Fine mapping of a major quantitative trait locus that regulates pod shattering in soybean

Legumes represent the second most important family of crop plants, accounting for ~27 % of the world’s crop production. While some legumes are grown as forages or vegetables, most crop legumes are grown for harvesting their nutritious seeds. The legume seeds are contained in the pod, which is composed of a single seed-bearing carpel that, when matures, splits open along two seams, a process called pod dehiscence or pod shattering. Pod shattering before or during harvest causes yield losses of grain legumes. Moreover, the dominant shattering trait of the wild progenitors is a limiting factor for efficient introgression of value-added traits into elite breeding lines. Knowledge of the genetic mechanisms underlying pod shattering will facilitate breeding of shattering-resistant varieties, expedite introgression of agronomically favorable traits from wild species to elite breeding lines, and enrich our understanding of the evolution of seed dispersal and crop domestication in diverse crop species. Here we report fine mapping of a major quantitative trait locus (designated as qPDH1) that regulates pod shattering in soybean (Glycine max). A combination of linkage and association mapping allowed us to delimit the qPDH1 locus within a 47-kb region on chromosome 16. The data reported here will facilitate positional cloning of the underlying gene and the development of breeder-friendly genetic markers for marker-assisted selection in soybean.     

Mimicry of lentil and the domestication of common vetch and grass pea

A hypothesis is proposed whereby weedy vetch (Vicia sativa L.) seed moved with seed of the cultivated lentil (Lens culinaris Medikus) as a tolerated weed during the spread of the lentil from the Fertile Crescent in the Near East to its current distribution. As a result, selection occurred in vetch weeds for a reduction in dormancy/hard-seededness, increased competitive ability and biomass, and phenological adaptation to new environments⇆redisposing the weed for domestication. The cropping of common vetch for forage in pure culture followed. Archaeological evidence of admixtures of grass pea (Lathyrus sativus L.) in Neolithic finds of lentil, pea (Pisum sativum L.) and bitter vetch (Vicia ervilia (L.) Wild.) suggests a similar process of selection in grass pea for a weedy habit from which domestication later occurred.     

Identification of AFLP and STS markers closely linked to the <Emphasis Type="Italic">def</Emphasis> locus in pea

The recessive mutation of the def gene of pea (Pisum sativum L.) leads to the loss of the hilum, the abscission zone between the seed and the pod. Thereby, it reduces the free dispersal of the seeds through pod shattering. As a prerequisite for a gene isolation via a map-based cloning approach, bulked segregant analysis followed by single plant analyses of over 200 homozygous individuals of a population of 476 F2 plants derived from a cross between 'DGV' (def wild-type) and 'PF' (def mutant), were used to detect markers closely linked to the def locus. The AFLP technique in combination with silver staining was used to maximize numbers of reproducible marker loci. Fifteen AFLP loci showed a genetic distance less than 5 and two of them less than 1 centiMorgans (cM) to the gene of interest. AFLPs were converted into sequence tagged sites (STSs) and into a newly refined AFLP-based single locus marker named the 'sequence specified AFLP' (ssAFLP).     

Genetic Changes Accompanying the Domestication of Pisum sativum: Is there a Common Genetic Basis to the ‘Domestication Syndrome’ for Legumes?

BACKGROUND AND AIMS: The changes that occur during the domestication of crops such as maize and common bean appear to be controlled by relatively few genes. This study investigates the genetic basis of domestication in pea (Pisum sativum) and compares the genes involved with those determined to be important in common bean domestication. METHODS: Quantitative trait loci and classical genetic analysis are used to investigate and identify the genes modified at three stages of the domestication process. Five recombinant inbred populations involving crosses between different lines representing different stages are examined. KEY RESULTS: A minimum of 15 known genes, in addition to a relatively few major quantitative trait loci, are identified as being critical to the domestication process. These genes control traits such as pod dehiscence, seed dormancy, seed size and other seed quality characters, stem height, root mass, and harvest index. Several of the genes have pleiotropic effects that in species possessing a more rudimentary genetic characterization might have been interpreted as clusters of genes. Very little evidence for gene clustering was found in pea. When compared with common bean, pea has used a different set of genes to produce the same or similar phenotypic changes. CONCLUSIONS: Similar to results for common bean, relatively few genes appear to have been modified during the domestication of pea. However, the genes involved are different, and there does not appear to be a common genetic basis to 'domestication syndrome' in the Fabaceae.     

