Screening techniques and sources of resistance to foliar diseases caused by major necrotrophic fungi in grain legumes

Summary Necrotrophic pathogens of the cool season food legumes (pea, lentil, chickpea, faba bean and lupin) cause wide spread disease and severe crop losses throughout the world. Environmental conditions play an important role in the development and spread of these diseases. Form of inoculum, inoculum concentration and physiological plant growth stage all affect the degree of infection and the amount of crop loss. Measures to control these diseases have relied on identification of resistant germplasm and development of resistant varieties through screening in the field and in controlled environments. Procedures for screening and scoring germplasm and breeding lines for resistance have lacked uniformity among the various programs worldwide. However, this review highlights the most consistent screening and scoring procedures that are simple to use and provide reliable results. Sources of resistance to the major necrotrophic fungi are summarized for each of the cool season food legumes. Marker-assisted selection is underway for Ascochyta blight of pea, lentil and chickpea, and Phomopsis blight of lupin. Other measures such as fungicidal control and cultural control are also reviewed. The emerging genomic information on the model legume, Medicago truncatula, which has various degrees of genetic synteny with the cool season food legumes, has promise for identification of closely linked markers for resistance genes and possibly for eventual map-based cloning of resistance genes. Durable resistance to the necrotrophic pathogens is a common goal of cool season food legume breeders.     

Integrated “omics” approaches to sustain global productivity of major grain legumes under heat stress

Grain legumes serve as key sources of dietary protein to the global human population. Consequence of high‐temperature (HT) stress is increasingly evident as drastically lost production of different crops including grain legumes worldwide, thus putting the global food security under great threat. In a changing climate scenario, cool season‐adapted grain legumes frequently encounter heat stress (HS) during their reproductive phase, thus witnessing serious yield losses. To combat the emerging challenges of HT stress, an integrated approach demanding collaborative efforts from various disciplines of plant science should be in place. This review summarizes major impacts of HT stress on grain legume, and captures the relevance of crop genetic resources to HS tolerance in these crops. Measurement of physiological traits assumes key place in view of ever‐increasing precision of next‐generation phenotyping assays. We also discuss the significance of genetic inheritance and QTL discovery and evolving “omics” science for developing HS tolerance grain legume crops.     

Legume breeding for rust resistance: lessons to learn from the model <Emphasis Type="Italic">Medicago truncatula</Emphasis>

Rusts are major biotic constraints of legumes worldwide. Breeding for rust resistance is regarded as the most cost efficient method for rust control. However, in contrast to common bean for which complete monogenic resistance exists and is efficiently used, most of the rust resistance reactions described so far in cool season food legumes are incomplete and of complex inheritance. Incomplete resistance has been described in faba bean, pea, chickpea and lentil and several of their associated QTLs have been mapped. However, the relatively large distance between the QTLs and their associated molecular markers hampers their efficient use for marker assisted selection. Their large genome size drastically hampers the development of genomic resource and limits the saturation of their genetic maps. The use of model plants such as the model legume Medicago truncatula may circumvent this drawback. The important genetic and genomic resources and tools available for this model legume can considerably speed up the discovery and validation of new genes and QTLs in resistance to legume pathogens. Here, the potential of M. truncatula as a model to study rust resistance in legumes, and to transfer rust resistance genes to cool season grain legumes is reviewed.     

Transfer of the Kosena Rfk1 gene,required in hybrid seed production,from oilseed rape to turnip rape

