Genetic resources in breeding for adaptation

Summary A crop's ability to productively exploit its environment depends on many adaptive features which are controlled by multiple genes, interacting among themselves and with the environment in complex ways.To promote widespread adoption, breeders frequently seek to develop broad adaptation in their varieties, often through the use of genes having a large effect on a single adaptive feature. Such genes may occur within the crop, its wild relatives, or unrelated taxa. Genes for many adaptive features (e.g. temperature tolerance) may be found in extreme environments. Others (e.g. photoperiod insensitivity) may have evolved away from primary centres of origin.Well characterized and documented ex situ germplasm collections aim to serve plant breeders' need for genes. Molecular marker and geographic information system (GIS) techniques are proving useful for locating and characterizing genetic diversity. New techniques (e.g. core collections and electronic information systems) are adding to the value of collections. Ex situ and in situ conservation of wild crop relatives are receiving increased attention. With all organisms becoming a potential source of genes for breeding, in situ ecosystem conservation is assuming added significance.Farmers, particularly in diverse, marginal environments in developing countries, continue to breed landraces adapted to their specific circumstances. In areas of high ecological diversity, a multitude of adaptive gene complexes have been selected within small geographic areas. Conventional breeding frequently neglects such farmers, and participatory methods based on locally adapted diversity, coupled with appropriate technical and policy support, may prove more effective in meeting their needs. Such dynamic, on-farm conservation and management systems would also enable genetic diversity to continue to evolve as a resource for conventional breeding.For genetic resources to remain a foundation for future sustainable agricultural development, complementary conservation and breeding strategies are needed.     

Isozyme variation in a number of populations of Theobroma cacao L. obtained through various sampling regimes

Summary Isozyme polymorphism in four enzyme systems was used to estimate the genetic diversity in nine populations of Theobroma cacao L. The distribution of allelic variation observed in this study was inconsistent with the currently held view that the Upper Amazon region of Peru is the centre of cacao genetic diversity. All the populations analysed from this area of Peru were genetically similar, and contained a low level of isozyme polymorphism. It is argued that this is a result of genuine low diversity in the region rather than a sampling artifact. If a centre of diversity of wild cacao truly exists, this work indicates that it lies further north in Ecuador and Colombia.Although they are morphologically highly variable, Trinitario cacao populations derived from cultivation are currently accorded low priority for conservation. This phenotypic variation was observed to be reflected in high levels of isozyme polymorphism. It is suggested that the present strategy for the conservation of cacao germplasm, which accords priority to the Upper Amazon region of Peru while neglecting the Carribbean should be reconsidered.     

Participatory crop improvement and formal release of <Emphasis Type="Italic">Jethobudho</Emphasis> rice landrace in Nepal

Jethobudho is an aromatic rice landrace of the Pokhara valley in middle hills of Nepal. Although local consumers are willing to pay a high price for its purchase, the landrace has a problem with quality variation. Decentralized participatory population improvement for specific market-identified traits was conducted on “Jethobudho” populations collected from farmers’ fields in seven geographic regions of the valley in Nepal. The preferred post harvest quality traits, field tolerance to blast and lodging, and superior post harvest quality traits of Jethobudho were established by a consumer market survey. These traits were used for screening the materials. 338 sub-populations of Jethobudho were evaluated for yield, disease, lodging resistance, and post harvest quality traits. Significant variation was found for culm strength, neck blast tolerance, awn characteristics, panicle length, number of grains per panicle, test grain weight and post harvest quality traits, whereas no significant variation was found in grain yield, plant height, tiller number, maturity period and leaf blast. Based on these identified traits and micro-milling evaluations, 183 populations were screened in on-farm and on-station nurseries, and in succeeding years populations were further screened by plant breeders and expert farmers in research trials, resulting in the selection of 46 populations for post harvest quality traits. Six accessions with similar agronomic traits, field tolerance to blast and lodging, and superior post harvest quality traits, were bulked and evaluated on-farm using participatory variety selection (PVS). The enhanced Jethobudho accessions were also evaluated for aroma using simple sequence repeat (SSR) and found to have unique aromatic genetic constitution. Community based seed production groups were formed, linked to the Nepal District Self Seed Sufficiency Programme (DISSPRO), and trained to produce basic seeds (truthfully labeled) of Jethobudho. The National Seed Board of Nepal released the enhanced landrace in the name of “Pokhareli Jethobudho” in 2006, as the first bulk variety of traditional high quality aromatic rice improved through participatory plant breeding to be formally released in Nepal for general cultivation under the national seed certification scheme. Landrace improvement is shown as an important option for supporting programmes for in situ conservation of landraces on-farm.     

