Unmanaged sexual reproduction and the dynamics of genetic diversity of a vegetatively propagated crop plant, cassava (Manihot esculenta Crantz), in a traditional farming system

Occurrence of intervarietal or interspecific natural crosses has been reported for many crop plants in traditional farming systems, underlining the potential importance of this source of genetic exchange for the dynamics of genetic diversity of crop plants. In this study, we use microsatellite loci to investigate the role of volunteer seedlings (plants originating from unmanaged sexual reproduction) in the dynamics of genetic diversity of cassava (Manihot esculenta Crantz), a vegetatively propagated crop, in a traditional farming system in Guyana. A previous field study showed that farmers incorporate such plants into the germplasm for vegetative propagation, and that many of them are likely to be assigned by farmers to recognized varieties. Under strict vegetative propagation clonality of varieties is expected. The high proportion of polyclonal varieties observed suggests that incorporation of seedlings into the germplasm for propagation is a frequent event. The molecular variability assessed with microsatellite markers shows that there is high differentiation among heterozygous varieties, whereas populations of seedlings do not depart from the proportions expected under Hardy-Weinberg assumptions. Assignment of seedlings to a recognized variety on the basis of morphological similarity greatly increases genetic diversity within the variety. We argue that recombination and gene flow play a major role in the dynamics of genetic diversity of cassava in traditional farming systems. Documenting unmanaged sexual reproduction and its genetic consequences is a prerequisite for defining strategies of in situ conservation of crop plant genetic resources.     

Dynamic Management of Maize Landraces in Central Mexico

Conservationists of crop genetic resources have feared that in situ conservation was not viable for agriculture precisely because of changes resulting from introduction of new varieties of existing crops, new crops, and new farm practices. In addition, conservation within farming systems necessarily implies a constantly changing crop population resulting from the processes of crop evolution. Even though in situ conservation of crop genetic resources is now generally understood to be dynamic, there are few examples of how evolution takes place in farmers fields. This study describes several changes in maize landraces in four communities along an altitude transect in Central Mexico (1200 to 2400 masl). While true modern varieties have not been widely adopted in the study region, farmer management results in numerous changes in maize landrace populations. Five types of dynamic management were observed: (1) purposeful hybridization between traditional and modern maize types, (2) possible creation of a new maize landrace by directional selection of the progeny of hybridization between two traditional landraces, (3) displacement of a local landrace by the introduction of a modern variety and a non-local landrace, (4) maintenance of stable populations of a locally dominant landrace, and (5) market-driven selection for a minor variety. We concur that in situ conservation of crops must be conceived as an open process where the objective is not to maintain historic varieties or static genetic conditions. Rather, in situ conservation of crops is totally in the hands of the farmer, although interventions may be designed to influence farmers’ management of agrobiodiversity.     

Genetic diversity of traditional South American Landraces of Cassava (Manihot esculenta Crantz): an analysis using microsatellites

The extent and structure of the genetic variability of traditional varieties of cassava (Manihot esculenta Crantz) have been little documented, despite considerable evidence for this crop? great varietal diversity in traditional agroecosystems. We used microsatellite markers to assess the genetic structure of traditional landraces of sweet and bitter cassava collected from five South American sites. As reference, we used a sample of 38 accessions from a world collection of cultivated cassava. For a total of 10 loci examined, we found 15 alleles that were not represented in this sample. Ten of these had been previously detected in wild Manihot species. The geographical structure of genetic variability was weak, but the genetic differentiation between bitter and sweet landraces was significant, suggesting that each form had evolved separately after domestication. Our results showed that traditional landraces form an important source of genetic diversity and merit more attention from managers of crop genetic resources.     

