Genetic Analysis of Hexaploid Wheat (Triticum aestivum L.) Using the Complete Sequencing of Chloroplast DNA and Haplotype Analysis of the Wknox1 Gene.

The aim of the presented study is a genetic characterization of the hexaploid wheat Triticum aestivum L. Two approaches were used for the genealogical study of hexaploid wheats-the complete sequencing of chloroplast DNA and PCR-based haplotype analysis of the fourth intron of Wknox1d and of the fifth-to-sixth-exon region of Wknox1b. The complete chloroplast DNA sequences of 13 hexaploid wheat samples were determined: Free-threshing-T. aestivum subsp. aestivum, one sample; T. aestivum subsp. compactum, two samples; T. aestivum subsp. sphaerococcum, one sample; T. aestivum subsp. carthlicoides, four samples. Hulled-T. aestivum subsp. spelta, three samples; T. aestivum subsp. vavilovii jakubz., two samples. The comparative analysis of complete cpDNA sequences of 20 hexaploid wheat samples (13 samples in this article plus 7 samples sequenced in this laboratory in 2018) was carried out. PCR-based haplotype analysis of the fourth intron of Wknox1d and of the fifth-to-sixth exon region of Wknox1b of all 20 hexaploid wheat samples was carried out. The 20 hexaploid wheat samples (13 samples in this article plus 7 samples in 2018) can be divided into two groups-T. aestivum subsp. spelta, three samples and T. aestivum subsp. vavilovii collected in Armenia, and the remaining 16 samples, including T. aestivum subsp. vavilovii collected in Europe (Sweden). If we take the cpDNA of Chinese Spring as a reference, 25 SNPs can be identified. Furthermore, 13-14 SNPs can be identified in T. aestivum subsp. spelta and subsp. vavilovii (Vav1). In the other samples up to 11 SNPs were detected. 22 SNPs are found in the intergenic regions, 2 found in introns, and 10 SNPs were found in the genes, of which seven are synonymous. PCR-based haplotype analysis of the fourth intron of Wknox1d and the fifth-to-sixth-exon region of Wknox1b provides an opportunity to make an assumption that hexaploid wheats T. aestivum subsp. macha var. palaeocolchicum and var. letshckumicum differ from other macha samples by the absence of a 42 bp insertion in the fourth intron of Wknox1d. One possible explanation for this observation would be that two Aegilops tauschii Coss. (A) and (B) participated in the formation of hexaploids through the D genome: Ae. tauschii (A)-macha (1-5, 7, 8, 10-12), and Ae. tauschii (B)-macha M6, M9, T. aestivum subsp. aestivum cv. 'Chinese Spring' and cv. 'Red Doly'.

Origin and Dynamics of Rwt6, a Wheat Gene for Resistance to Nonadapted Pathotypes of Pyricularia oryzae.

Avirulence of Eleusine isolates of Pyricularia oryzae on common wheat is conditioned by at least five avirulence genes. One is PWT3 corresponding to resistance gene Rwt3 located on chromosome 1D. We identified a resistance gene corresponding to a second avirulence gene, PWT6, and named it Rmg9 (Rwt6). Rwt6 was closely linked to Rwt3. A survey of the population of Aegilops tauschii, the D genome donor to common wheat, revealed that some accessions from the southern coastal region of the Caspian Sea, the birthplace of common wheat, carried both genes. Rwt6 and Rwt3 carriers accounted for 65 and 80%, respectively, of accessions in a common wheat landrace collection. The most likely explanation of our results is that both resistance genes were simultaneously introduced into common wheat at the time of hybridization of Triticum turgidum and A. tauschii. However, a prominent difference was recognized in their geographical distributions in modern wheat; Rwt3 and Rwt6 co-occurred at high frequencies in regions to the east of the Caspian Sea, whereas Rwt6 occurred at a lower frequency than Rwt3 in regions to the west. This difference was considered to be associated with range of pathotypes to which these genes were effective. A. tauschii accessions carrying Rwt3 and Rwt6 also carried Rwt4, another resistance gene involved in the species specificity. We suggest that the gain of the D genome should have given an adaptive advantage to the genus Triticum by conferring disease resistance.

Genome-Wide Association Study of Morpho-Physiological Traits in Aegilops tauschii to Broaden Wheat Genetic Diversity.