Rapid estimation of broad sense heritability of farmer-managed maize populations in the Central Valleys of Oaxaca, Mexico, and implications for improvement

A method for rapid estimation of broad senseheritability (H) was applied in farmers' maize fields in two communities inOaxaca, Mexico. Plant and ear traits were documented and H estimatescompared with narrow sense heritability (h²) from family analysis andpublished estimates. Absolute values of H estimates were larger thanh² estimates from this study and the literature. Relative ranking bytrait was the same as in the literature, though not the same as rankings ofh² estimates generated in this study. With an understanding of itslimitations, this rapid, economical estimation procedure could provideuseful initial information especially for collaborative crop improvement workbetween researchers and farmers or farming communities on-farm, a plantbreeding context for which little empirical information is available.     

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.     

Genetic divergence among cold tolerant rices (Oryza sativa L.)

Summary Genetic divergence was investigated among 17 rice varieties known to possess some degree of cold tolerance at different growth stages. The 17 varieties and their 102 F₁ hybrids with 6 male testers were studied for isozyme variation at 15 loci, spikelet fertility, and degree of cold tolerance at various stages. Multivariate analyses of the data provided several schemes of divergence based on various sources of evidence. All schemes gave similar results, and separated the varieties into japonica and indica groups. The japonica group displayed specific isozymes, a low F₁ fertility with indica testers, and a high degree of cold tolerance which was expressed in the F₁ progenies. The indica group displayed contrasting specific isozymes, a high F₁ fertility with indica testers, and a moderate to low degree of cold tolerance which was not expressed in the F₁ progenies. One variety, ARC 6000, displayed unique traits in most schemes and was classified into a distinct type based on the isozymes. The results emphasize that cold tolerance is a major trait for classification of rice into two varietal groups and that proper characterization of potential donors is essential in breeding. Isozyme studies are useful tools for this purpose.     

Increasing genetic diversity by participatory varietal selection in high potential production systems in Nepal and India

On-farm varietal biodiversity was studied through household surveys in two high potential production systems in Chitwan and Nawalparasi districts of the Nepal Terai and in Lunawada sub-district, Gujarat, India. Diversity was extremely low in Chaite rice in the Nepal study area (weighted diversity 0.04) and low in main season rice in the India study area (weighted diversity 0.34). In both areas, one cultivar dominated, CH 45 in Nepal and GR11 in India. In the India study area, biodiversity varied with the socio-economic group and better-off farmers had a greater varietal diversity. Participatory varietal selection (PVS) identified new varieties that farmers preferred. Their adoption by farmers increased on-farm varietal biodiversity within the three cropping seasons studied. Despite the commonly assumed uniformity of high potential production systems, the new varieties occupied specific niches in the farming system. Farmers' preferences for different varieties – there were large differences in quality traits and maturity period among the new varieties – should help to maintain biodiversity. Overall production is expected to increase as each niche becomes occupied increasingly by the best-adapted variety. PVS is a simple and powerful method of increasing food production in the high potential production systems that produce most of the developing world's grain. This revised version was published online in August 2006 with corrections to the Cover Date.     

