Genome-wide association study to identify chromosomal regions related to panicle architecture in rice (Oryza sativa)

Panicle architecture directly affects the grain yield of rice; meanwhile, crop domestication has increased the complexity of rice panicle size and branching pattern. Genome-wide association study (GWAS) was performed to investigate the genetic basis underlying panicle architecture using 183 rice accessions from around the world and a 7 K SNP array. Phenotyping was conducted at Texas A&M AgriLife Research Center in Beaumont, TX. GWAS was performed using MLM (Q?+?K) model using GAPIT software. At p-value?<?0.001, a total of 49 GWAS QTLs controlling various panicle architecture traits were mapped. Considering the recurring and linked SNPs across the panicle architecture traits, 42 independent QTL regions were identified. Among these, 27 QTL regions co-localized with known genes or previously reported QTLs or significant SNP markers, while 15 were potentially novel QTL regions. The results of our study offer useful information on genetic bases controlling panicle architecture in rice, which could be further validated and utilized for designing markers for use in markers assisted selection (MAS) in rice breeding programs.

     

Origin of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) domestication genes

A number of genes that contribute to the domestication traits of cultivated rice have been identified. These include Sh4, Rc, PROG1 and LABA1, which are associated with non-shattering rachis, white pericarp, erect growth and barbless awns, respectively. The mutations giving rise to the “domestication alleles” of these genes are either invariable in cultivated rice, or have variability that is strictly associated with the phenotypic trait. This observation forms the basis to those current rice domestication models that envisage a single origin for the domesticated phenotype. Such models assume that the domestication alleles are absent or rare in wild rice, emerged under cultivation and spread across all rice groups by introgressive hybridization. We examined whole-genome sequencing datasets for wild and cultivated rice to test the former two assumptions. We found that the rc and laba1 alleles occur in wild rice with broad geographical distribution, and reach frequencies as high as 13 and 15%, respectively. These results are in agreement with previous observations of the prog1 and sh4 domestication alleles in wild populations. We also show that the diversity of the genomic regions surrounding the rc, laba1, prog1 and sh4 alleles in wild accessions is greater than that in cultivated rice, suggesting that these alleles emerged prior to domestication. Our findings indicate that the possibility that independent rice groups obtained identical domestication alleles directly from the wild population needs to be considered.     

Agronomic and genetic characterization of wild emmer wheat (<Emphasis Type="Italic">Triticum turgidum</Emphasis> subsp<Emphasis Type="Italic">. dicoccoides</Emphasis>) introgression lines in a bread wheat genetic background

During a long evolutionary history across a range of environmental conditions in the Near East Fertile Crescent, wild emmer wheat (Triticum turgidum subsp. dicoccoides) has accumulated a wealth of genetic diversity and adaptations to multiple biotic and abiotic stress conditions. The joint effects of the domestication and subsequent selection in man-made agro-ecosystems have considerably reduced the genetic variation in cultivated crop species as compared to their wild progenitors. The wild emmer gene pool harbors a rich allelic repertoire for improving numerous agronomically important traits. A set of advanced wild emmer wheat introgression lines (ILs) was previously established on a bread wheat genetic background. It combines genetic diversity from the wild progenitor as well as the genetic background of the cultigen. Based on field evaluations, the set of ILs (divided hierarchically into different families based on the genetic background) has exhibited high phenotypic variance in agronomically important traits. Analyses of variance ?(ANOVA) have shown that the effect of the hierarchical genetic factors was significant in the vast majority of the traits. Similarly, characterizations by molecular markers of known traits have shown that the genetic profile differs between ILs from different genetic backgrounds. Multivariate principal component analysis has revealed interesting associations between vegetative and reproductive traits and illustrated the impact of some of these traits on ILs distribution in the PC1 and PC2. The exploitation of the wild emmer wheat ILs as a tool for reintroduction of alleles that were lost during domestication and possible implications for wheat breeding are discussed.     

