Variation in tolerance to radiant frost at reproductive stages in field pea germplasm

Radiant frost is a major abiotic stress, particularly at the reproductive stage, in field pea (Pisum sativum L.) grown in Mediterranean environments. Here, response to frost was studied for flowering stage (FS) organs (buds, flowers and set pods) and pod development stage (PDS) organs (flat, swollen and mature pods) under controlled conditions, with plants exposed to a minimum temperature of –4.8°C for 4 h. This frost treatment adversely affected seed yield through (i) abortion of buds, flowers and set pods (ii) death of pods and (iii) reduction in seed size. FS organs were more sensitive to frost than PDS organs. Genetic variation was observed among 83 accessions collected from 34 countries worldwide for survival of FS buds, flowers and set pods. In 60 of 83 accessions, no buds, flowers or set pods survived the frost treatment. Five accessions: ATC 104 (origin: United Kingdom), ATC 377 (Estonia), ATC 968 (Italy), ATC 3992 (Kazakhstan) and ATC 4204 (China), showed the highest frost tolerance of FS organs and lowest numbers of abnormal seeds. The frost tolerant accessions identified in this study may be useful as parents for breeding field pea varieties that will be less likely to suffer yield loss due to radiant frost during the reproductive stage.     

The inheritance of seed colour and vernalization requirement in Brassica napus using doubled haploid populations

Summary In the Mediterranean coastal region, spring barley cultivars are sown in the autumn, corresponding with the rainy season. Thirty eight cultivars out of 44 spring which were collected from North African countries including Morocco, Algeria, Tunisia, Libya and Egypt were of a single dominant spring gene type (+Sh ₂+). Almost the same situation was found in Spain, Italy and Greece, with exceptions for France and Turkey. Kernel row of North African cultivars was characterized by six-rowed type, though there was one exception in Egypt. In the South European countries of Spain, Italy and Greece, six-rowed type varieties were also prevalent. As to non-brittle rachis, in North African countries oriental type cultivars (Btbt ₂) were much higher in frequency than occidental type cultivars (btBt ₂), while in European countries, occidental type cultivars were rather dominant, except for Italian cultivars. In Turkey, located in the joint area between Europe and Asia, cultivars tested were mostly occupied by spring genotype other than +Sh ₂+, and also characterized by tworowed head and occidental type for non-brittle rachis. In the Mediterranean coastal region, especially in North Africa, restricted genotypes, +Sh ₂+ for spring growth habit and v for six-row may indicate adaptability favorable to the growth conditions in these areas. Furthermore, North African cultivars were characterized by the oriental type of non-brittle rachis. This may give us information about the migratory routes of the original landrace to North Africa.     

Factors affecting in vitro flowering and fruiting of green pea (Pisum sativum L.)

Multiple shoots were efficiently regenerated from cotyledonary node and shoot tip explants of Pisum sativum within 15 days on MS medium containing B₅ vitamins and supplelmented with 2.0 mgl^-1 6-benzylaminopurine. The elongated shoots produced on the same medium were excised and transferred to MS medium containing half strength ammonium nitrate (8.25 gml^-1) and supplemented with auxins (indole-3-butyric acid or naphthalene acetic acid) either alone or in combinations with gibberellic acid. Rooting and flowering were observed on the 7th and 15th day after their transfer to rooting medium. The flowers self-fertilised in vitro and produced mature pods within 25 days of rooting. These seeds were germinable both in vitro and in vivo. In vitro seeds sown in pots under field conditions developed into flowering plants, and subsequently produced pods with viable seeds. This revised version was published online in July 2006 with corrections to the Cover Date.     

Water Deficit during the Reproductive Period of Grass Pea (Lathyrus sativus L.) Reduced Grain Yield but Maintained Seed Size

The grain legume grass pea (Lathyrus sativus L.) is adapted to drought‐prone environments, but the extent and mechanisms of its tolerance are not well understood. In a pot experiment, water deficit was imposed on plants by withholding water from first flowering until predawn leaf water potential (LWP_pd) was ?3.12 MPa. Water deficit reduced dry matter, seed yield, harvest index and water use efficiency by 60 %, 87 %, 67 % and 75 %, respectively, when compared with the controls. Flower production stopped when LWP_pd fell to ?1.8 MPa. At LWP_pd?1.5 MPa, only 25 % of flowers resulted in filled pods (compared with 95 % filled pods in the control) with the rest aborted as flowers (48 %) or pods (27 %). Filled pods had more aborted ovules than controls, resulting in 29 % less seeds per pod. Water deficit reduced pollen viability, germination and the number of pollen tubes reaching the ovary by 13 %, 25 % and 31 %, respectively. Emergence from seeds produced from water‐deficient plants was 21 % less than controls, but seedling shoot dry mass was 18 % higher, in accordance with the 19 % higher seed mass. The sensitivity of flowering to drought limited pod numbers but enabled plants to retain existing pods and develop near‐normal seeds with low β‐N‐oxalyl‐l ‐α‐β‐diaminopropionic acid toxin concentrations. This trait is useful for farming systems reliant on harvested seed for the next crop and in cases where seed size influences the value of the product.     