SSR-based genetic diversity assessment among Tunisian winter barley and relationship with morphological traits

For studying genetic diversity caused byselection for adaptation and end-use, 17microsatellites (SSR), representative ofthe barley genome, were used in 26 barley(Hordeum vulgare L.) accessions andcultivars in Tunisia. Theaccessions/cultivars originate fromdifferent geographic regions and are ofdifferent end-use. For the 15 polymorphicSSR, the mean number of alleles per locuswas 3.6 and the average polymorphisminformation content was 0.45. Clusteranalysis based on SSR data and onmorphological data clearly differentiatethe genotypes according to their type(local landraces vs. varieties), row-numberand end-use. The correlation between bothdiversity measures was highly significant(r = 0.25, p<10^-5) and thecorrespondence between the clustering basedon SSR and morphological data wasrelatively good. Our results show the largegenetic diversity of the Tunisian barleycultivars and the association of thisdiversity with adaptation traits.     

Linkage analysis among loci for RAPDs, isozymes and some agronomic traits in Brassica campestris L.

Common bean, Phaseolus vulgaris L. is intercropped or relay cropped with maize in many Andean highlands of Colombia and Peru. Breeding beans for the target multiple cropping systems is essential for the development of productive and sustainable agriculture for the Andean smallholders. Outline of the breeding programme should follow the farming system approach with the establishment of on-farm trials and early farmers involvement. Bean breeding is oriented to minimize intercrop competition and to stabilize complementarity with maize. Genetic traits needed for improved varieties are divided as follows : traits not interacting with the cropping systems, traits specific to intercrops and traits related with socioeconomic and seed quality aspects. Screening, prebreeding and recombination nursery are better made under sole cropping while varietal improvement and on-farm trials are conducted under the target multiple cropping systems. Breeding schemes may involve recurrent, pedigree and bulk hybrid selection. The given application concerns the genetic improvement of P. coccineus, P. polyanthus and interspecific hybrids of P. vulgaris for both simultaneous and relay intercropping in Colombia and Peru. Earliness, cold tolerance, resistance to fungus diseases (mainly Ascochyta leaf blight and anthracnosis) and seed yield potential were the major objectives of the bean improvement programme. Priority has been given to the exploitation of the large diversity available in the secondary gene pool of common bean. This revised version was published online in July 2006 with corrections to the Cover Date.     

Challenges and opportunities for conservation of forest genetic resources

Increased use of forest resources and a shrinking forestland base threaten the sustainability of forest genetic resources and highlight the importance of conservation and sustainable management of these resources. As forest trees are normally the keystone species of forest ecosystems, their continued existence is essential for many floral and faunal associations of these ecosystems. Major concepts, challenges and opportunities for conservation of forest genetic resources are briefly discussed in this paper. The major challenges include population decline and population structure changes due to forest removal and conversion of forest land to other uses,forest fragmentation, forestry practices, climate change, disease conditions,introduced pests, atmospheric pollution, and introgressive hybridization. Developing scientifically sound conservation strategies, maintaining minimum viable population sizes, and deployment of genetically engineered organisms represent other important challenges in conservation. The usefulness of various biochemical and molecular genetic markers, adaptive traits, and genetic diversity measures for developing conservation strategies for in situ and ex situ genetic resource conservation are also discussed. Major opportunities for conservation of forest genetic resources include: use of molecular genetic markers and adaptive traits for developing conservation strategies; in situ conservation through natural reserves,protected areas, and sustainable forest management practices; ex situ conservation through germplasm banks, common garden archives, seed banks, DNA banks, and tissue culture and cryopreservation; incorporation of disease, pest, and stress tolerance traits through genetic transformation;plantation forestry; and ecological restoration of rare or declining tree species and populations. Forest genetic resource conservation and resource use should be considered complementary rather than contradictory to each other. This revised version was published online in July 2006 with corrections to the Cover Date.     

SSR analysis of chromosome 8 regions associated with aroma and cooked kernel elongation in Basmati rice