Radiant frost is a major abiotic stress, and one of the principal limiting factors for agricultural production worldwide, including Australia. Legumes, including field pea, faba bean, lentil and chickpea, are very sensitive to chilling and freezing temperatures, particularly at the flowering, early pod formation and seed filling stages. Radiant frost events occur when plants and soil absorb the sunlight during the day time and radiate heat during the night when the sky is clear and the air is still. Dense chilled air settles into the lowest areas of the canopy, where the most serious frost damage occurs. The cold air causes nucleation of the intracellular fluid in plant tissues and the subsequent rupturing of the plasma membrane. Among the cool season grain legume crops, chickpea, lentil and faba bean and field pea are the most susceptible to radiant frost injury during the reproductive stages. The more sensitive stages are flowering and podding. Frost at the reproductive stage results in flower abortion, poor pod set and impaired pod filling, leading to a drastic reduction in yield and quality. In contrast, in the UK and European countries, frost stress is related to the vegetative stages and, in particular, the effects of frost have been studied on cotyledon, uni/tri-foliolate leaf and seedling stages during the first few weeks of growth. Few winter genotypes have been identified as frost tolerant at vegetative stages. Vegetative frost tolerance is not related to reproductive frost tolerance, and hybrids from the vegetative frost-tolerant genotypes may not necessarily be tolerant at the reproductive stage. Tolerance to radiant frost has an inverse relationship with plant age. In the field, frost tolerance decreases from the vegetative stage to reproductive stage. Unlike wheat and barley, it is difficult to analyse and score frost damage in grain legume crops due to the presence of various phenophases on one plant at the reproductive stage. The extent of frost damage depends on the specific phenophases on a particular plant. However, current studies on genetic transformation of cold tolerant gene(s), membrane modifications, anti-freeze substances and ice nucleating or inhibiting agents provide useful information to improve our current understanding on frost damage and related mechanisms. The effects of frost damage on yield and grain quality illustrate the significance of this area of research. This review discusses the problem of radiant frost damage to cool season legumes in Australia and the associated research that has been carried out to combat this problem locally and worldwide. The available literature varies between species, specific climatic conditions and origin.     

Identifying and mapping genes of economic significance

An understanding of the genetic basis of characters of commercial importance is critical if a breeder is attempting to move such characters into breeding material. A number of particularly interesting characters or genes have been identified in cool season food legumes, and in pea many of these have been tagged by molecular markers such as allozyme or DNA polymorphisms. This process of mapping and tagging genes has been greatly accelerated by recent developments in molecular biology. It appears that markers will soon be available for many genes in lentil, faba bean, and chickpea and that genetic knowledge developed in one crop will have significant applications in the other cool season food legumes.     

Water use and water use efficiency of cool season grain legumes in low rainfall Mediterranean-type environments

The water use (E_t) and water use efficiency (WUE) of a range of cool season grain legume species (field pea [Pisum sativum L.], faba bean [Vicia faba L.], chickpea [Cicer arietinum L.], lentil [Lens culinaris Med.], albus lupin [Lupinus albus L.], dwarf chickling [Lathyrus cicera L.], ochrus chickling [Lathyrus ochrus L.], grass pea [Lathyrus sativus L.], narbon bean [Vicia narbonensis L.], common vetch [Vicia sativa L.], and purple vetch [Vicia benghalensis L.]) were examined on fine textured neutral to alkaline soils in the low to medium rainfall Mediterranean-type environments in south-western Australia at Merredin and Mullewa in two seasons. There was no difference in the total E_t between grain legumes at either site in either year. There was also no variation in soil water extraction between species on the shallow sandy loam soil at Merredin. However, C. arietinum, L. sativus and L. cicera had greater water extraction and P. sativum the least water extraction at Mullewa where soil conditions were less hostile and root penetration was not restricted. The pattern of water use varied markedly between the grain legume species examined. Grain yield was positively correlated with post-flowering water use (E_tpa) in both erect (r=0.59) and prostrate (r=0.54) grain legume species. Water use efficiencies for dry matter production (WUE_dm) of up to 30 kg ha⁻¹ mm⁻¹ for V. faba and V. narbonensis at Merredin, and water use efficiencies for grain yield (WUE_gr) of up to 16 kg ha⁻¹ mm⁻¹ for P. sativum and 13 kg ha⁻¹ mm⁻¹ for V. faba at Mullewa, were comparable to those reported for cereals and other grain legumes in previous studies in this and other environments. Potential transpiration efficiencies (TE) of 15 kg ha⁻¹ mm⁻¹ together with soil evaporation (E_s) values of 100–125 mm were estimated in this and associated studies, and can be used as benchmark values to assess the yield potential of cool season grain legume crops in low rainfall Mediterranean-type environments. The major traits of adaptation for grain legume species producing large yields in this short season environment are early flowering, and pod and seed set before the onset of terminal drought. Early phenology together with rapid ground cover and dry matter production allows greater water use in the post flowering period. This leads to greater partitioning of dry matter into seed, which is reflected in greater harvest index (HI) and WUE_gr, as was observed for V. faba and P. sativum. Improvement in the adaptation of other grain legume species to short season Mediterranean-type environments requires increased early growth for rapid ground cover and improved tolerance to low temperatures (especially for C. arietinum) during flowering and podding.     