Construction of a SSR-based genetic map and identification of QTL for domestication traits using recombinant inbred lines from a cross between wild and cultivated cowpea (<Emphasis Type="Italic">V. unguiculata</Emphasis> (L.) Walp.)

Cowpea (Vigna unguiculata (L.) Walp.) is a grain legume commonly grown and consumed in many parts of the tropics and subtropics. A genetic linkage map was constructed using simple sequence repeat (SSR) markers and a recombinant inbred (RI) population of159 individuals derived from a cross between the breeding line 524B, a California Blackeye, and 219-01, a perennial wild cowpea from Kenya. Out of 912 primer combinations predicted to amplify SSRs in cowpea, 639 reliably produced amplification products in PCR assays and 202 (31.6%) were polymorphic between the two parents. These polymorphic SSRs were used to construct a genetic map consisting of 11 linkage groups (LGs) spanning 677 cM, with an average distance between markers of 3 cM. Agronomic traits related to domestication (seed weight, pod shattering) were analyzed together with the genotypic data. Six quantitative trait loci (QTL) for seed size were revealed with the phenotypic variation ranging from 8.9 to 19.1%. Four QTL for pod shattering were identified with the phenotypic variation ranging from 6.4 to 17.2%. The QTL for seed size and pod shattering mainly cluster in two areas of LGs 1 and 10, facilitating the use of marker-assisted selection to eliminate undesirable wild phenotypes in breeding activities involving introgression of traits from wild germplasm. The generation of an SSR-based molecular map and additional trait-linked markers also contributes to the expanding tool kit available to cowpea breeders, especially in Africa.     

Pod shattering in grain legumes: emerging genetic and environment-related patterns

A reduction in pod shattering is one of the main components of grain legume domestication. Despite this, many domesticated legumes suffer serious yield losses due to shattering, particularly under arid conditions. Mutations related to pod shattering modify the twisting force of pod walls or the structural strength of the dehiscence zone in pod sutures. At a molecular level, a growing body of evidence indicates that these changes are controlled by a relatively small number of key genes that have been selected in parallel across grain legume species, supporting partial molecular convergence. Legume homologs of Arabidopsis thaliana silique shattering genes play only minor roles in legume pod shattering. Most domesticated grain legume species contain multiple shattering-resistance genes, with mutants of each gene typically showing only partial shattering resistance. Hence, crosses between varieties with different genes lead to transgressive segregation of shattering alleles, producing plants with either enhanced shattering resistance or atavistic susceptibility to the trait. The frequency of these resistance pod-shattering alleles is often positively correlated with environmental aridity. The continued development of pod-shattering-related functional information will be vital for breeding crops that are suited to the increasingly arid conditions expected in the coming decades.

Recent genetic, genomic, and phenotypic studies of pod shattering in grain legumes lay the foundation for breeding crops suited for increasingly arid conditions.     

The chickpea, summer cropping, and a new model for pulse domestication in the ancient near east

