between accessions from different countries were in descending order of importance: time to maturity, lowest pod height and 100-seed weight. The regional grouping indicates the importance of local adaptation through clusters of associated characters with phenological adaptation to the ecological environment as the major evolutionary force in the species.     

Selection for pod retention and pod indehiscence in lentils

Summary The genetic variation for seed yield losses and the response to selection for reduction of such losses was studied by a six-week delay in the timing of harvest in various segregating populations from four lentil crosses. The bulk segregating populations had previously been subjected to different numbers of selection by means of a delayed harvest. The loss in seed yield from a delayed harvest in the two seasons 1981/82 and 1982/83 accounted for 551 and 105 kg/ha seed, representing 34 and 11% of the yield from a correctly timed harvest respectively. Pod drop accounted for 65% of this loss, whereas dehiscence gave 35% of the loss in both seasons. The parents of cross 4 differed significantly for pod dehiscence, with genotype 74TA 550 showing relative indehiscence. Selection by means of a delayed harvest of bulk populations decreased pod dehiscence, mean seed weight and bulk mean yield.     

An Assessment of Variation for Nutritional and Non-nutritional Carbohydrates in Lentil Seeds (Lens culinaris)

The levels of the raffinose family of α-galactosides (raffinose, stachyose, verbascose and ciceritol) and of the nutritional carbohydrates (fructose, sucrose and starch) were determined in the seed of a range of lentil lines from germplasm collections and commercial Spanish cultivars. Significant levels of variation were found for all the measured characters, and particularly for the α-galactosides. The greatest variation was found in verbascose content, which ranged from about 1 % of the seed dry weight to undetectable amounts. The relationship between these chemical components and morphological seed characteristics was assessed. The only significant relationship between the different groups of characteristics was a positive correlation between the level of verbascose and the average seed weight for each genotype.