Variations in Sugar Content and Dry Matter Distribution in Roots and their Associations with Frost Tolerance in Certain Forage Legume Species

Enhancement of cold tolerance is an important aspect in breeding forage legumes in view of increasing interest in extending the cultivation of these crops. Three classes of characters — morphological, morpho-physiological and biochemical — were considered in selection for cold-tolerance between and within forage legume species under contrasting growing conditions in the field and a plastic house.
Significant correlation was found between cold damage and concentration of sugars m roots of various forage legume species which were grown under field conditions and subjected to natural frost. Both morphological as well as morpho-physiological characters were related to cold tolerance in Lathyrus ochrus (L.) DC. This indicated that chemical constituents cannot be used as a sole criterion to select for winter-hardy strains of this species. Root-shoot ratio showed a consistent relationship with tolerance to cold irrespective of the growing conditions and season. Hence, sugar concentration and root-shoot ratio are useful characters in testing species found cold tolerant by other methods. The root-shoot ratio could be used for testing cold tolerance under controlled conditions. The number of primary branches per plant, root weight and shoot weight may be used as morphological characters in selection for cold tolerance in L. ochrus under field conditions.     

Demand-driven breeding of food legumes for plant-nutrient relations in the tropics and the sub-tropics: serving the farmers; not the crops!

A number of improved cultivars of food legume crops have been developed and released in the tropics and the sub-tropics. Most of these cultivars have been developed through conventional breeding approaches based on the development of crop varieties under optimum soil fertility levels. Nevertheless, it is hardly possible to say that the varietal provisions made by the past approach have been readily accepted, and properly utilized to boost productivity of food legumes grown by resource-poor farmers. The approach itself did not fully appreciate the actual circumstances of the resource-poor farmers where marginal production systems prevail and the poorest farmers could not afford to use cultivars developed under optimum soil fertility level. Therefore, the limitations and strategic implications of past experiences made to develop crop cultivars need to be analyzed in order to formulate better strategies and approaches in the future. The main purpose of this article is to review the efforts made, the technical difficulties associated with the genetic improvement in food legumes as related to plant-nutrient relations, causes of limited breeding success and thereby draw lessons useful to designing future breeding strategies. The scope of nutrient deficiency stress and the approaches to breeding for plant-nutrient relations are discussed and the need for refining the approach and better targeting of the breeding methodologies suggested.     

Relay-intercropped forage legumes help to control weeds in organic grain production