The widely accepted models describing the emergence of domesticated grain crops from their wild type ancestors are mostly based upon selection (conscious or unconscious) of major features related either to seed dispersal (nonbrittle ear, indehiscent pod) or free germination (nondormant seeds, soft seed coat). Based on the breeding systems (self-pollination) and dominance relations between the allelomorphs of seed dispersal mode and seed dormancy, it was postulated that establishment of the domesticated forms and replacement of the wild ancestral populations occurred in the Near East within a relatively short time. Chickpea (Cicer arietinum L.), however, appears as an exception among all other "founder crops" of Old World agriculture because of its ancient conversion into a summer crop. The chickpea is also exceptional because its major domestication trait appears to be vernalization insensitivity rather than pod indehiscence or free germination. Moreover, the genetic basis of vernalization response in wild chickpea (Cicer reticulatum Ladiz.) is polygenic, suggesting that a long domestication process was imperative due to the elusive phenotype of vernalization nonresponsiveness. There is also a gap in chickpea remains in the archaeological record between the Late Prepottery Neolithic and the Early Bronze Age. Contrary to the common view that Levantine summer cropping was introduced relatively late (Early Bronze Age), we argue for an earlier (Neolithic) Levantine origin of summer cropping because chickpea, when grown as a common winter crop, was vulnerable to the devastating pathogen Didymella rabiei, the causal agent of Ascochyta blight. The ancient (Neolithic) conversion of chickpea into a summer crop required seasonal differentiation of agronomic operation from the early phases of the Neolithic revolution. This topic is difficult to deal with, as direct data on seasonality in prehistoric Old World field crop husbandry are practically nonexistent. Consequently, this issue was hardly dealt with in the literature. Information on the seasonality of ancient (Neolithic, Chalcolithic, and Early Bronze Age, calibrated 11,500 to 4,500 years before present) Near Eastern agriculture may improve our understanding of the proficiency of early farmers. This in turn may provide a better insight into Neolithic agrotechniques and scheduling. It is difficult to fully understand chickpea domestication without a Neolithic seasonal differentiation of agronomic practice because the rapid establishment of the successful Near Eastern crop package which included wheats, barley, pea, lentil, vetches, and flax, would have preempted the later domestication of this rare wild legume.     

Genomic dissection of pod shattering in common bean: mutations at non‐orthologous loci at the basis of convergent phenotypic evolution under domestication of leguminous species

The complete or partial loss of shattering ability occurred independently during the domestication of several crops. Therefore, the study of this trait can provide an understanding of the link between phenotypic and molecular convergent evolution. The genetic dissection of ‘pod shattering’ in Phaseolus vulgaris is achieved here using a population of introgression lines and next‐generation sequencing techniques. The ‘occurrence’ of the indehiscent phenotype (indehiscent versus dehiscent) depends on a major locus on chromosome 5. Furthermore, at least two additional genes are associated with the ‘level’ of shattering (number of shattering pods per plant: low versus high) and the ‘mode’ of shattering (non‐twisting versus twisting pods), with all of these loci contributing to the phenotype by epistatic interactions. Comparative mapping indicates that the major gene identified on common bean chromosome 5 corresponds to one of the four quantitative trait loci for pod shattering in Vigna unguiculata. None of the loci identified comprised genes that are homologs of the known shattering genes in Glycine max. Therefore, although convergent domestication can be determined by mutations at orthologous loci, this was only partially true for P. vulgaris and V. unguiculata, which are two phylogenetically closely related crop species, and this was not the case for the more distant P. vulgaris and G. max. Conversely, comparative mapping suggests that the convergent evolution of the indehiscent phenotype arose through mutations in different genes from the same underlying gene networks that are involved in secondary cell‐wall biosynthesis and lignin deposition patterning at the pod level.     

Identification of genome regions controlling cotyledon,pod wall/seed coat and pod wall resistance to pea weevil through QTL mapping

Pea weevil, Bruchus pisorum, is one of the limiting factors for field pea (Pisum sativum) cultivation in the world with pesticide application the only available method for its control. Resistance to pea weevil has been found in an accession of Pisum fulvum but transfer of this resistance to cultivated pea (P. sativum) is limited due to a lack of easy-to-use techniques for screening interspecific breeding populations. To address this problem, an interspecific population was created from a cross between cultivated field pea and P. fulvum (resistance source). Quantitative trait locus (QTL) mapping was performed to discover the regions associated with resistance to cotyledon, pod wall/seed coat and pod wall resistance. Three major QTLs, located on linkage groups LG2, LG4 and LG5 were found for cotyledon resistance explaining approximately 80 % of the phenotypic variation. Two major QTLs were found for pod wall/seed coat resistance on LG2 and LG5 explaining approximately 70 % of the phenotypic variation. Co-linearity of QTLs for cotyledon and pod wall/seed coat resistance suggested that the mechanism of resistance for these two traits might act through the same pathways. Only one QTL was found for pod wall resistance on LG7 explaining approximately 9 % of the phenotypic variation. This is the first report on the development of QTL markers to probe Pisum germplasm for pea weevil resistance genes. These flanking markers will be useful in accelerating the process of screening when breeding for pea weevil resistance.     

