Adaptation and yield stability of selected lines of Lathyrus spp. under rainfed conditions in West Asia

Summary Sixteen promising lines (selections) of Lathyrus spp. (chicklings), comprising 11 L. sativus, four L. ochrus, and one L. cicera, were evaluated under rainfed conditions in Syria. Two locations in each of four years were treated as separate environments, to give eight environments altogether. Genotype x environment (G×E) interactions were analysed using linear regression.There was considerable variation in herbage and seed yields within both lines and environments. Genotype x environment interactions were present for both herbage and seed yields; a little over half was accounted for by the linear regression. The non-linear component was also significant, although it was smaller than that of the linear component.The most stable herbage and seed yields were obtained from L. sativus. Two selections, 347 and 311, origmating from Syria and Turkey respectively, combined both high herbage and seed yields with wide adaptation and stability, and could be considered the most widely adapted lines. Other lines were identified as suitable for favourable and unfavourable environments; in particular, all four lines of L. ochrus have great potential in frost-free climates. The importance of genotype x environment interactions in future breeding strategies is discussed.     

The Gene Pools of the Grasspea (Lathyrus sativus L.)

Interspecific hybridization between the grasspea, Lathyrus sativus, and 15 wild species in Section Lathyrus is reported. Only two species, L. amphicarpos and L. cicera produced viable F¹ hybrids with low fertility when crossed with L. sativus as male parent. Crosses with six other species produced pods following pollination when L. sativus was the male, but seedlings were inviable, seeds did not germinate or pods were empty or had totally shrivelled seeds. When L. sativus was the female parent, only one cross with L. gorgoni produced an F¹, but the seedling was inviable. The germplasm resources of the grasspea are identified, with L. amphicarpos and L. cicera placed in the secondary gene pool and the other species in the tertiary gene pool. The definition of these germplasm resources is discussed in terms of grasspea improvement through plant breeding.     

Variation in the grass pea (Lathyrus sativus L.) and wild species

Summary Forty-nine accessions of Lathyrus sativus were studied for morphological variation. Data were analysed using Principal Components Analysis and Cluster Analysis. The variation in 14 species of Section Lathyrus was also evaluated in order to ascertain affinities between L. sativus and other species. L. sativus is a highly variable species, and there is a clear distinction between the blue-flowered forms from south-west Asia, Ethiopia and the Indian subcontinent, and the white and white and blue flowered forms with white seeds which have a more westerly distribution. Differences in vegetative parts may be due to selection for forage types. L. sativus appears to be closely related to L. cicera and L. gorgoni, and this relationship needs further investigation.     

Current status and future strategy in breeding grasspea (Lathyrus sativus)

Efforts in grasspea (Lathyrus sativus) improvement have increased since the development of lines that are very low in the neurotoxin Beta-N-oxalyl-L-alpha-beta-diamino propionic acid (ODAP); also referred to as Beta-N oxalyl-amino-L-alanine (BOAA). Many programs now address several related aspects of improvement simultaneously. These include reduced ODAP concentrations, insect and disease resistance, nitrogen fixation, agronomic practices, fodder and forage production, and components for increased yielding ability. The coordinated, multidisciplinary approach now being applied to the genetic improvement of grasspea should allow the potential of this largely neglected grain legume to be fully realized.     

Inheritance of neurotoxin (ODAP) content, flower and seed coat colour in grass pea (Lathyrus sativus L.)

Summary A strong epidemiological association is known to exist between the consumption of grass pea and lathyrism. A neurotoxin, -N-Oxalyl-L-, -diaminopropanoic acid (ODAP) has been identified as the causative principle. This study was undertaken to investigate the mode of inheritance of the neurotoxin ODAP, flower and seed coat colour in grass pea. Five grass pea lines with low to high ODAP concentration were inter-crossed in all possible combinations to study the inheritance of the neurotoxin. Parents, F₁ and F₂ progenies were evaluated under field condition and ODAP analyzed by an ortho-phthalaldehyde spectrophotometric method. Many of the progenies of low x low ODAP crosses were found to be low in ODAP concentration indicating the low ODAP lines shared some genes in common for seed ODAP content. The F₁ progenies of the low ODAP x high ODAP crosses were intermediate in ODAP concentration and the F₂ progenies segregated covering the entire parental range. This continuous variation, together with very close to normal distribution of the F₂ population both of low x low and low x high ODAP crosses indicated that ODAP content was quantitatively inherited. Reciprocal crosses, in some cases, produced different results indicating a maternal effect on ODAP concentration. Blue and white flower coloured lines of grass pea were inter-crossed to study the inheritance of flower colour. Blue flower colour was dominant over the white. The F₂ progenies segregated in a 13:3 ratio indicating involvement of two genes with inhibiting gene interactions. The gene symbol LB for blue flower colour and LW for white flower colour is proposed.     