Underground Vetch (Vicia sativa ssp. amphicarpa): A Potential Pasture and Forage Legume for Dry Areas in West Asia

Subterranean vetch [Vicia sativa ssp. amphicarpa (Dorth.) Aschers & Graebn.] is native to disturbed grasslands of the Mediterranean basin where heavy grazing, seasonal drought and erosion act as strong selection forces. It produces two pod types, above‐ground and 5 cm below the soil surface. Unlike subterranean clover (Trifolium subterranean L.), which buries its seeds after flowering above‐ground, subterranean vetch flowers and forms pods beneath the soil surface on underground stems. The aerial pods are produced after vegetative development ceases, while the underground pods are produced in ontogeny. The ability of this unusual vetch to survive in marginal areas with low rainfall (about 250 mm year^?1) and to produce nutritious herbage and pods is an important characteristic which helps address rehabilitation of degraded rangelands and increase feed production for small ruminants. Research at the International Center for Agricultural Research in the Dry Areas (ICARDA) during the 1988–93 growing seasons has assessed the herbage and seed productivity of underground vetch, its ability to grow in rotation with barley in marginal low‐rainfall areas, and its capacity to regenerate after heavy grazing. Drier conditions in 1989 favoured earlier underground flowering; the number of underground pods was higher than that of aerial pods. Grain yield of barley (var. Atlas) was around 2.0 t ha^?1 after underground vetch and only 1.2 t ha^?1 after barley. Grazing underground vetch had no effect on the productivity of the succeeding barley crop. The aerial and underground pods serve two distinct functions; aerial pods increase dissemination within suitable habitats, while underground pods increase the probability of plant survival under adverse conditions such as drought and heavy grazing. Underground vetch has two potential uses, namely the rehabilitation of marginal areas and production in rotation with barley.     

Genetic diversity assessment in cultivated cardoon by AFLP (amplified fragment length polymorphism) and microsatellite markers

Cultivated cardoon (Cynara cardunculus L. var. altilis DC) belongs, together with globe artichoke (C. (cardunculus L. var. sylvestris L.) and wild cardoon (C. cardunculus L. var., sylvestris (Lamk) Fiori). to the family Asteraceae (Compositae). Cultivated cardoon is of regional importance in Italy. Spain and southern France, where it is used for the preparation of traditional dishes. It has been shown to have potential as a source of oil from its seeds, inulin from its roots and various biopharmaceuticals from its leaves. Levels of genetic diversity and relatedness between eleven Italian and 10 Spanish accessions were assessed by DNA profiling with eight AFLP primer combinations and at five microsatellite loci. The AFLP analysis of genetic similarities showed that the Spanish and Italian accessions represent two distinct gene pools; substantial variation was present within each accession. On the other hand a limited variation was detectable by applying SSR markers.     

Production and characterization of hybrids between Cajanus cajan × C. reticulatus var. grandifolius

Cajanus reticulatus var. grandifolius, endemic to Australia and a wild relative of the cultivated species, C. cajan, was successfully crossed with the latter as the female parent. The major wild species characters such as persistent stipules, long pod hairs, pod shattering, brown seeds with grey speckles, and presence of seed strophiole were dominant in the hybrid. For growth and branching habit, and leaflet, flower, pod, and seed size, the hybrid was intermediate between the parents. The meiotic cells of the hybrid were found to have quadrivalents, trivalents, univalents,and showed reduced chromosome pairing as revealed by the increased number of rod bivalents per cell at metaphase-I, and stickiness and precocious movement of chromosomes to poles in the second division. In comparison to the parents, the hybrid had fewer pods and seeds. However,these anomalies in the interspecific hybrid are not significant enough to preclude the gene transfer from C. reticulatus to the cultivated species through a sexual route. This revised version was published online in August 2006 with corrections to the Cover Date.     