Aegilops tauschii, the D-genome donor of bread wheat, is a storehouse of genetic diversity that can be used for wheat improvement. This species consists of two main lineages (TauL1 and TauL2) and one minor lineage (TauL3). Its morpho-physiological diversity is large, with adaptations to a wide ecological range. Identification of allelic diversity in Ae. tauschii is of utmost importance for efficient breeding and widening of the genetic base of wheat. This study aimed at identifying markers or genes associated with morpho-physiological traits in Ae. tauschii, and at understanding the difference in genetic diversity between the two main lineages. We performed genome-wide association studies of 11 morpho-physiological traits of 343 Ae. tauschii accessions representing the entire range of habitats using 34,829 DArTseq markers. We observed a wide range of morpho-physiological variation among all accessions. We identified 23 marker-trait associations (MTAs) in all accessions, 15 specific to TauL1 and eight specific to TauL2, suggesting independent evolution in each lineage. Some of the MTAs could be novel and have not been reported in bread wheat. The markers or genes identified in this study will help reveal the genes controlling the morpho-physiological traits in Ae. tauschii, and thus in bread wheat even if the plant morphology is different.

Reproductive and genetic roles of the maternal progenitor in the origin of common wheat (Triticum aestivum L.).

Common wheat (Triticum aestivum L., AABBDD genome) is thought to have emerged through natural hybridization between Triticum turgidum L. (AABB genome) and Aegilops tauschii Coss. (DD genome). Hybridization barriers and doubling of the trihaploid F1 hybrids' genome (ABD) via unreduced gamete fusion had key roles in the process. However, how T. turgidum, the maternal progenitor, was involved in these mechanisms remains unknown. An artificial cross-experiment using 46 cultivated and 31 wild T. turgidum accessions and a single Ae. tauschii tester with a very short genetic distance to the common wheat D genome was conducted. Cytological and quantitative trait locus analyses of F1 hybrid genome doubling were performed. The crossability and ability to cause hybrid inviability did not greatly differ between the cultivars and wild accessions. The ability to cause hybrid genome doubling was higher in the cultivars. Three novel T. turgidum loci for hybrid genome doubling, which influenced unreduced gamete production in F1 hybrids, were identified. Cultivated T. turgidum might have increased the probability of the emergence of common wheat through its enhanced ability to cause genome doubling in F1 hybrids with Ae. tauschii. The ability enhancement might have involved alterations at a relatively small number of loci.

The role of reproductive isolation in allopolyploid speciation patterns: empirical insights from the progenitors of common wheat.

The ability to cause reproductive isolation often varies among individuals within a plant species. We addressed whether such polymorphism influenced speciation of the allopolyploid common wheat (Triticum aestivum L., AABBDD genome) by evaluating the expression of pre-pollination (outcrossing potential) and post-pollination (crossability) barriers in Aegilops tauschii Coss. (the D genome progenitor). In total, 201 Ae. tauschii accessions representing the entire natural habitat range of the species were used for anther length measurement and artificial crosses with a Triticum turgidum L. (the AB genome progenitor) tester. Intraspecific comparisons showed that both barriers were more strongly expressed in the TauL1 lineage than in the TauL2 lineage. The ability of Ae. tauschii to cause reproductive isolation in the hybridisation with T. turgidum might have markedly influenced common wheat's speciation by inducing lineage-associated patterns of gene flow. The TauL2 accessions with high potential for natural hybridisation with T. turgidum clustered in the southern coastal Caspian region. This provided phenotypic support for the derivation of the D genome of common wheat from southern Caspian populations. The present study underscored the importance of approaches that incorporate the genealogical and geographic structure of the parental species' reproductive isolation in understanding the mechanism of plant allopolyploid speciation.

Establishment of Imaging Spectroscopy of Nuclear Gamma-Rays based on Geometrical Optics.

Since the discovery of nuclear gamma-rays, its imaging has been limited to pseudo imaging, such as Compton Camera (CC) and coded mask. Pseudo imaging does not keep physical information (intensity, or brightness in Optics) along a ray, and thus is capable of no more than qualitative imaging of bright objects. To attain quantitative imaging, cameras that realize geometrical optics is essential, which would be, for nuclear MeV gammas, only possible via complete reconstruction of the Compton process. Recently we have revealed that "Electron Tracking Compton Camera" (ETCC) provides a well-defined Point Spread Function (PSF). The information of an incoming gamma is kept along a ray with the PSF and that is equivalent to geometrical optics. Here we present an imaging-spectroscopic measurement with the ETCC. Our results highlight the intrinsic difficulty with CCs in performing accurate imaging, and show that the ETCC surmounts this problem. The imaging capability also helps the ETCC suppress the noise level dramatically by ~3 orders of magnitude without a shielding structure. Furthermore, full reconstruction of Compton process with the ETCC provides spectra free of Compton edges. These results mark the first proper imaging of nuclear gammas based on the genuine geometrical optics.