The domestication of American wildrice

The expansion in wildrice (Zizania palustris) production and the associated research efforts represent the largest modern effort to domesticate a cereal grain. Wildrice growers brought the species under cultivation, but plant breeding and agronomic research have accelerated the domestication process. The domestication and commercialization of this diploid (2n = 30), protogynous, cross-pollinated, annual, aquatic cereal present an opportunity to examine crop evolution and domestication theory. Traits associated with the domesticated cereal grains are shattering resistance, tiller synchrony, and increased seed size. This syndrome of traits may result from automatic selection, the selective force applied by repeated cycles of planting of harvested seed. Positive responses from deliberate selection for these traits in wildrice populations indicate that a domestication ideotype is attainable through plant breeding and that founder effect in this diploid species may be negligible. Continued commercial production of wildrice in the Great Lakes region is not likely to further the domestication process, whereas automatic selection may be initiated in the emerging California wildrice industry.     

Sequence polymorphisms in wild,weedy, and cultivated rice suggest seed‐shattering locus sh4 played a minor role in Asian rice domestication

The predominant view regarding Asian rice domestication is that the initial origin of nonshattering involved a single gene of large effect, specifically, the sh4 locus via the evolutionary replacement of a dominant allele for shattering with a recessive allele for reduced shattering. Data have accumulated to challenge this hypothesis. Specifically, a few studies have reported occasional seed‐shattering plants from populations of the wild progenitor of cultivated rice (Oryza rufipogon complex) being homozygous for the putative “nonshattering” sh4 alleles. We tested the sh4 hypothesis for the domestication of cultivated rice by obtaining genotypes and phenotypes for a diverse set of samples of wild, weedy, and cultivated rice accessions. The cultivars were fixed for the putative “nonshattering” allele and nonshattering phenotype, but wild rice accessions are highly polymorphic for the putative “nonshattering” allele (frequency ~26%) with shattering phenotype. All weedy rice accessions are the “nonshattering” genotype at the sh4 locus but with shattering phenotype. These data challenge the widely accepted hypothesis that a single nucleotide mutation (“G”/“T”) of the sh4 locus is the major driving force for rice domestication. Instead, we hypothesize that unidentified shattering loci are responsible for the initial domestication of cultivated rice through reduced seed shattering.     

亚洲栽培稻主要驯化性状研究进展

水稻是研究谷类作物驯化的良好材料, 其中种子落粒性消失、休眠性减弱和株型上的变化是水稻驯化过程中的3个关键事件, 造就了高产、发芽整齐及可密植的现代水稻。落粒性丧失一直被认为是野生稻驯化形态学上的最直接证据, 而控制落粒的主要基因Sh4和qSH1分别暗示不同的水稻驯化历史。种子休眠性的减弱适应了现代农业生产上同步发芽的需求, Sdr4、qSD7-1和qSD12基因是目前已知的调控种子休眠性的3个关键位点。野生稻匍匐生长等特点与其长期所在的易变生境有关, 而栽培稻的直立生长形态则适应了农业上密植生产的需要, 受PROG1等基因控制。野生稻的异交特性促进了驯化基因在群体间传播, 而自花授粉则使驯化基因得以稳定遗传, 从而加快人工选择的累积。目前的水稻驯化研究侧重于单基因或一些中性标记, 而对控制驯化性状的网络化通路的进化研究却相对缺乏。随着功能基因组研究的深入, 驯化性状的分子机理将会被全面揭示, 而基于此的网络化通路研究必将更加真实地反应水稻驯化过程。文章综述了水稻关键驯化性状分子机理的研究进展, 为驯化基因网络的研究提供参考, 也为水稻分子设计改良提供新的思路。     

油菜抗裂果性研究简述

本文简述了油菜易裂果性的危害、抗裂果性研究的意义及国内外研究现状和进展;也介绍了我国现有的主要抗裂果性油菜资源,并提出国内应开展油菜抗裂果性遗传机理研究和转基因研究的建议。     

Pulse domestication before cultivation

Comparative studies of the pulses of the Middle East and of their wild progenitors indicate that the pattern of pulse domestication is completely different from that of cereals in the same region. Wild legumes are not suitable for cultivation because of their conspicuous seed dormancy. Pre-adaptation of wild pulses for cultivation through loss of the seed dormancy mechanism apparently occurred in wild populations and may have been influenced by the gathering practices of man in pre-agricultural times. Pod indehiscence was of low value in pulse domestication and had evolved after the crop was well established and widespread.     