Morphology and cytology of Cicer canariense,a wild relative of chickpea

Summary The short note describes the morphology and chromosome number of Cicer canariense Santos Guerra & Lewis. This taxon has been found to have a meiotic chromosome number of 2n<16 and not 2n<24, as reported earlier. At ICRISAT Center, while other perennial Cicers did not perform well, Cicer canariense flowered and produced seeds. The species grew vigorously and had multi-podded peduncles and multi-seeded pods. Such traits of Cicer canariense can be very useful in the genetic improvement of chickpeas.Submitted as JA 1456 by the International Crops Research Institute for the Semi-Arid Tropics (ICRISAT).     

Genetics of Seed Yield and its Components in Common Bean (Phaseolus vulgaris L.) of Middle-American Origin

Genetic variance, heritability, and expected response from selection arc useful in devising alternative methods and criteria of: selection. The objectives of this study were to estimate these for seed yield and its components from 200 F₂: populations involving 80 cultivars and lines of mostly small-seeded dry bush bean (Phaseolus vulgaris L.) of habits growth I, II, and III of Middle-American origin. All cultivars and lines were crossed in eight sets of ten parents each in a Design II mating system. The F₂ populations, without parents, were evaluated in the field in a replicates-in-sets design at two locations in Colombia in 1983. Estimates of additive genetic variance were significant for yield, pods/m², seeds/pod, and seed weight. Interaction with environments was also significant. Values for nonadditive genetic variance were not significant for either yield or yield components. The estimates of narrow sense heritability, based on the F₂ population mean and unbiased by genotype x environment interaction, were 0.21 ± 0.13 for yield. 20 ± 0.13 for pods/m², 0.57 ± 0.13 for seeds/pod, and 0.74 ± 0.15 for seed weight. The expected direct response from selection of the top 20 % of F₂ populations for yield per se would result in a 4.30 % increase in yield with a correlated response of 0.21 % in seed weight. In contrast, the expected gain from direct selection for seed weight would result in a 11.76 % increase in seed weight with a, correlated gain of 0.28 % for yield. Direct selection for pods/m² would decrease yield, seeds/pod and seed weight, while direct selection for seeds/pod would reduce pods/m² and seed weight but increase seed yield by 0.37 %. Data on yield from replicated trials in the early segregating generations could be utilized for identification and selection of promising crosses and families or lines with crosses for dry bean yield improvement.     

Genetic analyses of agronomic and seed quality traits of doubled haploid population in Brassica napus through microspore culture

Summary The results showed that the F1 genotype from the cross (Brassica napus cv. Zheshuang 758 × cv. Z-4115) had good response to embryogenesis, and their embryo yield and rate of plant regeneration reached 69.8 embryo/bud and 46.9%, respectively. Characters from the doubled haploid (DH) populations in B. napus were analyzed and it was showed that the means of each agronomic trait were between their parents, but they were nearer to the paternal in 6 agronomic traits (plant height, branch position, number of pods in the main raceme, length of pod, number of pods/plant and number of seeds/pod). The number of genes controlling each agronomic trait was analyzed based on the DH populations. The results showed that the number of genes controlling number of pods in the main raceme was the highest (15.6), and the least number of genes was involved for stem width (only 7.9). According to estimated coefficients of skewness and kurtosis of the traits tested, gene interaction was found to be absent for stem width, plant height, length of main raceme, number of primary and secondary branches, pod density in the main raceme and seed weight/plant. Complementary interaction was also observed in five agronomic traits (number of pods in the main raceme, length of pod, number of pods/plant, number of seeds/pod and 1000-seed weight). A significantly positive correlation was observed between seed yield/plant and four agronomic traits (length of main raceme, length of pod, number of pods/plant and 1000-seed weight). The experiment also showed that the erucic acid, glucosinolate, oil and protein contents of DH populations were 34.23%, 87.09 μmol/g, 44.09% and 42.67%, respectively. The numbers of genes controlling each quality trait were 7.8, 9.7, 9.4 and 8.7, respectively. Partial correlations between the seed quality traits and the agronomic characters of DH populations were analyzed. In this experiment, the partial correlations among seed quality traits were also analyzed and it was found that the oil content had a negative correlation with the other three seed quality traits.     