Introgression from modern hybrid varieties into landrace populations of maize (Zea mays ssp. mays L.) in central Italy

Landraces are domesticated local plant varieties that did not experience a deliberate and intensive selection during a formal breeding programme. In Europe, maize landraces are still cultivated, particularly in marginal areas where traditional farming is often practiced. Here, we have studied the evolution of flint maize landraces from central Italy over 50 years of on-farm cultivation, when dent hybrid varieties were introduced and their use was widespread. We have compared an 'old' collection, obtained during the 1950s, before the introduction of hybrids, and a recent collection of maize landraces. For comparison, a sample of maize landraces from north Italy, and of improved germplasm, including hybrids and inbred lines were also used. A total of 296 genotypes were analysed using 21 microsatellites. Our results show that the maize landraces collected in the last 5–10 years have evolved directly from the flint landrace gene pool cultivated in central Italy before the introduction of modern hybrids. The population structure, diversity and linkage disequilibrium analyses indicate a significant amount of introgression from hybrid varieties into the recent landrace populations. No evidence of genetic erosion of the maize landraces was seen, suggesting that in situ conservation of landraces is an efficient strategy for preserving genetic diversity. Finally, the level of introgression detected was very variable among recent landraces, with most of them showing a low level of introgression; this suggests that coexistence between different types of agriculture is possible, with the adoption of correct practices that are aimed at avoiding introgression from undesired genetic sources.     

Insights on the evolution of a vegetatively propagated crop species

The opportunity for gene flow between a vegetatively propagated crop and its wild relatives is expected to be much lower than for seed-propagated crops, since sexual reproduction in the crop occurs only infrequently. A study by Duputié and colleagues now demonstrates evidence of sexual reproduction between a vegetatively propagated crop and a closely related wild congener. Working in French Guiana, these workers have documented a hybrid zone arising from introgression between cassava (Manihot esculenta ssp. esculenta, Euphorbiaceae), which is propagated by stem cuttings, and wild Manihot populations growing in close proximity. Patterns of heterozygosity suggest that there are little-to-no barriers to reproduction between the crop and these wild populations. Previous work by these researchers has documented the importance of occasional sexual reproduction for the development of cassava varieties in traditional Amerindian farming systems. Taken together with their previous work, these new findings suggest that gene flow between wild Manihot populations and cassava plants could potentially play a much greater role in the crop's evolution than previously thought.     

Traditional Maize Storage Methods of Mayan Farmers in Yucatan, Mexico: Implications for Seed Selection and Crop Diversity

For small-scale farmers who maintain genetically diverse crop populations, aspects of the storage of their seeds and harvest may be just as important for successful farming as those related to productivity. The community of Yaxcaba, Yucatan, Mexico was studied to understand how the conditions under which Maya farmers store their maize harvests influence their seed selection practices and the diversity of maize varieties grown. Most farmers select their maize seed based primarily on ear characteristics and secondarily on grain characters. Farmers incorporate storability concerns by selecting for an archetypal healthy ear and by conducting selection in several steps between harvest and planting of the subsequent crop generation. Most farmers store their maize harvest in the husk, initially in the field and then in rustic granaries constructed of logs and palm thatch, in which farmers conserve separately their different seed lots. All local landrace populations show morphological adaptation (principally husk characteristics) for local storage conditions, indicating that storability has been an important selection pressure on traditional maize. Storability also appears to be a key factor working against the straightforward adoption of improved maize seed in Yaxcaba. Local farmers value many qualities of improved maize varieties, but their principal complaint is that improved maize cannot be conserved reliably under local storage conditions. Nearly all farmers who utilize improved seed stock in Yaxcaba grow locally adapted or ‘creolized’ versions of improved varieties, displaying characteristics of local maize landraces that facilitate their storage.     