Different sex phenotypes of Cucumis sativus L. and C. melo L. and their endogenous gibberellin activity

Summary Gibberellins were extracted from seeds and green plants of different sex types of the closely related species Cucumis sativus L. (cucumber) and C. melo L. (muskmelon). Both seeds and green plants of monoecious and andromonoecious lines of C. sativus contained significantly higher gibberellin levels at all growth stages tested than did a gynoecious line. Gibberellin activity in the monoecious and andromonoecious C. sativus line reached a maximum at the growth stage corresponding to flower differentiation at cotyledonary and first leaf axils. Vernalization of seeds of gynoecious plants resulted in increased male tendency correlated with increased gibberellin activity. In contrast, monoecious and andromonoecious lines of C. melo were gibberellin deficient relative to hermaphroditic and gynoecious lines.Journal Article No 5534 from the Michigan Agricultural Experiment Station. This work was supported by a grant from the Herman Frasch Foundation.     

Lathyrus improvement for resistance against biotic and abiotic stresses: From classical breeding to marker assisted selection

Summary Several Lathyrus species and in particular Lathyrus sativus (grass pea) have great agronomic potential as grain and forage legume, especially in drought conditions. Grass pea is rightly considered as one of the most promising sources of calories and protein for the vast and expanding populations of drought-prone and marginal areas of Asia and Africa. It is virtually the only species that can yield high protein food and feed under these conditions. It is superior in yield, protein value, nitrogen fixation, and drought, flood and salinity tolerance than other legume crops. Lathyrus species have a considerable potential in crop rotation, improving soil physical conditions; reducing the amount of disease and weed populations, with the overall reduction of production costs. Grass pea was already in use in Neolithic times, and presently is considered as a model crop for sustainable agriculture. As a result of the little breeding effort invested in it compared to other legumes, grass pea cultivation has shown a regressive pattern in many areas in recent decades. This is due to variable yield caused by sensitivity to diseases and stress factors and above all, to the presence of the neurotoxin β-N-oxalyl-L-α,β-diaminopropionic acid (β-ODAP), increasing the danger of genetic erosion. However, both L. sativus and L. cicera are gaining interest as grain legume crops in Mediterranean-type environments and production is increasing in Ethiopia, China, Australia and several European countries. This paper reviews research work on Lathyrus breeding focusing mainly on biotic and abiotic resistance improvement, and lists current developments in biotechnologies to identify challenges for Lathyrus improvement in the future.     

Diversity of white and narrow-leafed lupin genotype adaptive response across climatically-contrasting Italian environments and implications for selection

White lupin (Lupinus albus) and narrow-leafed lupin (L. angustifolius) have special interest as high-protein feed crops but their cultivation is limited by low grain yields. This study aimed to support breeding programs targeting Italy or other climatically variable south-European regions by investigating within-species adaptation patterns across contrasting Italian environments. An additional aim was comparing species for yielding ability. Eight narrow-leafed and six white lupin cultivars featuring different origin, phenological type (Mediterranean in both species; winter in white lupin; spring in narrow-leafed lupin) and plant architecture (determinate or indeterminate in both species; tall or dwarf in white lupin) were evaluated in a Mediterranean and a subcontinental-climate site under autumn and late-winter sowing. Additive main effects and multiplicative interaction was preferable to joint regression for modeling yield responses. In both species, cross-over GE interaction was observed (P < 0.05), autumn-sown Mediterranean and subcontinental environments were the most-contrasting for GE effects, and widely adapted material included cultivars of Mediterranean phenological type with indeterminate growth. Material with determinate growth was not among the best-yielding entries in any environment, whereas a dwarf winter-type white lupin entry was specifically adapted to autumn-sown subcontinental environments. White lupin displayed larger genetic variation than narrow-leafed lupin for phenology and other traits. Relationships of morphophysiological traits with grain yield were environment-specific and were locally high for some white lupin traits (early flowering, long reproductive phase, high aerial biomass, low proportion of pod wall). White lupin exhibited higher yielding ability than narrow-leafed lupin in all environments but the late-winter sown Mediterranean one, when comparing locally top-yielding cultivars.     