The domestication and dispersal of the cultivated ramie (Boehmeria nivea (L.) Gaud. in Freyc.) determined by nuclear SSR marker analysis

Ramie (Boehmeria nivea) was domesticated in China. However, the geographic region of domestication is not exactly known. Genetic diversity and population structure of ramie and their wild relatives were assessed using microsatellite markers. The 8 microsatellite primers revealed 96 alleles in 50 ramie populations, with an average of 10.25 alleles per locus. Cultivated ramie gene pool harbors approximately 82.9 % of the SSR diversity presented in wild B. nivea var. nivea. It is suggested that ramie has experienced a relatively moderate domestication bottleneck. The distribution of genetic diversity shows that genetic diversity is relatively high in populations along the Yangtze River compared to the peripheral ones. The scatter plots of principal coordinates analysis indicated that there are three well-supported varieties in B. nivea. The NJ tree and the distribution of genetic diversity showed that ramie has been domesticated in the middle and lower regions of the Yangtze valley, and populations in Sichuan province have been introduced from this region, forming naturalized populations.     

Quantitative Trait Loci for Seed Dormancy in Wild Barley (Hordeum spontaneum C. Koch)

A quantitative trait locus analysis was carried out to unravel the genetic basis of dormancy in wild barley (Hordeum spontaneum) from Israel. Two accessions, Ashkelon and Mehola, from divergent environments were crossed to produce a mapping population. A linkage map was produced from the F₂ population, and F₄ seeds were used for germination experiments. Five quantitative trait loci (QTL) were detected for dormancy across the different germination experiments. These QTL were found on chromosomes 1, 2, 5, 6 and 7. The variation explained by each QTL varied between 8 and 25%. Ashkelon alleles increased the germination except for the QTL on chromosome 5. Three out of these five QTL co-locate with QTL found earlier in cultivated barley crosses, although this does not necessarily imply that they would be the same loci. The level of dormancy is much higher in wild barley than in cultivated barley and wild barley may have alleles that have not yet been utilised in breeding for optimally dormant barley.     

Whole-genome analysis with SNPs from BOPA1 shows clearly defined groupings of Western Mediterranean, Ethiopian, and Fertile Crescent barleys

The discovery of Hordeum spontaneum C. Koch, a wild ancestor of cultivated barley, in Morocco in 1978 led to the proposal of a multicentric origin for this crop, as an alternative to the widely accepted theory of a single centre of domestication in the Fertile Crescent. Since this discovery, we have tested this hypothesis using the most advanced genetic techniques available at the time, from CM-proteins to RFLP and DNA-chloroplast markers. Nowadays, the availability of single nucleotide polymorphism (SNP) markers that are spread densely over the barley genome provides us with another powerful tool to give further support for the above. We have used 1,536 SNPs from the Barley Oligo Pool Assay 1 (BOPA1) of Illumina to characterize 107 wild and cultivated barley accessions from the Western Mediterranean, Fertile Crescent, Ethiopia, and Tibet. The results have confirmed that each location of the above-mentioned germplasm groups clusters separately. Analysis of molecular variance enabled us to focus on the chromosomal regions and loci that differentiated these groups of barley germplasm. Some of these regions contain vernalization and photoperiod response genes, some of the so-called domestication genes, as well as the most important quantitative trait locus for flowering time in the Mediterranean region. We have combined these results with other genetic evidence, and interpreted them in the framework of current theories on the onset of the Neolithic revolution in the Mediterranean region, to conclude that neither Ethiopia nor the Western Mediterranean can be ruled out as centres of barley domestication, together with the Fertile Crescent.     