Landraces of maize in Central Mexico: an altitudinal transect

Conservation of crop genetic resources is now considered an important component of sustainable agricultural development. If conservation of genetic resources for agriculture is to be successful, a more complete understanding of the dynamics affecting traditional (landrace) crop populations is needed. We conducted a study of maize-based agriculture in the Central Highlands of Mexico in communities at 2400, 1700, 1400, and 1200 masl to assess the status of traditional varieties in an area characterized by thorough integration into the national economy. Our research contradicts the view that modern varieties persist because of marginal conditions, deficient infrastructure, weaker markets, or traditional attitudes. One or two landraces dominated highland maize populations and farmers appeared to be more conservative in terms of their emphasis on traditional maize varieties than at lower elevations. The dominance of traditional varieties in the highlands is well known but poorly explained, and the coexistence of traditional and modern varieties in the mid-elevations was unexpected. Our highland study area has good roads, is near Mexico City, and is less than 50 km away from four major crop research institutes that have done maize breeding since 1950’s. We suggest that in situ conservation of maize genetic resources in the highlands is sustained because the landraces there have good agronomic performance and are highly valued by farmers for their end-use qualities. At the mid-elevations, competition between local and modern maize was sharpest, and farmers have found that both landraces and improved varieties suit their needs, hence enhancing genetic diversity. Interventions and incentives would appropriately be carried out here to assure in situ conservation of locally adapted landraces of maize.     

Population Structure of Barley Landrace Populations and Gene-Flow with Modern Varieties

Landraces are heterogeneous plant varieties that are reproduced by farmers as populations that are subject to both artificial and natural selection. Landraces are distinguished by farmers due to their specific traits, and different farmers often grow different populations of the same landrace. We used simple sequence repeats (SSRs) to analyse 12 barley landrace populations from Sardinia from two collections spanning 10 years. We analysed the population structure, and compared the population diversity of the landraces that were collected at field level (population). We used a representative pool of barley varieties for diversity comparisons and to analyse the effects of gene flow from modern varieties. We found that the Sardinian landraces are a distinct gene pool from those of both two-row and six-row barley varieties. There is also a low, but significant, mean level and population-dependent level of introgression from the modern varieties into the Sardinian landraces. Moreover, we show that the Sardinian landraces have the same level of gene diversity as the representative sample of modern commercial varieties grown in Italy in the last decades, even within population level. Thus, these populations represent crucial sources of germplasm that will be useful for crop improvement and for population genomics studies and association mapping, to identify genes, loci and genome regions responsible for adaptive variations. Our data also suggest that landraces are a source of valuable germplasm for sustainable agriculture in the context of future climate change, and that in-situ conservation strategies based on farmer use can preserve the genetic identity of landraces while allowing adaptation to local environments.     

Development of simple sequence repeat (SSR) markers for the assessment of gene flow and genetic diversity in pigeonpea (Cajanus cajan)

Pigeonpea (Cajanus cajan) is an important subsistence crop in India where traditional landraces and improved hybrids are grown alongside each other. Gene flow may result in genetic erosion of these landraces and their wild relatives, whilst transgene escape from future genetically engineered varieties is another potential hazard. To assess the impact of these factors gene flow needs to be measured. A set of 10 simple sequence repeat markers have been developed, which exhibit polymorphism across a range of pigeonpea varieties. Use of these markers also offers an efficient system for the assessment of genetic diversity within populations of pigeonpea.     

Genetic diversity and population structure of common bean (<Emphasis Type="Italic">Phaseolus vulgaris</Emphasis> L<Emphasis Type="Italic">.</Emphasis>) landraces from the East African highlands

The East African highlands are a region of important common bean production and high varietal diversity for the crop. The objective of this study was to uncover the diversity and population structure of 192 landraces from Ethiopia and Kenya together with four genepool control genotypes using morphological phenotyping and microsatellite marker genotyping. The germplasm represented different common bean production ecologies and seed types common in these countries. The landraces showed considerable diversity that corresponded well to the two recognized genepools (Andean and Mesoamerican) with little introgression between these groups. Mesoamerican genotypes were predominant in Ethiopia while Andean genotypes were predominant in Kenya. Within each country, landraces from different collection sites were clustered together indicating potential gene flow between regions within Kenya or within Ethiopia. Across countries, landraces from the same country of origin tended to cluster together indicating distinct germplasm at the national level and limited gene flow between the two countries highlighting divided social networks within the regions and a weak trans-national bean seed exchange especially for landrace varieties. One exception to this may be the case of small red-seeded beans where informal cross-border grain trade occurs. We also observed that genetic divergence was slightly higher for the Ethiopian landraces compared to Kenyan landraces and that Mesoamerican genotypes were more diverse than the Andean genotypes. Common beans in eastern Africa are often cultivated in marginal, risk-prone farming systems and the observed landrace diversity should provide valuable alleles for adaptation to stressful environments in future breeding programs in the region.     