Photosynthesis is Reduced,and Seeds Fail to Set and Fill at Similar Soil Water Contents in Grass Pea (Lathyrus sativus L.) Subjected to Terminal Drought

Grass pea (Lathyrus sativus L.) is an indeterminate grain legume considered adapted to dry environments, but the mechanisms of its adaptation are not well understood. Grass pea plants were exposed to terminal drought from podding, and the development of water deficit was measured together with its effects on leaf photosynthesis, stomatal conductance, carbon remobilisation to the seeds, flower production and abortion, pod production and abortion, seed set, seed growth and the neurotoxin β‐N‐oxalyl‐L‐a, β‐diaminopropionic acid (β‐ODAP) concentration. Predawn leaf water potential (Ψ_leaf), stomatal conductance (gs), rate of leaf photosynthesis (Pn), flower production, pod production, filled pod number, seed number, seed size and yield decreased, while flower abortion, pod abortion and seed abortion increased, and the concentration of β‐ODAP was unchanged under terminal drought conditions. gs and Pn began to decrease at a higher plant‐available soil water content (PAWC) (67.2 ± 2.3 % and 62.9 ± 2.3 %) than Ψ_leaf (43.7 ± 1.1 %). Flowers and pods ceased being produced only when the PAWC decreased below 40.1 ± 4.6 % and 35.3 ± 3.0 %, respectively, but seed set and seed growth ceased when PAWC decreased below 55.5 ± 1.6 % and 58.0 ± 3.7 %, respectively. The mobilization of ¹³C labelled assimilates from the stems was greater under terminal drought than under well‐watered conditions, but the transfer to the seed was small. We conclude that seed set and seed growth decreased as the soil dried due to a reduction in current photosynthesis as a result of stomatal closure.     

Effect of plant density on selection for seed yield in two population types of Phaseolus vulgaris L.

Summary Pedigree selection for seed yield, using early generation yield tests, was practiced from the F₂ to F₇ in two populations of common bean (Phaseolus vulgaris L.) in three plant densities: 66, 133, and 266 thousand plants ha^-1 at CIAT-Palmira, Colombia. The six highest yielding lines selected from each plant-density, along with 13 parents, were evaluated in a 7×7 lattice design with three replications at the three densities utilized for selection, and at 399,000 plants ha^-1 in 1988 and 1989.Based on the mean performance of selected lines and the mean of the parents, selection for seed yield was effective in all densities in both populations. However, none of the lines selected from the population within the race Mesoamerica (TC 4673) significantly outyielded their best parent under any plant density. The highest yielding lines selected from the interracial population (TR 4635) outyielded their best parent irrespective of the plant-density used for selection. The highest yielding line originated from the highest density used for selection. Low density was neither good for selection nor for evaluation and identification of high-yielding cultivars of common bean. There was no significant difference between the mean yield of lines selected at the intermediate and high population densities. The effects of plant density, year, and their interactions were significant for seed yield.     

Location-Specific and Across-Location Selections for Seed Yield in Populations of Common Bean, Phaseolus vulgaris L.

Two populations of common bean, Phaseolus vulgaris L., were used to make three independent location-specific selections at Palmira, Quilichao, and Popayán, Colombia. Also, three across-location selections were made by alternating generations between Palmira and Popayan, between Quilichao and Popayán, and among Palmira, Quilichao, and Popayán. The seven highest yielding lines selected from each environment and population along with 14 parents and two checks were evaluated for seed yield, seed weight, maturity, and reaction to an-thracnose, angular leaf spot, and common bacterial blight. Mean squares due to evaluation and selection environments, selected lines, and their interactions were significant for all traits. Mean yields of selected lines were higher than the mean of parents in both populations. Among location-specific selections, lines yielded higher at their respective selection site. Moreover, selections made at Popayán were higher yielding than those made at Quilichao or Palmira when tested across locations and years. Use of Popayán, a relatively wetter and cooler site, either alone or in combination with other locations resulted in higher yield, higher seed weight, and higher resistance to anthracnose. On the average, location-specific selections were lower yielding than the across-location selections. Among across-location selections, those selected between Palmira and Popayán were the highest yielding in both populations. Selections at Palmira and across all three locations resulted in greater stability.     

Pre and posthaustorial resistance to rusts in <Emphasis Type="Italic">Lathyrus cicera</Emphasis> L.