Identification of artichoke SSR markers: molecular variation and patterns of diversity in genetically cohesive taxa and wild allies

A set of 24 microsatellite markers was identified in the artichoke genome, using various approaches. A genomic library allowed the development of 14 SSR markers, whereas the other 10 were obtained from gene intron/UTR regions or from other species. Allelic variation was scored in C. cardunculus (artichoke, cultivated cardoon, and wild cardoon) samples, and in other wild Cynara allies. For the 23 polymorphic loci, a total of 165 alleles were scored, 135 of which in the artichoke primary genepool, and the remaining ones in the other Cynara species. Some allele combinations were able to identify artichoke varietal types, and some alleles were unique to specific groups. This makes these markers potentially useful in product traceability and in contributing to the saturation of genetic maps. The percentage of shared alleles between C. cardunculus taxonomic groups, and Nei’s genetic distances indicated that wild cardoons from the Eastern Mediterranean were more closely related to artichoke and less to cultivated cardoon in comparison to wild cardoons from the Western Mediterranean, and the genetic distance between the two wild cardoon genepools was rather high. The UPGMA dendrogram based on Nei’s genetic distances revealed that artichokes formed a fairly defined cluster, whereas Eastern wild cardoons occupied another branch, and Western wild cardoons were clustered together with cultivated cardoons. The transferability of microsatellite markers to other Cynara wild species was quite good. Sequencing alleles at three loci showed that, apart from microsatellite length variation, point mutations and insertion/deletions were quite abundant especially when comparing C. cardunculus to the other Cynara species. In the sequenced regions, some SNPs were identified which distinguished artichoke on one side, and cultivated and wild cardoon on the other, while other SNPs were apportioned according to the geographic distribution of Cynara wild species.     

How many tough-rachis mutants gave rise to domesticated barley?

It is commonly agreed that cultivation of wild barley preceded the selection of the domesticated, non-brittle spike type. However, how common was wild barley cultivation before domestication and how many domesticated mutants gave rise to the barley crop could not be inferred from botanical and archaeological evidence. Some clues, nevertheless, can be obtained from the pattern of allozyme diversity in wild and cultivated barley obtained by Kahler and Allard (1981). Parallel variation, in terms of number of alleles per locus and frequency of the various alleles, was found in wild and domesticated barley. This similarity has been taken as an indication of multiple domestications and the frequency of the rarest alleles has been used to estimate that about 100 tough-rachis different mutants were necessary for the inclusion of the allozyme diversity of the wild barley in the domesticated crop. Assuming mutation rate of 10^–6 in the locus governing tough rachis, the plant population required to generate these 100 mutants in one year would extend over about 200 hectares, or 10 hectares if the 100 mutants have been formed over a period of 20 years. The simplified calculations suggest that prior to domestication cultivation of wild barley was not a common practice.     

Development and use of microsatellite markers for genetic diversity analysis of cañahua (<Emphasis Type="Italic">Chenopodium pallidicaule</Emphasis> Aellen)

Cañahua (Chenopodium pallidicaule Aellen) is a poorly studied, annual subsistence crop of the high Andes of South America. Its nutritional value (high in protein and mineral content) and ability to thrive in harsh climates make it an important regional food crop throughout the Andean region. The objectives of this study were to develop genetic markers and to quantify genetic diversity within cañahua. A set of 43 wild and cultivated cañahua genotypes and two related species (Chenopodium quinoa Willd. and Chenopodium petiolare Kunth) were evaluated for polymorphism using 192 microsatellite markers derived from random genomic cañahua sequences produced by 454 pyrosequencing of cañahua genomic DNA. Another 424 microsatellite markers from C. quinoa were also evaluated for cross-species amplification and polymorphism in cañahua. A total of 34 polymorphic microsatellite marker loci were identified which detected a total of 154 alleles with an average of 4.5 alleles per marker locus and an average heterozygosity value of 0.49. A cluster analysis, based on Nei genetic distance, clearly separated from wild cañahua genotypes from the cultivated genotypes. Within the cultivated genotypes, subclades were partitioned by AMOVA analysis into six model-based clusters, including a subclade consisting sole of erect morphotypes. The isolation by distance test displayed no significant correlation between geographic collection origin and genotypic data, suggesting that cañahua populations have moved extensively, presumably via ancient food exchange strategies among native peoples of the Andean region. The molecular markers reported here are a significant resource for ongoing efforts to characterize the extensive Bolivian and Peruvian cañahua germplasm banks, including the development of core germplasm collections needed to support emerging breeding programs.     