Diversity and structure of landraces of Agave grown for spirits under traditional agriculture: A comparison with wild populations of A. angustifolia (Agavaceae) and commercial plantations of A. tequilana

Traditional farming communities frequently maintain high levels of agrobiodiversity, so understanding their agricultural practices is a priority for biodiversity conservation. The cultural origin of agave spirits (mezcals) from west-central Mexico is in the southern part of the state of Jalisco where traditional farmers cultivate more than 20 landraces of Agave angustifolia Haw. in agroecosystems that include in situ management of wild populations. These systems, rooted in a 9000-year-old tradition of using agaves as food in Mesoamerica, are endangered by the expansion of commercial monoculture plantations of the blue agave variety (A. tequilana Weber var. Azul), the only agave certified for sale as tequila, the best-known mezcal. Using intersimple sequence repeats and Bayesian estimators of diversity and structure, we found that A. angustifolia traditional landraces had a genetic diversity (H(BT) = 0.442) similar to its wild populations (H(BT) = 0.428) and a higher genetic structure ((B) = 0.405; (B) =0. 212). In contrast, the genetic diversity in the blue agave commercial system (H(B) = 0.118) was 73% lower. Changes to agave spirits certification laws to allow the conservation of current genetic, ecological and cultural diversity can play a key role in the preservation of the traditional agroecosystems.     

Wheat gene bank accessions as a source of new alleles of the powdery mildew resistance gene <Emphasis Type="Italic">Pm3</Emphasis>: a large scale allele mining project

Background  

In the last hundred years, the development of improved wheat cultivars has led to the replacement of landraces and traditional varieties by modern cultivars. This has resulted in a decline in the genetic diversity of agriculturally used wheat. However, the diversity lost in the elite material is somewhat preserved in crop gene banks. Therefore, the gene bank accessions provide the basis for genetic improvement of crops for specific traits and and represent rich sources of novel allelic variation.     

Estimating maize genetic erosion in modernized smallholder agriculture

Replacement of crop landraces by modern varieties is thought to cause diversity loss. We studied genetic erosion in maize within a model system; modernized smallholder agriculture in southern Mexico. The local seed supply was described through interviews and in situ seed collection. In spite of the dominance of commercial seed, the informal seed system was found to persist. True landraces were rare and most informal seed was derived from modern varieties (creolized). Seed lots were characterized for agronomical traits and molecular markers. We avoided the problem of non-consistent nomenclature by taking individual seed lots as the basis for diversity inference. We defined diversity as the weighted average distance between seed lots. Diversity was calculated for subsets of the seed supply to assess the impact of replacing traditional landraces with any of these subsets. Results were different for molecular markers, ear- and vegetative/flowering traits. Nonetheless, creolized varieties showed low diversity for all traits. These varieties were distinct from traditional landraces and little differentiated from their ancestral stocks. Although adoption of creolized maize into the informal seed system has lowered diversity as compared to traditional landraces, genetic erosion was moderated by the distinct features offered by modern varieties.     

Maize genetic diversity in traditionally cultivated polycultures in an isolated rural community in Mexico: implications for management and sustainability

ABSTRACT

Background: Maize in Mexico exhibits great genetic diversity, maintained by traditional practices of indigenous and non-indigenous communities, the same practices that have led to crop diversification over centuries. As one of the main staple crops worldwide, safeguarding the genetic diversity of maize is paramount to food security.

