Exploring the natural variation for reproductive thermotolerance in wild tomato species

Climate change has become a serious threat for crop productivity worldwide. The increased frequency of heat waves strongly affects reproductive success and thus yield for many crop species, implying that breeding for thermotolerant cultivars is critical for food security. Insight into the genetic architecture of reproductive heat tolerance contributes to our fundamental understanding of the stress sensitivity of this process and at the same time may have applied value. In the case of tomato (Solanum lycopersicum), germplasm screenings for thermotolerance have often used yield as the main measured trait. However, due to the complex nature of yield and the relatively narrow genetic variation present in the cultivated germplasm screened, there has been limited progress in understanding the genetic basis of reproductive heat tolerance. Extending the screening to wild accessions of related species that cover a range of climatic conditions might be an effective approach to find novel, more tolerant genetic resources. The purpose of this study was to provide insight into the sensitivity of individual reproductive key traits (i.e. the number of pollen per flower, pollen viability and style protrusion) to heat-wave like long-term mild heat (LTMH), and determine the extent to which genetic variation exists for these traits among wild tomato species. We found that these traits were highly variable among the screened accessions. Although no overall thermotolerant species were identified, several S. pimpinellifolium individuals outperformed the best performing cultivar in terms of pollen viability under LTMH. Furthermore, we reveal that there has been local adaptation of reproductive heat tolerance, as accessions from lower elevations and higher annual temperature are more likely to show high pollen viability under LTMH.     

Construction of a crop—wild hybrid population for broadening genetic diversity in cultivated sunflower and first evaluation of its combining ability: the concept of neodomestication

The genetic base of sunflower elite lines is very narrow, due to many years of selection and breeding. To broaden the genetic diversity of the cultivated sunflower, in 1995 73 wild sunflower populations were crossed with 3 cultivated lines (Testers), and 219 hybrid offspring’s were evaluated in the field. GCA and SCA effects were computed suggesting for all traits a genetic potential for improvement through selection. Study of the hybrids revealed that the wild accessions bear different genetic abilities to combine with the testers for traits of morphological architecture, phenology and yield (seed weight and seed oil). The variance due to GCA and SCA showed that gene action was additive for days to flowering, branching and plant height. Genotypes derived from the same geographic origin may have either good or poor general combing ability. The correlation between GCA and per se genotype performance was positive for all traits except for seed oil content. This was the first attempt to evaluate wild-cultivated hybrids in sunflower on a large scale and will be the starting point for the management of hybrid Helianthus annuus populations for breeding. GCA and SCA estimations will facilitate the definition of strategies to manage and exploit the natural diversity for this crop.     

Gene flow between cultivated sunflower and <Emphasis Type="Italic">Helianthus petiolaris</Emphasis> (Asteraceae)

Helianthus petiolaris (Asteraceae) native to North America has naturalized in Argentina. The extensive overlapping with sunflower crop regions, the coincidence of life cycles and the common pollinators facilitate interspecific crosses with sunflower, H. annuus var. macrocarpus. To estimate the occurrence of crop-to-wild and wild-to-crop gene flow, off-type plants in progenies of pure stands of both species flowering in coincidence and the presence of crop alleles in H. petiolaris populations were investigated in 26 wild populations and nine crop fields. Morphological traits and RAPD markers were used to attempt hybrid characterization. Off-type individuals were found in frequencies of 0.5 and 0.3% among crop progenies and wild populations, respectively. Off-type plants showed intermediate values for metric morphological traits and low fertility traits. Some off-type plants proved to carry crop alleles based on molecular analysis. The average frequency of cultivar-marker introgression across the wild populations was very low (0.02). Although observed hybridization rates seem to be low, the extension of crop-wild overlapping in Argentina make hybrid formation a noticeable process. Therefore, the new sunflower varieties and eventually GM varieties would transfer their traits through pollen flow and they would persist in H. petiolaris populations.     