First On-Site True Gamma-Ray Imaging-Spectroscopy of Contamination near Fukushima Plant.

We have developed an Electron Tracking Compton Camera (ETCC), which provides a well-defined Point Spread Function (PSF) by reconstructing a direction of each gamma as a point and realizes simultaneous measurement of brightness and spectrum of MeV gamma-rays for the first time. Here, we present the results of our on-site pilot gamma-imaging-spectroscopy with ETCC at three contaminated locations in the vicinity of the Fukushima Daiichi Nuclear Power Plants in Japan in 2014. The obtained distribution of brightness (or emissivity) with remote-sensing observations is unambiguously converted into the dose distribution. We confirm that the dose distribution is consistent with the one taken by conventional mapping measurements with a dosimeter physically placed at each grid point. Furthermore, its imaging spectroscopy, boosted by Compton-edge-free spectra, reveals complex radioactive features in a quantitative manner around each individual target point in the background-dominated environment. Notably, we successfully identify a "micro hot spot" of residual caesium contamination even in an already decontaminated area. These results show that the ETCC performs exactly as the geometrical optics predicts, demonstrates its versatility in the field radiation measurement, and reveals potentials for application in many fields, including the nuclear industry, medical field, and astronomy.

Salt tolerance during germination and seedling growth of wild wheat Aegilops tauschii and its impact on the species range expansion.

Adaptation to edaphic stress may have a key role in plant species range expansion. Aegilops tauschii Coss., the common wheat's D-genome progenitor native to the Transcaucasus-Middle East region, is a good model to study the relationships between soil salinity and plant distributions: one of its intraspecific sublineages, TauL1b, drove the long-distance eastward expansion of this species range reaching semi-arid-central Asia. Salt tolerance during germination and seedling growth was evaluated in 206 Ae. tauschii accessions by treating seeds with NaCl solutions differing in concentrations. Differences in natural variation patterns were analyzed between sublineages and associated with natural edaphic condition variables, and then compared with reproductive trait variation patterns. The natural variations observed in NaCl-induced-stress tolerance had clear geographic and genetic structure. Seedling growth significantly increased in the TauL1b accessions that were collected from salt-affected soil habitats, whereas germinability did not. Principal component analysis suggested that the NaCl-induced-stress tolerances and reproductive traits might have had a similar degree of influence on Ae. tauschii's eastward range expansion. Adaptation to salt-affected soils through increased seedling growth was an important factor for the species' successful colonization of the semi-arid central Asian habitats. TauL1b accessions might provide useful genetic resources for salt-tolerant wheat breeds.

Intraspecific lineage divergence and its association with reproductive trait change during species range expansion in central Eurasian wild wheat Aegilops tauschii Coss. (Poaceae).

BACKGROUND: How species ranges form in landscapes is a matter of long-standing evolutionary interest. However, little is known about how natural phenotypic variations of ecologically important traits contribute to species range expansion. In this study, we examined the phylogeographic patterns of phenotypic changes in life history (seed production) and phenological (flowering time) traits during the range expansion of Aegilops tauschii Coss. from the Transcaucasus and Middle East to central Asia. RESULTS: Our comparative analyses of the patterns of natural variations for those traits and their association with the intraspecific lineage structure showed that (1) the eastward expansion to Asia was driven by an intraspecific sublineage (named TauL1b), (2) high seed production ability likely had an important role at the initial dispersal stage of TauL1b's expansion to Asia, and (3) the phenological change to early flowering phenotypes was one of the key adaptation events for TauL1b to further expand its range in Asia. CONCLUSIONS: This study provides for the first time a broad picture of the process of Ae. tauschii's eastward range expansion in which life history and phenological traits may have had respective roles in its dispersal and adaptation in Asia. The clear association of seed production and flowering time patterns with the intraspecific lineage divergence found in this study invites further genetic research to bring the mechanistic understanding of the changes in these key functional traits during range expansion within reach.

The cuticular wax inhibitor locus Iw2 in wild diploid wheat Aegilops tauschii: phenotypic survey, genetic analysis, and implications for the evolution of common wheat.