Lathyrus cicera has a high potential as fodder crop in dry areas, but can in particular environments be damaged by rust. Little is known on the availability of resistance against rust fungi and the underlying mechanisms in L. cicera germplasm. The present study assessed and characterised macro and microscopically the resistance to rust fungi Uromyces pisi and U. viciae-fabae, in a collection of L. cicera accessions. A wide range of disease reaction was found in the germplasm collection against the different rust species. L. cicera accessions were highly resistant to U. viciae-fabae being hypersensitive response the most frequent reaction. On the contrary, most accessions showed a compatible interaction with U. pisi, with varying levels of partial resistance, although cases of hypersensitivity were also identified. Differences on germination, orientated germ tube growth and appressoria differentiation were observed but were in general of marginal importance to explain the resistance to U. pisi among the L. cicera accessions. Resistance was due, to a combination of pre and post-haustorial mechanisms.     

Interspecific hybridization in Cucumis L. II. The crossability of species,an investigation of in vivo pollen tube growth and seed set

Summary The crossability of 12 Cucumis species of African and Asiatic origin was studied in a diallel cross, in order to find ways to realise the cross between the common cucumber (C. sativus L.) and its wild relatives which carry resistances against diseases and pests.Self-pollinations and cross-pollinations within species gave normal pollen tube growth and seet set. The different accessions within a species, as a rule reacted alike in interspecific crosses. In crosses between African species different crossing patterns were found, viz. bilateral congruity, bilateral incongruity and unilateral incongruity. Within C. sativus all accessions intercrossed freely, except one, which displayed unilateral incongruity.Good seed was harvested from several crosses and in some cases embryo culture was needed for further development of seeds. No good seeds were obtained from any cross between a species of the African group and C. sativus L.     

Variation and association of morphological and biochemical characters in grass pea (Lathyrus sativus L.)

Fifty grass pea land race populations of Ethiopia that were selected based on administrative regions and different altitude classes were used in this study to determine variability in morphological characters, Oxalyl Diamino Propanoic acid(ODAP) – a causative agent of lathyrism, and protein contents and their associations. Highly significant differences were observed among the populations for most of the morphological characters. Populations collected from Gondar region and the higher altitude group (>2550 m) showed the highest mean and coefficient of variation indicating the presence of high genetic diversity in this region and altitude group. ODAP analysis from single plants showed significant variation both within a population and between populations. Four individual plants with low ODAP contents ranging from 0.149% to 0.182% (range within safe level) were identified in populations collected from different regions, indicating the diversity and variation of this trait with in one population and between regions. These low ODAP lines are considered important for further breeding. The association of ODAP with grain yield, biomass, plant height and seed size was negative, suggesting that the selection of tall and late maturing varieties with large seed size and high grain yield potential will enable the development of varieties with low ODAP content. This revised version was published online in August 2006 with corrections to the Cover Date.     

Collection and evaluation of grasspea (Lathyrus sativus L.) germplasm of central Italy

Characterization of 16 grasspea (Lathyrus sativus L.) accessions collected in central Italy (Marche region) was performed for morphological and agronomic traits and for 3‐(‐N‐oxalyl)‐L‐2,3 diaminoprop‐ionic acid (ODAP) content. Field trials, carried out in two locations, revealed high among‐population genetic variation. In particular, an evident differentiation between commercial populations, cultivated by market‐oriented farms, and household populations (not market‐oriented) was found. Genotype × environment interaction was negligible. Medium‐high values of ODAP content were observed following colorimetric and capillary electrophoresis analyses. A high positive correlation between the two methods was found (r = 0.83**), but the colorimetric values showed, on average, significant 14% lower ODAP values. This research represents a pre‐breeding activity aimed at developing a breeding programme for the preservation and utilization of Italian grasspea germplasm within sustainable or organic agricultural systems.     

Self-fertility and the distribution of seed yield in Vicia faba L.

Summary Five lines of field beans (Vicia faba L.) bred and selected for a high level of self-fertility have demonstrated heritable differences in distribution of seed yield on the reproductive nodes when grown under conditions of selfpollination. Fully self-fertile genotypes produce a large proportion of the seed yield on the lower nodes. The breeding and selection for both self-fertility and the distribution of seed yield is discussed.     