Natural diversity of inflorescence architecture traces cryptic domestication genes in barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.)

Many-grained mutants occurring spontaneously among their less well-endowed field mates may have appeared to early farmers as fortunate twists of fate foreboding wealth and abundance. In domesticated barley, the number of kernel rows in spike can be tripled by recessive mutant alleles at the Six-rowed spike 1 (vrs1) locus that abolish the suppression of lateral spikelet fertility. In another barley row-type, so called intermedium-spike (int), lateral floret size is often intermediate between six-and two-rowed types. Phenotypic and sequence analyses of our intermedium-spike collection revealed that other genes can increase the size of florets and even stimulate occasional grain setting in lateral spikelets. Here, we show that a complete six-rowed phenotype occurs in a diverse panel of intermedium-spike barley carrying wildtype Vrs1 in the presence of the Int-c.a allele of the intermedium spike-c (int-c) gene, previously considered only as a modifier of lateral spikelet fertility. Int-c.a-type alleles had arisen before domestication and are associated with the enlargement of lateral florets in wild barley, suggesting that natural selection/evolution acts towards reduced lateral floret size. Since Int-c.a cannot overcome the suppression of lateral florets in the genomic background of wild barleys, we infer the existence of other gene loci, at which novel alleles or allelic combinations were selected for after domestication, to increase grain number of barley independently of Vrs1.     

Diversity in Allium ampeloprasum: from small and wild to large and cultivated

Allium ampeloprasum evolved as a complex of different cyto- and morpho-types widely distributed either in the wild or domesticated range of the Mediterranean regions. The assessment of genetic and phylogenetic relationships between Tunisian A. ampeloprasum and specimens from different origins and with variable degree of domestication can promote conservation and breeding. Minisatellite M13, microsatellite (GTG)₅ and nucleotide sequence analysis of the internal transcribed spacer region (ITS) were used to assess DNA polymorphism and genetic diversity. M13 and (GTG)₅ molecular markers efficiently discriminated A. ampeloprasum gene-pool from A. sativum. Geographic genetic patterns of variation of the wild gene-pool were not detected. However, domesticated A. ampeloprasum (great headed garlic, kurrat and leek) clustered consistently within the ampeloprasum group. A. sativum was found to be closer to A. ampeloprasum than A. fistulosum and A. schoenoprasum. A high number of single point mutations (SNPs) was recorded over the ITS1-2 spacer sequence. Most of these SNPs were heterozygous only in great headed garlic. It is inferred that heterozygosity played the major role in promoting great headed garlic domestication. Thus, great headed garlic adaptation to horticultural conditions along with its yield trait sizes are mainly associated to heterozygosity rather than to polyploidy.     

Genetic identity and relationships of Iranian apple (Malus?×?domestica Borkh.) cultivars and landraces, wild Malus species and representative old apple cultivars based on simple sequence repeat (SSR) marker analysis