In organic grain production, weeds are one of the major limiting factors along with crop nitrogen deficiency. Relay intercropping of forage legume cover crops in an established winter cereal crop might be a viable option but is still not well documented, especially under organic conditions.Four species of forage legumes (Medicago lupulina, Medicago sativa, Trifolium pratense and Trifolium repens) were undersown in six organic wheat fields. The density and aerial dry matter of wheat, relay-intercropped legumes and weeds were monitored during wheat-legume relay intercropping and after wheat harvest until late autumn, before the ploughing of cover crops.Our results showed a large diversity of aerial growth of weeds depending on soil, climate and wheat development. The dynamics of the legume cover crops were highly different between species and cropping periods (during relay intercropping and after wheat harvest). For instance, T. repens was two times less developed than the other species during relay intercropping while obtaining the highest aerial dry matter in late autumn. During the relay intercropping period, forage legume cover crops were only efficient in controlling weed density in comparison with wheat sole crop. The control of the aerial dry matter of weeds at the end of the relay intercropping period was better explained considering both legumes and wheat biomasses instead of legumes alone. In late autumn, 24 weeks after wheat harvest, weed biomass was largely reduced by the cover crops. Weed density and biomass reductions were correlated with cover crop biomass at wheat harvest and in late autumn. The presence of a cover crop also exhibited another positive effect by decreasing the density of spring-germinating annual weeds during the relay intercropping period.     

Integrating genotype,fungicide, and spray schedule reduced gall (Physoderma) disease progression and enhanced grain yield in faba bean

Journal of Crop Science and Biotechnology - Faba bean is a cool season food legume crop. However, productivity of the crop is constrained by faba bean gall (FBG), which is an emerging disease that...     

Integrated breeding approaches for improving drought and heat adaptation in chickpea (Cicer arietinum L.)

Chickpea (Cicer arietinum L.) is a dry season food legume largely grown on residual soil moisture after the rainy season. The crop often experiences moisture stress towards end of the crop season (terminal drought). The crop may also face heat stress at the reproductive stage if sowing is delayed. The breeding approaches for improving adaptation to these stresses include the development of varieties with early maturity and enhanced abiotic stress tolerance. Several varieties with improved drought tolerance have been developed by selecting for grain yield under moisture stress conditions. Similarly, selection for pod set in the crop subjected to heat stress during reproductive stage has helped in the development of heat‐tolerant varieties. A genomic region, called QTL‐hotspot, controlling several drought tolerance‐related traits has been introgressed into several popular cultivars using marker‐assisted backcrossing (MABC), and introgression lines giving significantly higher yield than the popular cultivars have been identified. Multiparent advanced generation intercross (MAGIC) approach has been found promising in enhancing genetic recombination and developing lines with enhanced tolerance to terminal drought and heat stresses.     

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.     

Resistance to insect pests: What do legumes have to offer?

Summary Insect pests are major problems for all crops, worldwide. In this review we will focus on legumes, which are attacked by a range of insect pests including pod/seed feeders, defoliators and sap feeders. We review the history of breeding for resistance to insect pests in legumes, which has had mixed success, and discuss further opportunities in this area. We also review the extraordinary array of direct and indirect mechanisms contributing to insect defence in legumes, the understanding and exploitation of which offer opportunities for both legume and non-legume crops. There is also good potential to improve insect resistance in legume crops through a detailed understanding of the signaling pathways that regulate induced responses to insect feeding, and recent progress in this area, primarily obtained from non-legume systems, is reviewed. The importance legumes play in farming systems, their wide range of novel chemistry and the emergence of model systems suitable for genomic approaches present opportunities for research in this area strongly linked to breeding programs to help develop legume crops with enhanced insect resistance. CSIRO’s right to retain a non-exclusive, royalty-free licence in and to any copyright is acknowledged.     

Screening techniques and sources of resistance to foliar diseases caused by fungi and bacteria in cool season food legumes

Screening techniques are an important component of the overall strategy of breeding for resistance to diseases in cool season food legumes. Suitable screening methods have been developed for several major foliar diseases of chickpea, pea, faba bean, and lentil, and sources of resistance have been identified. International cooperation plays an important role in promoting research and keeping collections of cultivated species and their wild relatives. New biotechnological approaches are promising for enhancing the practical use of genes for resistance.     