Aroma and cooked kernel elongation (CKE) are the two most important quality traits, which differentiate the highly valued Basmati rice from other rice types. Previous studies on genetic analysis have shown that genes/QTLs for these two traits are linked and present on chromosome number 8. We have evaluated the genetic diversity in 33 rice genotypes representative of the traditional Basmati (TB), cross-bred Basmati derived from indica × Basmati rice crosses and non-Basmati (indica and japonica) rice varieties for chromosome number 8 using 26 SSR markers including a specific marker (SCU-SSR1) for RG28 locus; the results have been compared with whole genome based SSR allelic data. The 26 SSR markers (24 polymorphic and 2 monomorphic) amplified a total of 106 alleles; 21 of these alleles were detected to be unique, present in only one genotype. The number and size of the alleles, and polymorphism information content (PIC) values ranged between 1–8, 87–312 and 0–0.736 bp, respectively. SCU-SSR1 marker amplified a total of three alleles (128, 129 and 130 bp). All the TB varieties except Basmati 217 (129 bp) and 7/13 cross-bred Basmati varieties had the 130 bp allele. Alleles of 129 and 128 bp were present in majority of the indica and japonica varieties, respectively. The average pair-wise Jaccard similarity coefficients for TB, indica and japonica varieties were 0.512, 0.483 and 0.251, respectively. Average similarity coefficient between TB and japonica was higher (0.236) compared to that between TB and indicas (0.150). Genetic relationships as determined by Principal Component Analysis (PCA, NTSYS-pc), PowerMarker tree, and Structure analyses, clearly showed high-level differentiation between TB and indica rice varieties, which formed two distinct clusters. The cross-bred Basmati and japonica rice genotypes were placed between these two clusters. Basmati 217 and Ranbir Basmati were quite divergent from rest of the TB varieties. Some of cross-bred Basmati varieties including Super, CSR30 and kernel were closer to TB. Indica rice varieties, CSR10 (salt tolerant variety) and Pokkali (salt tolerant landrace) formed a separate distinct cluster. The Pritchard structure analysis divided the rice genotypes in four major sub-populations of TB, cross-bred Basmati, indica and japonica (including Ranbir Basmati and Basmati 217) rice varieties. Chromosome 8 data-set showed a positive correlation (Mantel test, r = 0.739) with the allelic data-set for 30 SSR markers well-distributed on 12 rice chromosomes indicating a higher level of similarity between the two. The study demonstrates the distinctness of TB from other rice types (indica and japonica) and also provides several novel markers for differentiation between TB rice supplies from cheaper cross-bred Basmati and long-grain non-Basmati varieties at commercial level.     

The nabicol: A horticultural crop in northwestern Spain

Nabicol (B. napus L. var. pabularia) is a traditional crop in the Northwest of Iberian Peninsula (South of Galicia and North of Portugal) where it is grown during the winter season on small farms and gardens using traditional varieties. A collection of 36 populations of nabicol from Galicia (northwestern Spain) was evaluated during 2002 and 2003 in two locations and two growing seasons (spring/summer and autumn/spring) for 28 agronomic and morphological traits. The objectives of this study were to: (i) evaluate a collection of nabicol landraces from northwestern Spain, (ii) determine the suitability of this germplasm as a summer crop and (iii) study the genetic diversity among local populations. Significant differences were observed among populations for most traits. Genotype × environment interaction was significant for most of them. Spring/summer growing season could be recommended for growing nabicol but resistance to Lepidoptera pests attacking Brassica crops should be improved. Most populations had an agronomic value similar to the commercial variety. The most promising variety for horticultural use was MBG-BRS0063, which showed the highest yield. Morphological and agronomic data were subjected to cluster analysis and four groups were defined with a group clustering most populations. The low genetic diversity could be explained because populations were collected in close geographical areas and the selection made by farmers was always for a horticultural use. These results give information about the diversity and breeding value of the nabicol Spanish germplasm, which could be useful in breeding programs.     

Isozyme polymorphism in three gene pools of cultivated chicory (Cichorium intybus L.)

Summary Polymorphism of ten enzymes, acid phosphatase (APH), isocitrate dehydrogenase (IDH), leucine aminopeptidase (LAP), phosphorylase (PP), superoxide dismutase (SOD), malic enzyme (ME), glutamate oxaloacetate transaminase 1 and 2 (GOT-1, GOT-2) and phosphoglucomutase 1 and 2 (PGM-1 and PGM-2), was investigated in three gene pools of cultivated chicory, including six cultivated wild chccory, eight industrial chicory and eight Brussels chicory varieties. LAP, APH, PP and PGM-2 showed high phenotypic polymorphism whilst GOT-1 and ME had poor polymorphism. For three enzyme coding loci Lap, Pgm-1 and Got-2, allele frequencies were determined. Isozyme composition in the three chicory gene pools was significantly different, showing, respectively, high, intermediate and poor average amount of phenotypic polymorphism in cultivated wild chicory, industrial chicory and Brussels chicory. Isozyme variation within and between varieties of the three gene pools is discussed in relation to breeding practices.     