In order to shed light on the role of Iran in apple evolution and domestication, we chose to investigate the relationships of a collection of 159 accessions of wild and domesticated apples including Iranian indigenous apple cultivars and landraces, selected wild species, and old apple scion and rootstock cultivars from different parts of the world. The majority of the wild species belonged to M. sieversii, which is widely believed to be the main maternal wild ancestor of domestic apples, from Kazakhstan and M. orientalis, which is one of the probable minor ancestors of domestic apples, from Turkey and Russia located on the east and west of Iran, respectively. The accessions were assigned into six arbitrary populations for the purpose of generating information on genetic parameters. Nine simple sequence repeat (SSR) loci selected from previous studies in apple were screened over DNA extracted from all the accessions. Results showed that all SSR loci displayed a very high degree of polymorphism with 11–25 alleles per locus. In total, there were 153 alleles across all loci with an average of 17 alleles per locus. The SSR allelic data were then used for estimation of population genetic parameters, including genetic variation statistics, F-statistics, gene flow, genetic identity, genetic distance and then cluster analysis using POPGENE 1.32 software. The F-statistics and gene flow in particular, showed that there was more intra-population than between population variation. The genetic identity and genetic distance estimates, and the dendrogram generated from the un-weighted pair group arithmetic average (UPGMA) method of cluster analysis showed that the Iranian cultivars and landraces were more closely related to M. sieversii from Central Asia (east of Iran) and M. orientalis native to Turkey and Russia than to other accessions of Malus species. Also, the old apple cultivars from different parts of the world have a closer genetic relationship to M. sieversii, M. orientalis and the Iranian apples, than to other wild species. Based on these results, we suggest that the Iranian apples may occupy an intermediate position between the domesticated varieties and wild species. We propose that Iran could be one of the major players in apples’ domestication and transfer from Central Asia to the western countries.     

Analysis of average standardized SSR allele size supports domestication of soybean along the Yellow River

Soybean (Glycine max) was domesticated in China from its wild progenitor G. soja. The geographic region of domestication is, however, not exactly known. Here we employed the directional evolution of SSR (microsatellite) repeats (which mutate preferentially into longer alleles) to analyze the domestication process and to infer the most ancestral soybean landraces. In this study, the average standardized SSR allele sizes across 42 SSR loci in 62 accessions of G. soja were determined, and compared with those in 1504 landraces of G. max, collected from all over China and representing the diversity in the gene bank. The standardized SSR allele size in the landraces (0.009) was significantly (P = 8.63 × 10^?58) larger than those in G. soja (?0.406). Pairwise comparisons between inferred clusters and sub-clusters of Chinese landraces indicated that the average standardized SSR allele size also increased with the further differentiation of landraces populations. Spring-sowed types had the shortest size, followed by summer-sown types, while the sub-cluster of autumn-sown type had the largest length. The spring-sowed landraces located near the middle region along the Yellow River had the smallest allele sizes, indicating that this is the most ancestral population of cultivated soybean. We concluded that soybean was most likely domesticated in the middle region of the Yellow River in central China, initially as a spring-sown type.     

Inter-MITE polymorphisms of a newly identified MITE show relationships among sugarcane (Saccharum) species

Sugarcane, one of the most important tropical crops, belongs the genus Saccharum. This genus consists of six species, four cultivated and two wild. The domestication histories of the four cultivated Saccharum species is an interesting and important topic of study. Previous studies have categorized the four cultivated species into two groups, one consisting only of S. edule and the other comprising S. officinarum, S. sinense and S. barberi. All four species have inherited the genomic DNA of S. robustum, one of two wild relative species. Saccharum species have large genomes with complex structures, as evidenced by chromosomes with a high degree of polyploidy, alloploidy and aneuploidy. Miniature inverted-repeat transposable elements (MITEs) are class II (DNA) transposons that disperse throughout the plant genome. In this study, a Tourist family MITE sequence with 18/19-bp terminal inverted repeats and a 2-bp target site duplication was newly identified from genomic DNA of S. robustum. The abundant accumulation of this MITE sequence in the sugarcane genome enabled the application of inter-MITE polymorphism (IMP) analysis to Saccharum. IMP analysis revealed the genetic relationships among all six Saccharum species and the domestication histories of the four cultivated species.     