Aims: This study evaluated the genetic diversity and population structure of traditionally cultured maize landraces in a rural seasonal agricultural community in Veracruz, Mexico, in order to learn how traditional practices shape these landraces, and propose strategies for their preservation.

Methods: We analysed 118 individual maize samples belonging to five morphotypes (white, yellow, black, red and mottled) with eight microsatellite markers.

Results: We encountered high genetic diversity, according to expected heterozygosity (H_e = 0.61). However, inbreeding coefficient and gene flow values suggested the existence of assortative mating, which causes low genetic differentiation. Population structure analysis identified three genetic pools, independent of grain colour. These findings suggest that all morphotypes belong to the same population, which is sub-structured due to assortative mating and gene flow related to local agronomic management.

Conclusions: Current management practices in this community could lead to genetic erosion. In order to preserve diversity, wider regional seed exchange and selection for morphological diversity could be implemented.     

中国农作物及其野生近缘植物多样性研究进展

本研究围绕粮食、经济、果树、蔬菜、饲草与绿肥、花卉六大类作物,以物种多样性和遗传多样性等为主线,开展了中国农作物及其野生近缘植物多样性研究,取得重大进展。科学界定和系统规范了种质资源概念和范畴,提出了层次结构的理论,查清了中国粮食与农业植物有9631个物种及其分布、特征特性及用途;深入研究了中国农作物种质资源的本底,首次明确了中国农作物总计有528种（类）,涉及1339个栽培物种和1930个野生近缘植物物种;优化了农作物种以下农艺性状分类技术指标,阐明了中国农作物地方品种的987个变种、978个变型、1223个农艺性状特异类型的植物学特征、生物学特性及其遗传多样性;得出了中国是禾谷类作物裸粒基因、糯性基因、矮秆基因和育性基因等特异基因的起源中心或重要起源地之一的结论;首次编撰出版了《中国作物及其野生近缘植物》专著6卷,并编撰了《中国粮食与农业植物物种名录》,形成了具有中国特色的作物种质资源学科理论体系。为农作物种质资源的高效利用、保障粮食安全和农业可持续发展奠定了基础。     

Farmers Drive Genetic Diversity of Thai Purple Rice (Oryza sativa L.) Landraces

Purple or black rice (Oryza sativa L.) is a culturally important germplasm in Asia with a long history of cultivation in northern Thailand. Purple rice is identified by the color of the rice pericarp, which varies from purple to black with the accumulation of phenolic acids, flavonoids, and anthocyanins. In the present study, we assessed molecular variation within and between wetland purple rice landraces germplasm from northern and northeastern Thailand using 12 microsatellite loci. All purple rice varieties surveyed showed high levels of homozygosity within varieties and strong genetic differentiation among varieties, indicating the fixation of genetic differences among them. This pattern is consistent with purple rice farming practices in northern Thailand, where a small portion of harvested seed is selected and replanted based on farmers’ preferences. The reduced genetic diversity and high homozygosity observed for purple rice is also consistent with patterns expected for this inbreeding crop. Genetic differentiation among the varieties showed some degree of structuring based on their geographical origin. Taken together, these data highlight that the genetic diversity and structure of wetland purple rice landraces is shaped by farmer utilization and cultivation through local cultural practices, and that conservation should focus on ex situ conservation across its cultivation range, along with on-farm, in situ conservation based on farmers’ seed-saving practices. In situ conservation may prove especially valuable for preserving the genetic identity of local varieties and promote adaptation to local environments.