Isozyme variation in Helianthus petiolaris and sunflower,H. annuus

Summary Helianthus petiolaris is a wild species used in genetic improvement of sunflower, as a donor of cytoplasmic male sterility and of genetic resistance to diseases. Isozyme variation for ADH, ACP, EST, GDH, LAP, PGI, PGD and SKDH in this species was studied using starch gel electrophoresis. The patterns thus obtained were compared with zymograms of inbred lines, hybrids and open pollinated varieties of H. annuus. The same alleles for EST and SKDH isozymes were found in both species, while ACP showed an allele that has not been found in sunflower. The rest of the isozyme systems showed both common alleles and characteristic ones for each species. ACP, GDH and PGD were monomorphic in H. petiolaris, while ADH and LAP were monomorphic in H. annuus. The isozyme markers obtained here could be useful in breeding programs involving interspecific crosses, and studies on introgression and on genetic variation in other populations of this wild species.     

Contribution of autochthonous maize populations for adaptation to European conditions

Early vigor, earliness and cold tolerance are the main potential contributions of European maize (Zea mays L.) for breeding programs for adaptation to areas with short growing seasons and cold springs. The objective of this research was to determine the potential contributions of populations from different European regions to breeding for adaptation. Six Spanish and six French maize populations differing on variability for earliness, vigor and cold tolerance were crossed in a complete diallel without reciprocals. The populations and their crosses were evaluated in the field and in a cold chamber. Minimum temperatures were the main environmental trait affecting genotype × environment interaction, probably due to the cold sensitivity of the genotypes with the best performance in the field. The best population cross, based on specific heterosis for adaptation-related traits in the field, was Viana × Rastrojero, but this cross was cold sensitive. Tuy × Lazcano should be the best choice for a breeding program for adaptation, based on performance in the field and cold tolerance. As conclusions, there was variability for earliness, vigor and cold tolerance among the populations and crosses involved in this study, being tolerant to cold conditions the populations with medium growing cycle originated in areas with short growing seasons. The highest yielding crosses were cold sensitive.     

Levels and distribution of genetic diversity of coconut (Cocos nucifera L., var. Typica form typica) from Sri Lanka assessed by microsatellite markers

The coconut variety Typica, form typica, commonly known as Sri Lanka tall coconuts is the most widely exploited and grown variety in Sri Lanka. Under the coconut bio-diversity conservation programme, several Typica populations have been collected by island-wide surveys and planted ex situ. Thirty-three coconut populations were subjected to microsatellite assay with eight coconut-specific microsatellite primer pairs in order to study the levels and distribution of genetic variation of the collected materials for formulating future collection strategies and selecting parents for the breeding programme. A total of 56 alleles were detected ranging from 3 to 10 alleles per primer pair with an average of 7 alleles per locus. Overall a very high level of genetic diversity was detected (0.999) for all the populations studied ranging from 0.526 for population Debarayaya to 0.683 for population Dickwella. Only four introduced coconut populations, i.e. Clovis, Margeret, Dickwella, Mirishena and an embryo-cultured population were clearly separated from the resulting dendrogram. A very high level of within population variation (99%)accounted for native populations suggests a common history and a restricted genetic base for native Sri Lankan tall coconuts. Categorization of alleles into different classes according to their frequency and distribution confirmed the results of the dedrogram and concluded the adequacy of single large collection from the entire target area to represent the total genetic diversity in Sri Lanka. This study discusses useful information regarding conservation and breeding of coconut in Sri Lanka. This revised version was published online in August 2006 with corrections to the Cover Date.     