BACKGROUND: Cuticular wax production on plant surfaces confers a glaucous appearance and plays important roles in plant stress tolerance. Most common wheat cultivars, which are hexaploid, and most tetraploid wheat cultivars are glaucous; in contrast, a wild wheat progenitor, Aegilops tauschii, can be glaucous or non-glaucous. A dominant non-glaucous allele, Iw2, resides on the short arm of chromosome 2D, which was inherited from Ae. tauschii through polyploidization. Iw2 is one of the major causal genes related to variation in glaucousness among hexaploid wheat. Detailed genetic and phylogeographic knowledge of the Iw2 locus in Ae. tauschii may provide important information and lead to a better understanding of the evolution of common wheat. RESULTS: Glaucous Ae. tauschii accessions were collected from a broad area ranging from Armenia to the southwestern coastal part of the Caspian Sea. Linkage analyses with five mapping populations showed that the glaucous versus non-glaucous difference was mainly controlled by the Iw2 locus in Ae. tauschii. Comparative genomic analysis of barley and Ae. tauschii was then used to develop molecular markers tightly linked with Ae. tauschii Iw2. Chromosomal synteny around the orthologous Iw2 regions indicated that some chromosomal rearrangement had occurred during the genetic divergence leading to Ae. tauschii, barley, and Brachypodium. Genetic associations between specific Iw2-linked markers and respective glaucous phenotypes in Ae. tauschii indicated that at least two non-glaucous accessions might carry other glaucousness-determining loci outside of the Iw2 locus. CONCLUSION: Allelic differences at the Iw2 locus were the main contributors to the phenotypic difference between the glaucous and non-glaucous accessions of Ae. tauschii. Our results supported the previous assumption that the D-genome donor of common wheat could have been any Ae. tauschii variant that carried the recessive iw2 allele.

Genetic mechanisms of allopolyploid speciation through hybrid genome doubling: novel insights from wheat (Triticum and Aegilops) studies.

Polyploidy, which arises through complex genetic and ecological processes, is an important mode of plant speciation. This review provides an overview of recent advances in understanding why plant polyploid species are so ubiquitous and diverse. We consider how the modern framework for understanding genetic mechanisms of speciation could be used to study allopolyploid speciation that occurs through hybrid genome doubling, that is, whole genome doubling of interspecific F1 hybrids by the union of male and female unreduced gametes. We outline genetic and ecological mechanisms that may have positive or negative impacts on the process of allopolyploid speciation through hybrid genome doubling. We also discuss the current status of studies on the underlying genetic mechanisms focusing on the wheat (Triticum and Aegilops) hybrid-specific reproductive phenomena that are well known but deserve renewed attention from an evolutionary viewpoint.

Genetic basis for spontaneous hybrid genome doubling during allopolyploid speciation of common wheat shown by natural variation analyses of the paternal species.

The complex process of allopolyploid speciation includes various mechanisms ranging from species crosses and hybrid genome doubling to genome alterations and the establishment of new allopolyploids as persisting natural entities. Currently, little is known about the genetic mechanisms that underlie hybrid genome doubling, despite the fact that natural allopolyploid formation is highly dependent on this phenomenon. We examined the genetic basis for the spontaneous genome doubling of triploid F1 hybrids between the direct ancestors of allohexaploid common wheat (Triticum aestivum L., AABBDD genome), namely Triticumturgidum L. (AABB genome) and Aegilopstauschii Coss. (DD genome). An Ae. tauschii intraspecific lineage that is closely related to the D genome of common wheat was identified by population-based analysis. Two representative accessions, one that produces a high-genome-doubling-frequency hybrid when crossed with a T. turgidum cultivar and the other that produces a low-genome-doubling-frequency hybrid with the same cultivar, were chosen from that lineage for further analyses. A series of investigations including fertility analysis, immunostaining, and quantitative trait locus (QTL) analysis showed that (1) production of functional unreduced gametes through nonreductional meiosis is an early step key to successful hybrid genome doubling, (2) first division restitution is one of the cytological mechanisms that cause meiotic nonreduction during the production of functional male unreduced gametes, and (3) six QTLs in the Ae. tauschii genome, most of which likely regulate nonreductional meiosis and its subsequent gamete production processes, are involved in hybrid genome doubling. Interlineage comparisons of Ae. tauschii's ability to cause hybrid genome doubling suggested an evolutionary model for the natural variation pattern of the trait in which non-deleterious mutations in six QTLs may have important roles. The findings of this study demonstrated that the genetic mechanisms for hybrid genome doubling could be studied based on the intrinsic natural variation that exists in the parental species.