Genetic diversity in wild cereals: regional and local studies and their bearing on conservation ex situ and in situ

The current alarming global crisis and extinction of biodiversity affect negatively the planet's biosphere. Conservation of biodiversity is one attempt to alleviate the pending extinction of the biosphere by humans. Genetic diversity, the basis of evolution by natural selection, is gravely threatened in the progenitors of cultivated plants and its exploration, evaluation, conservation in situ and ex situ is imperative to guarantee sustainable development. This is illustrated by the population genetics and ecology of two important progenitors of cereals wild, wheat and barley. The wild cereals are rich in adaptive genetic diversity in the Fertile Crescent, primarily in Israel, which is their center of origin and diversity. The 40–55% intrapopulation diversity level in the wild cereals contrasts sharply with the average of 80% in outbreeders. Genetic diversity in wild wheat and barley is structured, particularly in wild emmer wheat, as an 'archipelago' ecological and genetic structure. These include central, semi-isolated and ecologically peripheral and marginal isolated populations, where specific alleles and allele combinations predominate as coadapted blocks of genes, adaptive to diverse ecological stresses. These involve both physical (climatic and edaphic) and biotic (pathogens and parasites) stresses at macro- and microgeographical scales. Complementary in situ and ex situ conservation is imperative across the geographic range of these species, to safeguard their immensely important genetic resources for crop improvement.     

Evidence for two domestication events of hyacinth bean (<Emphasis Type="Italic">Lablab purpureus</Emphasis> (L.) Sweet): a comparative analysis of population genetic data

Studying molecular genetic relationships can substantially contribute to the understanding of the pathways of domestication of a species. Although an increasing number of molecular genetic studies have been performed on Lablab purpureus (hyacinth bean), many covered germplasm of restricted geographic origin or limited intra-specific systematic position. Integrating the molecular diversity found with phenotypic or morpho-agronomic diversity is also deficient. This investigation combines findings of eight molecular genetic studies that include about 400 accessions of both wild and cultivated germplasm, thus providing the largest assessment of diversity in Lablab purpureus to date. In particular, results from a recent molecular investigation (Robotham and Chapman 2015) are revisited and reinterpreted by integrating them with known phenotypic diversity. Wild accessions clearly fall into two types, with characteristic pods—2-seeded and 4-seeded. The large majority of cultivated types are more closely related to 4-seeded pod-types. Certain cultivated 2-seeded pod-type accessions from Ethiopia are genetically closer to wild 2-seeded pod-types. These two major phenotypes are reflected in two chloroplast DNA haplotypes A and B. Hence, two domestication events appear to exist in L. purpureus based on this combined data. No other geographic patterns of diversity, which might assist to trace the dispersal of L. purpureus, were found as cultivated accessions predominantly fell into 2-3 major groups. In all studies, the greatest genetic diversity was found in Africa, making Ethiopia one of the probable centers of domestication.     

Population genetic structure of in situ wild Sorghum bicolor in its Ethiopian center of origin based on SSR markers

In situ population studies of wild relatives of crops are crucial for the conservation of plant genetic resources, especially in regions with high genetic diversity and a risk of local extinction. Ethiopia is the center of origin for sorghum, yet little is known about the genetic structure of extant wild populations. Using 9 Simple Sequence Repeat loci, we characterized 19 wild populations from five regions, 8 local cultivar populations from three regions, and 10 wild sorghum accessions from several African countries. To our knowledge, this is the most comprehensive study to date of in situ wild sorghum populations in Africa. Genetic diversity corrected for sample size was significantly greater in the wild populations in situ than in local cultivars or the accessions. Approximately 41 % of the genetic variation in the wild plants was partitioned among populations, indicating a high degree of differentiation and potential value for germplasm conservation, and the average number of migrants (N_m) per generation was 0.43. Cluster analyses showed that some wild populations were grouped by geographic region, whereas others were not, presumably due to long-distance seed movement. Four wild populations from disjunct regions formed a unique cluster with an Ethiopian accession of subsp. drummondii and probably represent a weedy race. STRUCTURE and other analyses detected evidence for crop-wild hybridization, consistent with previous molecular marker studies in Kenya, Mali, and Cameroon. In summary, in situ wild sorghum populations in Ethiopia harbor substantial genetic diversity and differentiation, despite their close proximity to conspecific cultivars in this crop/wild/weedy complex.     