Characterization of common bean wild populations for their in situ conservation in Northwestern Argentina

In situ conservation of wild species is a method of conservation that allows keeping populations in their natural environments, and set the strategies for maintaining the natural populations. The Active Bank of Northwestern Argentina (BANOA) is in charge of the in situ conservation of wild populations of common bean (Phaseolus vulgaris L.) in Northwestern Argentina (NOA), and has an ex situ collection of 401 landraces and 221 wild accessions from the NOA. We evaluated the phenotypic diversity of 68 common bean wild populations from the NOA both in protected and unprotected areas, finding a moderate variation among them. Ten phenotypic reproductive characteristics related to pod and seed displayed significant differences in the analysis of variance; these traits together with the seed weight were the basis for the multivariate analysis. The cluster analysis ordered the populations in 12 groups but trends in geographical distribution or phenotypical variation were not recognized. For the conservation in situ of the wild bean populations, their diversity should be considered. Two types of populations can be highlighted: (i) candidates for in situ conservation in order to preserve the novel variation generated by convergence with cultivated sympatric germplasm (populations 433, 437, 471, 509, 513 and 517) and (ii) those whose phenotype represents clearly the wild status and should be preserved in situ as such in their current status (populations 480, 495, 496, 525 and 533).     

Crossability between cultivated and wild tuber-and non-tuber-bearing Solanums

The cultivated potato has over 200 extant wild relatives many of which contain genes valuable for disease resistance, hardiness, processing and agronomic traits. Crossability of these wild species directly with the cultivated potato is complicated by several reproductive phenomena such as stylar and ploidy barriers and Endosperm Balance Numbers (EBN). However, a systematic analysis of crossability with many of these wild relatives has never fully been examined. Reciprocal crosses were made between cultivated potato and over 400 wild potato accessions; stylar barriers and 2n gamete production were examined as was the fertility of many of the putative hybrids. Generally, the seed/fruit ratio increased the more closely related the species were to the cultivated potato. However, a few crosses were successful in spite of predicted failure due to ploidy or EBN differences. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of acclimation on chilling tolerance in Asian cultivated and wild rice

Tolerance to low temperature can be enhanced by pre-exposure to low, non-harmful temperatures, a process known as acclimation. The effects of acclimation on chilling tolerance in species from tropical and subtropical regions are not well understood. Therefore, we investigated whether acclimation improved chilling tolerance at the plumule and seedling stages in Asian rice. A comparison of the effect of acclimation between chilling tolerance scores from a tolerant (A58) and a susceptible (W107) strain demonstrated that acclimation enhances chilling tolerance especially in the tolerant strain, indicating that considerable genetic variation in acclimation capacity exists. The genetic variation in chilling tolerance in acclimated plants was investigated using 57 strains derived from cultivated (Oryza sativa L. ssp. japonica and ssp. indica) and wild (O. rufipogon Griff.) rice. The genetic differentiation of the japonica and indica subspecies of cultivated rice with regard to chilling tolerance was more prominent in the presence than in the absence of acclimation. Furthermore, latitudinal clines for chilling tolerance were observed in both acclimated plumules and seedlings of wild rice, whereas without acclimation, a latitudinal cline was observed only at the plumule stage. This suggests that acclimation capacity, as well as intrinsic chilling tolerance, might contribute to local adaptation. QTLs involved in acclimated chilling tolerance at the plumule stage are different from those involved in intrinsic chilling tolerance. The present results indicate that the maximum genetic potential for chilling tolerance is achieved by acclimation, giving cautions for evaluation of genetic resources in rice.     

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.     

Vigour/tolerance trade‐off in cultivated sunflower (Helianthus annuus) response to salinity stress is linked to leaf elemental composition

Developing more stress‐tolerant crops will require greater knowledge of the physiological basis of stress tolerance. Here, we explore how biomass declines in response to salinity relate to leaf traits across 20 genotypes of cultivated sunflower (Helianthus annuus). Plant growth, leaf physiological traits and leaf elemental composition were assessed after 21 days of salinity treatments (0, 50, 100, 150 or 200 mM NaCl) in a greenhouse study. There was a trade‐off in performance such that vigorous genotypes, those with higher biomass at 0 mM NaCl, had both a larger absolute decrease and proportional decrease in biomass due to increased salinity. More vigorous genotypes at control were less tolerant to salinity. Contrary to expectation, genotypes with a low increase in leaf Na and decrease in K:Na were not better at maintaining biomass with increasing salinity. Rather, genotypes with a greater reduction in leaf S and K content were better at maintaining biomass at increased salinity. While we found an overall trade‐off between sunflower vigour and salt tolerance, some genotypes were more tolerant than expected. Further analysis of the traits and mechanisms underlying this trade‐off may allow us to breed these into high‐vigour genotypes in order to increase their salt tolerance.     