     

Thai elite cassava genetic diversity was fortuitously conserved through farming with different sets of varieties

Many in situ conservation programs have been developed to preserve plant landrace diversity and to promote its sustainable utilization, but little is known about the effectiveness of the developed programs in conserving plant genetic diversity. We investigated the effectiveness of an unregulated (i.e., unplanned or open) conservation system maintained by Thai farmers in conserving Thai elite cassava (Manihot esculenta Crantz) varieties. Specifically, we compared genetic diversity of 266 cassava clones that were collected from 80 farms in eight provinces with 16 cassava landraces and varieties released since the 1970s through genotyping with 35 informative simple sequence repeat (SSR) markers. The SSR analysis revealed a large regional heterogeneity in cassava diversity, with a strong genetic differentiation of the assayed clones among the 80 farms (19.8 %) and across the eight provinces (11.8 %). Significant associations were also found between SSR variation and farm agro-ecological factors or some farming practices. However, there was no significant genetic differentiation (0.9 %) between the 266 farm clones and 16 reference varieties. These findings suggest that the Thai elite cassava genetic diversity was fortuitously conserved by the farmers through farming with different sets of varieties. Implications of these findings are discussed with respect to on-farm conservation of plant genetic resources.     

Challenges for transgene detection in landraces and wild relatives: learning from 15 years of debate over GM maize in Mexico

Maize is one of the world’s five staple cereals and its traditional varieties constitute a global resource critical to future agricultural development. Fifteen years ago, claims that transgenes had spread into traditional landrace maize in Mexico started an international discussion on the scale and significance of transgene flow from genetically modified (GM) crops to centres of crop origin and genetic diversity. The initial discovery of transgenes in landrace maize sparked an intense environmental dispute in which the culture and traditions of indigenous people were seen as threatened by the unchecked spread of biotechnological inventions from multinational corporations. This dispute was reflected in a political and legal battle over the regulatory status of GM crops in Mexico, which continues today as approvals of GM maize for cultivation remain subject to contestation in the courts. These legal, political and environmental disputes have been fanned by the existence of a significant scientific controversy over the methods for GM detection. The use of various approaches and a lack of harmonized methods specific for monitoring and detection of transgenes in landraces has generated both positive and negative results for GM contamination in Mexico over the years. In this paper, we review the peer-reviewed literature on transgene detection in Mexican maize and highlight the challenges associated with transgene detection in landraces. In doing so, we identify the key methodological aspects under dispute and pinpoint the research bottlenecks and needs for building the capacity to effectively monitor transgene escape from GM crops to wild relatives or landraces.     

新疆若羌县与且末县抗逆农作物种质资源调查

若羌县、且末县地处塔克拉玛干沙漠南缘,是新疆干旱半干旱生态区的典型代表,境内生态类型多样,气候干旱少雨、土壤次生盐渍化重,至今仍存在有一定规模的传统农业耕作模式,保留着丰富多样的抗逆农作物种质资源。本研究在农作物种质资源普查基础上,分析了两县的社会经济发展、农业生产及种植结构调整的演变规律;调查收集抗逆种质资源121份,隶属11科28属34种,分析评价了部分作物种质资源的抗逆性,以及与当地的生态条件、农业生产模式和百姓生活习惯的内在联系,探讨了主要农作物品种演替的阶段性变化,讨论了新疆干旱区抗逆农作物种质资源的收集、保护和利用问题。     

重庆部分地区的种质资源调查荞麦篇

"第三次全国农作物种质资源调查与收集"重庆项目组,于2015年10月至11月初在重庆市巫山县、奉节县和石柱县3个县开展了农业生物资源的系统调查,通过深入村社和农户,调查了解作物种类与品种多样性情况,收集和挖掘古老、特有和稀少种质资源。本次调查共收集到农作物地方品种和野生近缘植物种质资源341份,其中粮油作物136份、蔬菜86份、果树57份、茶树5份、烟草22份、牧草30份、麻类5份。本文重点介绍在粮油作物中的荞麦种质资源调查和资源收集情况,在收集到的19份荞麦资源中,既有珍稀、古老的地方品种,也包括特有野生资源。荞麦承载了彝族等少数民族的文化内涵,随着现代农业的发展,荞麦的古老、珍稀和特有种质资源正在流失,建议相关部门因地制宜,在发展现代农业生产的同时,采取必要的经济措施,对古老、珍稀和特有的荞麦种质资源,进行农户或原生境保护,为进一步开发利用提供种质资源基础。