Relationship between morphological variation and geographical origin or selection history in Lupinus pilosus

Patterns of morphological diversity were examined in Lupinus pilosus in relation to geographical origins or selection history of accessions. There was significant variation among accessions for most architectural and reproductive characters in a field experiment in Perth, Western Australia. Late flowering was associated with resetted early growth, profuse branching and collection in higher rainfall sites. High yield and profuse podding on the main stem was associated with large pods and leaves, many seeds per pod, greater height to the first main-stem pod, and collection in higher altitude sites. Ten groups of accessions, identified by hierarchical cluster analysis, accounted for 79% of genotype and 69% of genotype x character sums of squares. One group, possibly taxonomically distinct from the others, consisted exclusively of wild types from northern Israel with poor pod set and low seed yield, poor nodulation, pale foliage, and small leaves. Wild types from Turkey and Crete had rosetted early growth and were late flowering, and the Turkish group had extremely rapid mid-season growth. Ornamental types from Europe or Australia with pink, purple or white flowers clustered together in an early flowering group with vigorous early growth. A smooth-seeded type of L. pilosus was discovered in a group of short, late flowering and rough-seeded types from Syria.     

Variation and covariation in Vicia faba L. populations of Mediterranean origins

Summary Variation and covariation in 15 Vicia faba L. ssp. major populations from Italy. Greece and Egypt were analyzed. Ten quantitative morphobiological characters of agronomical and nutritional interest were observed. Differences among countries of origin were highly significant for all characters except for lowest podded node. Phenotypic distances among populations, both within and among origins, estimated on canonical variates evidenced that the Egyptian populations from a distinct group and that a relevant degree of similarity exists among the Greek and the Italian meterial. The covariation structure within populations, which was studied by means of factor analysis, was comparatively simpler in the Egyptian material than in that form the other two Mediterranean countries. The differences evidenced can be attributed to different gene associations between characters and suggest the possibility of obtaining, through recombination, genotypes agronomically more favorable.Research work supported by C.N.R., Italy, special grant. Sub-Project 1, Paper No. 299.     

Biplot analysis of trait relations of white lupin in Spain

A 2-year study of autumn-sown white lupin (Lupinus albus L.) was conducted under rainfed Mediterranean conditions in southern Spain. 108 landraces were evaluated for yield and major phonological and plant structural characteristics. Path coefficient and Genotype-Trait (GT) biplot analysis were used. Applying both types of analyses to the multiple trait data revealed that GT biplot graphically displayed the interrelationships among traits and facilitated visual comparison of accessions and selection. Wide variation was observed for trait phenology (dates of flowering and maturity), plant structure (plant and first order inflorescence height) and yield characters (first order branches, first order inflorescence and total yield). It was found that the accessions with the highest yields were the tallest and the latest to flower, both on the mainstem and on the lowest first order branch, and also were the latest to mature. GT biplots also had suggested as the accessions have been displayed according to their origins. The results showed that foreign germplasm may be an important material for improving some yield components such as number of pods and seeds per plant of locally adapted lupins in Spain. This revised version was published online in July 2006 with corrections to the Cover Date.     

Field performance of natural narrow-leafed lupin from the northwestern Spain

Summary Narrow-leafed lupin or lupine (Lupinus angustifolius L.) is one of the three species of the genus Lupinus that grows naturally in Galicia (northwestern Spain). In this region, with more than one million of cattle heads, almost 20% of Spanish total, there is no cultivation of any protein legume for feed. Lupins are widely used as a source of protein and energy in livestock feed and would contribute to the supply of vegetable protein at low cost. A 2-year study of natural narrow-leafed lupin germplasm was conducted under winter Mediterranean conditions in northwestern Spain. Fifty-nine wild populations and two cultivars developed in Poland were evaluated for major morphological, phenological, and agronomical characteristics. The objectives were to assess the agronomic potential of this germplasm for the future production and breeding, and to know the relationships among populations studied. Lupin populations showed significant differences for most of the traits studied. Genotype × environment interaction was significant for plant height, green and dry mass, pod length, seeds per pod, seed length and width, and water permeability of seed coat, indicating that some genotypes were better fitted to a specific environment than others. Three narrow-leafed lupin populations (LUP-0098, LUP-0138 and LUP-0200) were identified with suitable agronomical characteristics for their possible future cultivation in the region and breeding purposes. The accession LUP-0138 presented the best behaviour for seed production. These results open the way for the winter narrow-leafed lupin to become a promising crop in the future for the northwestern Spain.     