Morphological differences among Raphanus raphanistrum populations and their relationship to related crops

Wild radish (Raphanus raphanistrum) has developed introgressed populations after hybridization with its cultivated counterpart (R. sativus) in California. Hybridization with various Brassica and Sinapis species is also possible. To determine if hybridization is responsible of the genetic diversity of European populations, six wild radish populations with distinct morphological traits were sampled from geographically distant regions in Europe. Plants were cultivated in an oilseed rape field and in insect‐proof cages. Silique and flower morphology, growth, and reproductive traits were measured. The wild radish populations could be discriminated by the morphological traits, but not related to geographic regions. In particular, populations of one region showed wide variability in terms of silique shape and growth behaviour, and small‐sized flowers. Although the origin of morphological diversity in wild radish is unclear, i.e. native or due to gene flow from the cultivated radish or other Brassicaceae, significant morphological divergence was found that could have relevant effects on plant ecology and adaptation.     

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

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

Phaseolin Diversity in the Tepary Bean, Phaseolus acutifolius A. Gray

The tepary bean (Phaseolus acutifolius A. Gray) is grown mainly in and regions of Mexico and the southwestern U.S. as a subsistence crop by small farmers. It is also a store of genetic variability for traits such as disease and pest resistance and stress tolerance to improve the common bean (P. vulgaris L.). To determine geographic patterns of variation and the influence of domestication on genetic variability, the genetic diversity of phaseolin, the major seed storage protein, was characterized among 55 wild and 8 cultivated teparies using polyacrylamide gel electrophoresis. Fifteen electrophoretic phaseolin patterns were identified among wild forms, whereas only one pattern was exhibited by cultivars. This result suggests a single domestication in this species leading to a strong reduction in diversity. An additional finding is the divergence m phaseolin types between populations east and west of the Sierra Madre Occidental mountains.     

Development of drought‐tolerant breeding lines derived from Helianthus annuus × H. argophyllus interspecific crosses

Helianthus argophyllus is a wild species known as “silver sunflower”, which possesses several traits, including morphological traits that increase drought tolerance. Therefore, introgressions between chosen cultivated lines and two H. argophyllus accessions were made, and segregating generations were established. Important agronomic traits including single heading, high pollen fertility, silver canopy (indicating more cuticular wax), and content of cuticular waxes and oil were selected over six segregating generations. The resulting F₆ lines showed introgression of water saving traits, as they had lower excise leaf water loss, with comparable yield to standard checks. However, these F₆ lines were late maturing and showed poor flowering synchronization between the cultivated and introgressed lines. Introgressed line “D‐22” was particularly promising as a breeding line, with superior agronomic and drought resistance traits. This line had the potential to be used as an inbred parental line for introgression of drought resistance traits into elite sunflower germplasm. Combining ability analysis of the introgressed lines further showed their potential for heterosis breeding or to be used as parental lines in breeding programme.     

Genetic resources of salt tolerance in wild Hordeum species

Salt tolerance was evaluated in 340 accessions of Hordeum, consisting of 41 brittle-rachis forms of Hordeum vulgare L. subsp. vulgare (H. agriocrithon) accessions, 154 H. vulgare L. subsp. spontaneum (H. spontaneum) accessions, and 145 accessions of ten other species or subspecies of wild Hordeum. Germination was carried out at concentrations of 171, 257, and 342 mM NaCl. The levels of salt tolerance for seed germination in wild Hordeum species were generally lower than those found by Mano et al. (1996) in cultivated barley; the NaCl tolerance level of the different species were as follows: H. agriocrithon > H. spontaneum > other wild Hordeum species. In addition, leaf injury index was used to assess tolerance at the seedling stage after treatment with 500 mM NaCl solution for four weeks. The levels of salt tolerance at the seedling stage in wild Hordeum species were generally higher than those found by Mano & Takeda (1995) in cultivated barley. Most wild Hordeum species showed high NaCl tolerance at the seedling stage and are considered good sources of germplasm for salt tolerance breeding. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Evidence of a partial reproductive barrier between wild and cultivated pearl millets (Pennisetum glaucum)