Integrated gene resource management of underutilized legumes in India

The Indian gene centre possesses a rich legume biodiversity––1,152 species comprising cultivated, underutilized edible and forage legumes. Majority of the underutilized food legumes are widely distributed as wild species in various agro-ecological regions of peninsular India. Indian legume species (62%) contribute to the food and health security of ethnic communities. A total of 66,546 accessions of legume gene resources including underutilized species are conserved in the National Gene Bank. Collection, characterization and conservation efforts regarding the diversity of these beans are described. The importance of genetic variation in legumes and their wild relatives as a source of desirable resistance to pests and diseases in a changing climate scenario is discussed. Information on legumes used in Indian and modern systems of medicine and ethno-botany as well as the scope for bio-prospecting are presented. Advanced biotechnological applications in legume research for sustainable utilization of these resources are highlighted. An integrated gene resource management strategy to combat malnutrition, identify gene resources for legume improvement and enhance their value as traditional food and medicine is described.     

Biological nitrogen fixation in three long-term organic and conventional arable crop rotation experiments in Denmark

Biological nitrogen (N) fixation (BNF) by legumes in organic cropping systems has been perceived as a strategy to substitute N import from conventional sources. However, the N contribution by legumes varies considerably depending on legumes species, as well as local soil and climatic conditions. There is a lack of knowledge on whether the N contribution of legumes estimated using short-term experiments reflects the long-term effects in organic systems varying in fertility building measures. There is also limited information on how fertilizer management practices in organic crop rotations affect BNF of legumes. Therefore, this study aimed to estimate BNF in long-term experiments with a range of organic and conventional arable crop rotations at three sites in Denmark varying in climate and soils (coarse sand, loamy sand and sandy loam) and to identify possible causes of differences in the amount of BNF. The experiment included 4-year crop rotations with three treatment factors in a factorial design: (i) rotations, i.e. organic with a year of grass-clover (OGC), organic with a year of grain legumes (OGL), and conventional with a year of grain legumes (CGL), (ii) with (+CC) and without (−CC) cover crops, and (iii) with (+M) and without (−M) animal manure in OGC and OGL, and with (+F) mineral fertilizer in CGL. Cover crops consisted of a mixture of perennial ryegrass and clover (at the sites with coarse sand and sandy loam soils) or winter rye, fodder radish and vetch (at the site with loamy sand soil) in OGC and OGL, and only perennial ryegrass in CGL at all sites. The BNF was measured using the N difference method. The proportion of N derived from the atmosphere (%Ndfa) in aboveground biomass of clover grown for an entire year in a mixture with perennial ryegrass and harvested three times during the growing season in OGC was close to 100% at all three sites. The Ndfa of grain legumes in both OGL and CGL rotations ranged between 61% and 95% depending on location with mostly no significant difference in Ndfa between treatments. Cover crops had more than 92% Ndfa at all sites. The total BNF per rotation cycle was higher in OGC than in OGL and CGL, mostly irrespective of manure/fertilizer or cover crop treatments. There was no significant difference in total BNF between OGL and CGL rotations, but large differences were observed between sites. The lowest cumulated BNF by all the legume species over the 4-year rotation cycle was obtained at the location with sandy loam soil, i.e. 224–244, 96–128, and 144–156 kg N ha⁻¹ in OGC, OGL and CGL, respectively, whereas it was higher at the locations with coarse sand and loamy sand soil, i.e. 320–376, 168–264, and 200–220 kg N ha⁻¹ in OGC, OGL and CGL, respectively. The study shows that legumes in organic crop rotations can maintain N₂ fixation without being significantly affected by long-term fertilizer regimes or fertility building measures.     