Characterization of single nucleotide polymorphism in Tunisian grapevine genome and their potential for population genetics and evolutionary studies

In this study, two gene fragments corresponding to the VvMYBA1 and VvMYBA2 loci were sequenced on a sample of grapes including cultivated and wild accessions originating from Tunisia, Germany and France. A total of 42 SNPs were detected in the sequenced fragments giving an average of 1 SNP every 33 bp. High level of polymorphism was observed in the samples either in cultivated or wild accessions. Pattern of nucleotide diversity indicates a non departure from neutrality expectations for wild grapevine sample for gene VvMYBA1 and VvMYBA2 and for cultivated sample for gene VvMYBA1. However, a linkage to a selective sweep was revealed for cultivated grapevine gene pool in gene VvMYBA2. A genetic structure of the studied sample according to accession taxonomic status was revealed by the UPGMA clustering with a considerable overlap. This result was confirmed by significant but low genetic differentiation values between cultivated and wild sample. The number of migrants Nm based on sequence data information between Tunisian cultivars and Tunisian wild accessions showed a low level of gene flow between those germplasms. This finding indicates that Tunisian cultivars do not derive directly from local wild populations but could mostly correspond to imported materials introduced during historical times. However, the possibility that some cultivars derived from ancestral events of local domestication or cross hybridization with native wild plants was not completely excluded for Tunisian grapevine accessions.     

Genetic differentiation and diversity of phenotypic characters in Chinese wild soybean (<Emphasis Type="Italic">Glycine soja</Emphasis> Sieb. et Zucc.) revealed by nuclear SSR markers and the implication for intraspecies phylogenic relationship of characters

Genetic diversity is reduced from wild soybean to cultivars and from landraces to modern varieties. However, intraspecies genetic diversity loss between characters or phenotypes also existed in wild soybean. We revealed the phylogenic relationship of character types in Chinese wild soybean using 42 SSR markers. We conjectured that white flower, no-seed bloom, grey pubescence, and four seed coat colours were evolutionarily acquired phenotypes. There were a small decrease (∆H = 0.1–6.46%) of gene diversity and a moderate reduction (∆Na = 10.81–53.54%) of number of alleles but a violent loss (80.74–98.59%) of unique alleles in the acquired phenotypes. Our results seemed to suggest that ovoid and elliptic leaves were differentiated at the earliest and subsequently lanceolate leaf appeared before the domestication of soybeans within wild soybean, and that G. gracilis type was another earliest type, maybe emerged since the appearance of soybeans or it was concomitant with the domestication of soybean.     

Using diversity of the chloroplast genome to examine evolutionary history of wheat species

Chloroplast microsatellites (SSRs) are conserved within wheat species, yet are sufficiently polymorphic between and within species to be useful for evolutionary studies. This study describes the relationships among a very large set of accessions of Triticum urartu Thum. ex Gandil., T. dicoccoides (Körn. ex Asch. et Graebn.) Schweinf., T. dicoccon Schrank, T. durum Desf., T. spelta L., and T. aestivum L. s. str. based on their cpSSR genotypes. By characterising the chloroplast diversity in each wheat species in the evolutionary series, the impact on diversity of major evolutionary events such as domestication and polyploidyisation was assessed. We detected bottlenecks associated with domestication, polyploidisation and selection, yet these constrictions were partially offset by mutations in the chloroplast SSR loci that generated new alleles. The discrete cpSSR alleles and haplotypes observed in T. urartu and Aegilops tauschii, combined with other species specific polymorphisms, provide very strong evidence that concur with current opinion that neither species was the maternal and thus cytoplasmic donor for polyploid wheats. Synthetic hexaploid wheats possessed the same chloroplast haplotypes as their tetraploid progenitors demonstrating how the novel synthetic wheat lines have captured chloroplast diversity from the maternal parents, the chloroplast is maternally inherited and novel alleles are not created by genomic rearrangements triggered by the polyploidisation event.