Summary The occurrence of seed malformation in association with reduced thousand grain weight and germination ability has been observed in crosses between cultivated female plants and wild male plants. A survey of 16 cultivated accessions (P. glaucum subsp. glaucum) and 11 wild accessions (P. glaucum subsp. monodii) ranging over the whole species diversity showed this postzygotic incompatibility was general, but its intensity varied greatly with the cultivated female accession used and very little with the wild male parent origin. About 15% of the 123 cultivated x wild crosses observed gave normal seeds. Seed malformation has never been observed in crosses between cultivated accessions and appeared independent of genetic distances between the parents. The reciprocal crosses between wild female plants and cultivated male plants gave normal-looking seeds with good germination but consistently reduced thousand grain weight. Both seed malformation and seed small size are an expression of a genetic imbalance. These slight reproductive barriers seem to have been built during the domestication process.Abbreviation ICRISAT International Crop Research Institute for the Semi-Arid Tropics     

A non-invasive crop ideotype to reduce invasive potential

Summary In plant breeding programs, qualitative and quantitative traits confer market value and, thus, constitute the basis for developing breeding criteria during crop domestication. Some traits such as high male/female fertility are advantageous in the wild and could enable the evolution of cultivated crops into invasive weeds. Other traits, e.g. sterility, are not expected to confer invasiveness. To date there has been very limited involvement in invasion risk assessment by plant breeders. Thus, in this paper we propose that trait-based selection of potential crop species be coupled with species design in the creation of a “non-invasive crop ideotype” as an avenue to reduce invasiveness during domestication. The non-invasive crop ideotype embodies the ideal characteristics for a crop to excel in cultivated environment(s) but minimizes the likelihood it will establish and spread in non-cultivated environments, constituting the underlying foundation for all breeding objectives, choice(s) of breeding methodologies, and propagation techniques for non-invasive crop release. Using ornamental (floriculture) horticultural crops as an example, we identify 10 traits to be used individually or in combination to reduce invasiveness while retaining commercial value: reduced genetic variation in propagules, slowed growth rates, non-flowering, elimination of asexual propagules, lack of pollinator rewards, non-shattering seed, non-fleshy fruits, lack of seed germination, sterility, and programmed death (apotopsis). A non-invasive crop ideotype would constitute the underlying foundation for all breeding objectives, the choice(s) of breeding methodologies, and propagation technique(s). The ideotype should be flexible and should adjust to species- and crop-specific traits to account for the intended use. For example, development of sterile cultivars may have negligible effects in reducing invasiveness if the crops spread vegetatively. A non-invasive crop ideotype may increase the direct participation of plant breeders, who are the professionals directly involved in the collection, development and release of new crops, in reducing the invasive potential of ornamental crops. Future research is required to determine the feasibility of incorporating each trait into various crops, use of classical or molecular techniques for creation of non-invasive crops, trait stability (lack of genotype × environment interaction over years and locations), consumer acceptance, and long-term viability.     

The Inheritance of Earliness and Fruit Weight in Crosses between Cultivated Tomatoes and two Wild Species of Lycopersicon

The inheritance of earliness and fruit Weight in tomato was studied in tour interspecific crosses. Two cultivated varieties, i.e.‘HS 101’ and “HS 102′, were hybridized with two wild species. L. birsutum f. glabratum‘B 6013’ and L. pimpinellifolium‘A 1921′. Six generations of these crosses were evaluated for these traits and the estimates of gene effects were derived from the generation mean using an epistatic (six parameters) model. There were very wide differences between cultivated and wild species for earliness and fruit weight and in the segregating populations, plants with delayed maturity and smaller fruit size were recorded with high frequency. It was found that the inheritance pattern was mainly governed by additive gene action. Epistatic effects also contributed towards the inheritance of both traits.     