Genetic variation in F<Subscript>2</Subscript> populations and their potential in the improvement of grain yield in tef (<Emphasis Type="Italic">Eragrostis tef</Emphasis>)

Tef is a staple cereal of Ethiopia in high demand by consumers. In order to cope up with this high consumer demand, productivity per unit of land must increase through the development and use of high-yielding varieties. To this effect, the National Tef Research Project has long been striving towards the development of high yielding varieties through direct selection from germplasm and concentrating favourable alleles through hybridization and selection, despite the tedious crossing technique. The objective of this study was to assess the degree of genetic variation in F₂ populations of tef as a basis for improving grain yield. F₂ populations from 12 crosses and their parents were grown at the Debre Zeit Agricultural Research Center, Ethiopia, and assessment was made on eight traits on individual plant basis. Eleven of the 12 crosses showed substantial genetic variation for grain yield and its components, indicating the potential for improvement through selection. Moreover, grain yield, plant weight and yield related traits showed moderate to high heritability values (17–80%). In all the crosses, tiller number, panicle weight, yield per panicle and panicle length showed significant (P ≤ 0.05) and positive association with grain yield. Considering the degree of genetic variation and heritability values, emphasis should be given to selected crosses in an effort to developing high-yielding tef varieties.     

Allozyme differentiation of European populations and cultivars of Lolium perenne L., and the relation to ecogeographical factors

Summary Sixty Lolium perenne populations were screened for allozyme diversity at five loci. Objective was to determine whether allozyme diversity could be used as selection criterion for genebank accessions of L. perenne. Subsampling of locations was tested with allozyme analysis to determine whether genetically different populations could be collected at one location. Correlations between allelic frequencies and environmental factors and morphological data were established, to find ecogeographical patterns in the observed variation. Results indicated that with few examples each allelic variant could be observed in each population screened; very few unique alleles were found. Differences between populations were largely due to differences in allelic frequencies. Few correlations were found with environmental factors and morphological data. For some allele frequencies a north-south cline was observed. Generally, allozymic data of the five screened loci did not appear to be useful for the selection of accessions for genebank storage. Significant genotypic differences between populations collected at one location could be established. In general these results agreed with earlier results concerning phenotypic variation in the same populations.     

Outcrossing in celery (Apium graveolens)

Summary Isozyme marker loci were used to measure outcrossing in artificial and natural populations of celery (Apium graveolens). In three separate experiments involving populations of uniform density and flowering time, outcrossing rate ranged from 47 to 87% and the mean was 71.4%. In a wild population, outcrossing was estimated at 32.4 and 53.1%, using different homozygotes as the fixed female, and the weighted mean was 49.4%. A slight correlation of outcrossing rate with population density was observed. Mean outcrossing rates estimated from departures from panmictic expectations at polymorphic isozyme loci among open-pollinated plant introductions were 67.6% (celeries), 79.8% (celeriacs), and 85.1% (smallages). Based on these results, it was concluded that celery is best described as intermediate with respect to self- and cross-pollination.     

Genetic variability and heritability estimates for hardseededness and flowering in balansa clover (Trifolium michelianum Savi) populations

Suitability of annual pasture legume species like balansa clover (Trifolium michelianum Savi) in southern Australian farming systems depends on their hardseededness and time to flowering. Late maturing varieties with increased rate of hardseed breakdown, particularly in the late summer—early autumn period are desirable to ensure reliable regeneration in permanent pastures. Two half-sib family balansa clover populations were used to estimate heritability for the rate of breakdown of hardseededness and time to flowering. High narrow-sense heritability estimates (90.5–96.1%, rate of hardseed breakdown; 85.9–94.5%, time to flowering) were obtained, and were mainly attributed to additive gene effects. There was no relationship between rate of hardseed breakdown and time to flowering in Group1 half-sib family, which indicated that selection for both traits could be undertaken independently. In half-sib family Group 2 the relationship between the two traits suggested the possibility of selecting late maturing cultivars with increased rate of hardseed breakdown. Significant differences in time to flowering and rate of hardseed breakdown both within and between the two half-sib family groups suggested that phenotypic recurrent selection would be effective in improving the traits in these populations.