Potential for wild species in cool season food legume breeding

Wild species which are crossable to cultivated pea, lentil, and chickpea have been collected and are maintained in major germplasm collections throughout the world. Wild species of Vicia crossable to the cultivated faba bean have not been found. The primary, secondary, and tertiary gene pools of the cool season food legumes represent potential genetic diversity that may eventually be exploited in cultivated types to overcome biotic and abiotic stresses. Technical difficulties in obtaining hybrids beyond those within the primary gene pool is a major obstacle. Reproductive isolation, embryo breakdown, hybrid sterility, and limited genetic recombination are major barriers to greater use of wild germplasm. Conventional crossing has been successful in producing interspecific hybrids in Lens, Cicer and Pisum and those hybrids are being evaluated for desired recombinants. In vitro culture of hybrid embryos has been successful in overcoming barriers to wider crosses in Lens. The successful transfer of genes from wide sources to cultivated types can be assisted by repeated backcrossing and selection designed to leave behind undesired traits while transferring genes of interest. Molecular marker assisted selection may become a valuable tool in the future use of wild species. In general, too little is known about the possible genetic variation available in wild species that could be valuable in developing resistance to biotic and abiotic stresses. Current efforts on the use of wide hybridization in the cool season food legumes are reviewed and discussed.     

Standorterhebungen zur Stickstoffdynamik nach Anbau von Körnerleguminosen

Field studies on nitrogen dynamics after cultivation of grain legumes Field trials were conducted in order to study the nitrogen dynamics in soil after cultivation of grain legumes and to investigate the possibility of reduction of nitrate leaching due to catch crops or suitable following crops. Accordingly, in 1989/90 soil samples were taken on 12 farms at depths of 0–80 cm in 4 week intervals and analysed for NO₃-N. Furthermore, Brassica napus and Sinapis alba were sown after grain legumes on two farms, and at the experimental station Roggenstein field trials were carried out with different catch crops (Sinapis alba, Raphanus sativus, Lolium multiflorum and Pisum sativum) after grain peas. Considerable amounts of nitrogen (100–150 kg N/ha) in the form of crop residues (straw and grains) were left on the fields cultivated with grain legumes. After harvesting, nitrate content in the soil layer 0–80 cm was on grain legume fields almost twice as high as on fields cultivated with winter wheat. During autumn, the soil nitrate contents increased remarkably. In the soil layer 0–80 cm the maximum values rose to 140 kg N/ha after peas, to 120 kg N/ha after faba beans and only to 65 kg N/ha after winter wheat. The more intensive N-mineralization after peas compared to faba beans is due to a lower C/N-ratio of crop residues and an earlier harvest time of 2-3 weeks of peas. In winter extremely high N-leaching was measured on fallow land after cultivation of grain legumes. Cultivation of catch crops makes it possible to retain up to 110 kg N/ha in plant material. Raphanus sativus and Sinapis alba are most suitable for this purpose due to their high N-uptake even when they are sown late. Ploughing up catch crops in autumn results in a fast mineralization of their immobilized nitrogen. This implies the risk of N-leaching into deeper soil layers during winter, depending on the amount of rainfall and water capacity of the soil. Particularly on soils with low water capacity, early N-mineralization needs to be prevented by cultivating catch crops which freeze off or survive in winter. Cultivation of Brassica napus (winter form) after grain legumes leads to an extensive uptake of soil nitrate before the beginning of the seepage period, and therefore almost excludes enhanced N-leaching.     

Advances in “omics” approaches to tackle drought stress in grain legumes

Grain legumes being affordable sources of proteins, vitamins and essential micronutrients are key to human nutrition worldwide. However, frequent drought episodes present serious threat to grain legume production worldwide. Advances in legume omics in concert with evolving phenotyping and breeding techniques hold great promise to improve drought response of these crops. These resources could underpin prebreeding efforts to expedite discovery and deployment of novel drought tolerance traits into elite backgrounds. Fast-track transfer of traits that confer drought tolerance using marker technologies has been demonstrated in grain legumes like chickpea. However, complex genetic architecture of drought tolerance demands embracing more efficient tools like genomic selection (GS) for accelerated trait improvement. Recent studies on GS for addressing complex traits like drought tolerance have yielded encouraging results in these crops. Recently, speed breeding (SB) protocols have also been optimized for the improvement of long-day/day-neutral grain legumes. Efficacy of SB protocols with regard to complex traits awaits further evidences though. There remains immense scope for integrating SB with GS and gene editing to deliver drought-tolerant cultivars.