Comparative assessment of genetic diversity between wild and cultivated barley using gSSR and EST‐SSR markers

Barley is an economically important cereal crop especially for feed and malt production, but its value as food is increasing due to various health benefits. Wild barley is the progenitor of modern day barley cultivars possessing a rich source of genetic variation for various biotic and abiotic stresses. Species‐specific molecular markers have great potential for efficient introgression of these important traits from wild to cultivated barley. In the present study, 140 microsatellite markers were screened to assess the genetic variation and species‐specific markers between wild and cultivated germplasm. Of these 140, a polymorphic set of 48 genomic (gSSR) and 16 EST‐SSRs amplified a total of 685 alleles. Cluster analysis discriminated all 47 accessions and classified wild and cultivated genotypes into two distinct groups, according to their geographic origin. Our analysis indicated that gSSRs were more informative than EST‐based SSRs. Results from PCoA analysis for species‐specific alleles clearly suggest that wild barley genotypes contain a higher number of unique alleles.     

Reproductive stage tolerance to salinity and alkalinity stresses in rice genotypes

Salinity and alkalinity (sodicity) seriously threaten rice production in south Asia. Improving screening methodologies for identifying sources of tolerance is crucial for breeding salt tolerant rices. Rice genotypes of varying tolerance (tolerant, semi‐tolerant and sensitive) were screened in saline soil of electrical conductivity, ECe 4 and 8 dS/m and alkali soil of pH 9.5 and 9.8 in lysimeters. Vegetative growth events were less affected by both the stresses in comparison to reproductive stage. Grain yield was reduced by 26.7%, 45.7% and 50.3% at ECe 8 dS/m in three tolerance groups respectively. At pH 9.8 the reduction was 25.1%, 37.2% and 67.6% in the three groups respectively. Higher floret fertility contributed to higher seed set and grain yields in tolerant genotypes whereas higher spikelet sterility led to poor seed set and lower grain yields in sensitive genotypes. The 1000 grains weight was also significantly reduced at ECe 4 or pH 9.8. Screening at reproductive stage for morphological traits like floret fertility is thus more useful to identify rice genotypes tolerant to both salinity and alkalinity stress. Genotypic (G) and environmental (E) effects and GE interactions were highly significant for the growth attributes and grain yield. Based on analysis of variance, genotypes tolerant to salinity and alkalinity as well to both the stresses were identified.     

Response of cotton genotypes to water and heat stress: from field to genes

Cotton is a crop of tropical and subtropical regions but the seed cotton yield is highly influenced by abiotic stresses like drought and heat. Response of cotton genome to abiotic stresses is highly complex and involve many genes. A comprehensive study, involving cotton genotypes developed through conventional and synthetic tetraploid method, was designed to (i) study the introgression of heat and water stress tolerance by using wild relatives (ii) evaluate genetic markers for marker assisted selection against water and heat stress. Two separate experiments for water and heat stress tolerance with a common control were established. Treatments in each experiment include a control and a stress treatment. Heat stress was applied by sowing crop two month earlier than the control treatment, whereas water stress was imposed by withholding alternate irrigation. Analyses of variance depicted highly significant (P ≤ 0.01) effect of genotypes and both stresses on boll retention, boll weight and seed cotton yield. Interaction of genotypes with stress in both experiments was also highly significant (P ≤ 0.01). Genotypes derived from interspecific crosses performed consistently in stress conditions compared to control which prove it a reliable method to introgress stress related genes from wild parents. Four genes reported for water stress tolerance and five genes reported for heat stress tolerance were evaluated by field results for efficient marker assisted selection (MAS). Results verified drought stress genes but heat stress genes could not explain genetic variability caused by heat stress. It is concluded from the results that separate genes may be responsible for heat stress tolerance for vegetative and reproductive stages, therefore, selection criteria should include both the traits.