Evaluation of Pedigree and Single Seed Descent Selection Methods for Cultivar Development in Cowpea (Vigna unguiculata L. Walp)

Three crosses of cowpea (Vigna unguiculata [L.] Walp) were used to assess the effectiveness of two selection procedures in cowpea. The selection procedures were pedigree selection (PD) and single seed descent selection (SSD). Lines developed through each procedure were yield tested and selected at F₆ and F₇ for PD and SSD, respectively. The selected lines for both procedures were compared at F₈ for grain yield per plant (YLD), number of pods per plant (NPO), and pod development period (PDP), in two locations. Lines developed through both procedures differed more in their population means than in the performance of their top 10% or highest yielding lines. There was no consistent relationship between magnitude of genetic variability and percentage of superior lines obtained from each procedure. Both procedures have been effective in producing superior genotypes for yield and number of pods. The PD method produced superior transgressive segregants in two out of the three crosses for PDP. SSD allowed a more rapid generation than PD.     

Comparison of intra- and inter-pool crosses in faba beans (Vicia faba L.). I. Hybrid performance and heterosis in Mediterranean and German environments

Two recently detected, promising CMS systems in faba bean have opened up new possibilities in hybrid breeding. This study was conducted to determine hybrid performance and heterosis of 33 intra-pool and 66 inter-pool crosses from a factorial of six European small-seeded (Minor), six European large-seeded (Major), and eight Mediterranean faba bean lines. The parent lines and F₁ crosses were evaluated at two German and seven Mediterranean environments for anthesis, maturity, yield, yield components and plant height. Yield in the Mediterranean environments was 60% of that in the German environments. European Minor lines and their intra-pool crosses were unadapted to Mediterranean environments and yielded only 31 % compared with the Mediterranean lines and their crosses, whereas European Major genotypes yielded 58%. In German environments the European crosses yielded 107% of the Mediterranean crosses. Heterotic yield increase over the parental mean was largest for European Minor x Mediterranean inter-pool crosses, amounting to 95% in the Mediterranean and 73% in German environments. Our results confirmed the expectation of an increased heterosis in inter-pool over intra-pool hybrids and poor adaptation of European genotypes in Mediterranean environments. In the latter, Mediterranean intra-pool hybrids were the most promising, whereas in Germany inter-pool hybrids of the European Minor × Mediterranean type were most promising.     

Selection for multiple disease resistance reduces cucumber yield potential

Summary Experiments were conducted in Wisconsin and Michigan to determine whether selection for multiple disease resistance adversely affects yielding ability of commercial cucumber (Cucumis sativus var. sativus L.; CS) lines. Inbred F₄ and F₅ lines were developed from C. sativus var. hardwickii (R.) Alef. (CH) x CS which had been either selected or not selected for resistance to the causal organisms of scab, anthracnose, and downy mildew. The exotic CH germplasms possesses a multiple fruiting habit with high yield potential and the CS genotypes include several disease resistance. In each comparison between selected and unselected progeny, the unselected families either significantly outyielded, or were not significantly different than their selected counterparts. In no case did the selected progeny outyield the unselected progeny. Since this was observed for both fruit number per plant and total fruit weight per plant, and it occurred despite differences in fruit size, we conclude that selection for disease resistance led to a reduction in yield potential in these populations.Received for publication-. The cost of publishing this paper was defrayed in part by the payment of page charges. Under postal regulations, this paper therefore must be hereby marked advertisement solely to indicate this fact.     

Grass pea (<Emphasis Type="Italic">Lathyrus sativus</Emphasis>): Is there a case for further crop improvement?

An ideal legume for resource-poor farmers, grass pea [Lathyrus sativus] is a drought tolerant crop that thrives with minimal external inputs. It is grown on 1 million ha throughout South Asia, mainly as a relay crop after rice. It is also grown extensively in Ethiopia, where it is an important legume for human consumption. Traditionally used for human consumption and as a source of animal feed, this protein-rich legume is favoured for its excellent flavour. There is great potential for an expansion in the utilization of grass pea in drought-prone economies, such as Ethiopia. However, L. sativus produces small quantities of a neurotoxin, β-N-oxalyl –L-α-diaminopropanoic acid (ODAP), which, when consumed alone in large quantities, may cause ‘lathyrism’, an irreversible paralysis of the legs. Lathyrism is a medical condition closely associated with poverty in rural areas. Despite efforts in some countries to discourage production of L. sativus, poor farming communities continue to rely on the crop to supplement their meagre diets. As local land races with high toxin levels generally outperform introduced varieties with lower toxin levels, farmers have little option but to continue to grow their locally-adapted varieties. While there has been some success in breeding grass pea lines with low levels of ODAP, crop improvement programmes are scarce and under-resourced. Facing rising food prices and more frequent natural disasters associated with climate change, we should not neglect the ability of grass pea to provide human and animal feed, under conditions unsuitable for economic production of other legume crops.