Pulse domestication and cereal domestication: How different are they?

Evidence is brought to indicate that the domestication of lentil and pea is not very different from that of wheat and barley. All these Near East crops are characterized by basically the same domestication traits the key elements of which are breakdown of the wild mode of seed dispersal and loss of germination regulation. It is argued that both in the pulses and in the cereals these traits evolved in the same way. The changes are best explained by assuming that mutations causing the loss of the wild-type adaptations were automatically selected for soon after people transferred the wild progenitors into a system of planting and reaping.     

‘兰箭3号’箭筈豌豆荚果发育动态及腹缝线结构研究

箭筈豌豆(Vicia sativa)是高海拔地区重要的一年生豆科牧草,但荚果成熟时的开裂现象会造成种子的严重损失。该研究以栽培品种‘兰箭3号’为对象,对其荚果在发育过程中的形态特征、水分含量、腹缝线表面结构及腹缝线横截面解剖结构的动态变化进行观察分析,以探讨箭筈豌豆荚果的裂荚机理,为生产中确定种子收获的适宜时间提供理论依据。结果显示:(1)‘兰箭3号’约在盛花后25~30d荚果变为浅棕色,此时荚果已完成生理成熟,且荚果的大小和干重均达到最大值,含水量降到最小值;盛花后25d荚果腹缝线出现裂缝,盛花后35d腹缝线完全裂开。(2)‘兰箭3号’于盛花后20d腹缝线处离层细胞开始解体;盛花后25d,内、中、外果皮的薄壁细胞均开始失水皱缩,其中内果皮的薄壁细胞部分已开始破裂,离层细胞及其下面的薄壁细胞完全解体,外部果瓣缘细胞内侧细胞壁破裂,但外侧异常加厚的细胞壁仍然保持完整并连接两个果瓣,使荚果不开裂;盛花后30~35d,内、中、外果皮的薄壁细胞完全失水,细胞壁皱缩在一起,同时外部果瓣缘细胞外侧细胞壁断裂成两部分,荚果的两个果瓣裂开。研究表明,盛花后25~30d荚果失去绿色变为浅棕色时是‘兰箭3号’的适宜收获时间,且离层和细胞失水产生的机械拉力是导致箭筈豌豆荚果开裂的主要原因,推测外部果瓣缘细胞外侧增厚融合的细胞壁很可能是‘兰箭3号’抵抗裂荚的关键结构。     

Formation Mechanism and Occurrence Law of Pod Shattering in Soybean: A Review

Seed shattering refers to the phenomenon in which the pods split along the abdominal and back sutures before the crop is received, so that the seeds are spread. Seed shattering is vital to the reproduction of their offspring in wild plants, but it is also the main cause of crop yield loss reason. Pod-explosion resistance is a complex process of physical and physiological and biochemical reactions. Soybean seed shattering phenomenon is widespread, which severely restricts the development of soybean industry. Seed shattering (pod cracking or fruit dropping) is essential for the reproduction of its offspring in wild plants, but it is also the main cause of crop yield loss. This article analyzes the morphology and structure of pods related to seed shattering from the morphology of pods. On the basis of the regularity of the occurrence of seed shattering and the summary of phenotypic index identification methods, physiologically introduced the regulation mechanism of key enzymes and endogenous hormones on seed shattering. The localization, labeling and cloning of seed shattering genes are introduced in molecular biology. The study focused on reviewing the latest advances in the research on soybean seed shattering characteristics, and discussed with the research results of related crops. Finally, the research and application of soybean seed shattering resistance were prospected for several aspects.