The reproductive biology of three perennial Sesbania species (Leguminosae)

Summary The reproductive biology of Sesbania sesban, S. goetzei and S. keniensis (Leguminosae; Papilionoideae) was studied. Fifty percent flowering was observed at 102 to 153 days after germination for S. sesban accessions; 96 to 146 days for S. goetzei and 131 to 176 days for S. keniensis accessions. Flowers opened in the afternoon and remained fresh for 2–3 days. Bee species including Xylocopa sp., Apis mellifera, Megachile bituberculata and Chalicodoma sp. visited the flowers. Hand pollination experiments showed that all three species were self and cross compatible. The percentage of fully developed seeds was greater in pods formed after cross pollination compared to self pollination. No evidence was found for stigmatic or stylar self incompatibility. Outcrossing is probably the common method of reproduction under natural conditions, although in isolated trees substantial seed set by selfing might occur. Pod production under natural conditions was 34% for S. sesban; 49% for S. goetzei and 39% for S. keniensis. Considerable variation was found in pod production under open pollination between accessions of the same species. Selective abortion was observed within pods, with more mature seeds formed at the distal end of the pod.     

Impact of heat stress on pod-based yield components in field pea (Pisum sativum L.)

Elevated temperatures associated with climate change result in crops being exposed to frequent spells of heat stress. Heat stress results in reduced yield in field pea (Pisum sativum L.); it is therefore important to identify cultivars with improved pod and seed retention under heat to mitigate this loss. Objectives were to investigate the effect of heat stress on phenology, yield and pod-based yield components. Sixteen pea cultivars were evaluated at normal and late (hot) seeding dates in the field in Arizona 2012 and in growth chambers with two temperature regimes (24/18°C and 35/18°C day/night temperature for 7 days) during reproductive development. We measured variation in the pattern of pod retention at four-node positions on plants, seed retention by ovule position (stylar, medial and basal) within pods and screened cultivars for pod retention, seed retention and yield. Heat stress reduced seed yield by accelerating the crop lifecycle and reducing pod number and seed size. Heat stress had the most damaging effect on younger reproductive growth (flowers and pods developed later), resulting in ovary abortion from developing flowers. Heat also accelerated seed abortion in all ovule positions within pods. Two high-yielding cultivars under control temperature, “Naparnyk” and “CDC Meadow”, maintained high yield in heat, and “MFR043” had the lowest yield. Cultivars “40-10” and “Naparnyk” retained the most ovules and seeds per pod, and “MFR043” aborted seeds when exposed to heat. In half of the cultivars, ovules at the basal peduncle end of pods were likely to abort while ovules at the medial and stylar end positions developed into seeds. For seven of the field cultivars, ovules at the medial pod position also produced mature seeds. Cultivars “40-10”, “Naparnyk” and “CDC Meadow” had greater pod and ovule retention or maintained high yield under heat stress, and were identified as heat-tolerant cultivars. Our results allow for a better understanding of pod-based yield components in field pea under heat stress and developing heat-tolerant cultivars.     

Flowering, Pod Set and Reproductive Success in Soya Bean

Much of the variation in yield of soya bean [Glycine max (L.) Merrill] and other grain crops is associated with changes in the number of pods and seeds per unit area. Photosynthesis and seed characteristics are the primary determinants of pod and seed number, but recent research suggests that the temporal distribution of flower production may also play an important role. Soya bean has a long flowering period (up to 40 days or more), but most of the flowers are produced in a much shorter time. The length of the period varies among cultivars, growth habits (indeterminate and determinate) and environments. The reproductive success of flowers produced early in the period is usually greater than those produced later. Little is known, however, of the regulation of the temporal distribution of flower production or its potential role in determining pod and seed number at maturity. Research is needed to determine first, if the temporal distribution can be manipulated to increase pod and seed set and secondly, if such increases would result in higher yields.     

Durum Wheat under Mediterranean Conditions as Affected by Seed Size

This study was carried out in order to determine the effect of seed size on the growth and yield of durum wheat. Four field experiments, composed of six durum wheat (Triticum turgidum L. var. durum) varieties and three seeding sizes were conducted in north‐east Spain in 2000 and 2001 in randomized complete block designs. The growth of seedlings was dramatically affected by seed size. Large seeds produced greater plot stands, but the plants had fewer tillers, leaves and spikes and less green area and dry weight than plots from small seeds. Grain yield was 16 % greater in plots from large seeds, which resulted in greater biomass, green area index, spikes per m² and heavier kernels than in plots from small seeds. Kernel weight was the yield component most related to grain yield in the three seed sizes. The percentage of yield variation explained by kernel weight increased as the weight of kernels increased. This was a consequence of the use of larger seeds, the same effect being observed when the comparison was made between varieties with different kernel weights. Selection for heavy kernel varieties may help to improve the yield of durum wheat in Mediterranean environments similar to that prevailing in north‐east Spain.     

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.