Evolution of polyploid triticum wheats under cultivation: the role of domestication, natural hybridization and allopolyploid speciation in their diversification.

The evolution of the polyploid Triticum wheats is distinctive in that domestication, natural hybridization and allopolyploid speciation have all had significant impacts on their diversification. In this review, I outline the phylogenetic relationships of cultivated wheats and their wild relatives and provide an overview of the recent progress and remaining issues in understanding the genetic and ecological factors that favored their evolution. An attempt is made to view the evolution of the polyploid Triticum wheats as a continuous process of diversification that was initiated by domestication of tetraploid emmer wheat and driven by various natural events ranging from interploidy introgression via hybridization to allopolyploid speciation of hexaploid common wheat, instead of viewing it as a group of discrete evolutionary processes that separately proceeded at the tetraploid and hexaploid levels. This standpoint underscores the important role of natural hybridization in the reticulate diversification of the tetraploid-hexaploid Triticum wheat complex and highlights critical, but underappreciated, issues that concern the allopolyploid speciation of common wheat.

Applicability of Aegilops tauschii drought tolerance traits to breeding of hexaploid wheat.

Few genes are available to develop drought-tolerant bread wheat (Triticum aestivum L.) cultivars. One way to enhance bread wheat's genetic diversity would be to take advantage of the diversity of wild species by creating synthetic hexaploid wheat (SW) with the genomic constitution of bread wheat. In this study, we compared the expression of traits encoded at different ploidy levels and evaluated the applicability of Aegilops tauschii drought-related traits using 33 Ae. tauschii accessions along with their corresponding SW lines under well-watered and drought conditions. We found wide variation in Ae. tauschii, and even wider variation in the SW lines. Some SW lines were more drought-tolerant than the standard cultivar Cham 6. Aegilops tauschii from some regions gave better performing SW lines. The traits of Ae. tauschii were not significantly correlated with their corresponding SW lines, indicating that the traits expressed in wild diploid relatives of wheat may not predict the traits that will be expressed in SW lines derived from them. We suggest that, regardless of the adaptability and performance of the Ae. tauschii under drought, production of SW could probably result in genotypes with enhanced trait expression due to gene interactions, and that the traits of the synthetic should be evaluated in hexaploid level.

Population structure of wild wheat D-genome progenitor Aegilops tauschii Coss.: implications for intraspecific lineage diversification and evolution of common wheat.

Aegilops tauschii Coss. is the D-genome progenitor of hexaploid wheat. Aegilops tauschii, a wild diploid species, has a wide natural species range in central Eurasia, spreading from Turkey to western China. Amplified fragment length polymorphism (AFLP) analysis using a total of 122 accessions of Ae. tauschii was conducted to clarify the population structure of this widespread wild wheat species. Phylogenetic and principal component analyses revealed two major lineages in Ae. tauschii. Bayesian population structure analyses based on the AFLP data showed that lineages one (L1) and two (L2) were respectively significantly divided into six and three sublineages. Only four out of the six L1 sublineages were diverged from those of western habitats in the Transcaucasia and northern Iran region to eastern habitats such as Pakistan and Afghanistan. Other sublineages including L2 were distributed to a limited extent in the western region. Subspecies strangulata seemed to be differentiated in one sublineage of L2. Among three major haplogroups (HG7, HG9 and HG16) previously identified in the Ae. tauschii population based on chloroplast variation, HG7 accessions were widely distributed to both L1 and L2, HG9 accessions were restricted to L2, and HG16 accessions belonged to L1, suggesting that HG9 and HG16 were formed from HG7 after divergence of the first two lineages of the nuclear genome. These results on the population structure of Ae. tauschii and the genealogical relationship among Ae. tauschii accessions should provide important agricultural and evolutionary knowledge on genetic resources and conservation of natural genetic diversity.

Evolutionary dynamics of wheat mitochondrial gene structure with special remarks on the origin and effects of RNA editing in cereals.

We investigated the evolutionary dynamics of wheat mitochondrial genes with respect to their structural differentiation during organellar evolution, and to mutations that occurred during cereal evolution. First, we compared the nucleotide sequences of three wheat mitochondrial genes to those of wheat chloroplast, alpha-proteobacterium and cyanobacterium orthologs. As a result, we were able to (1) differentiate the conserved and variable segments of the orthologs, (2) reveal the functional importance of the conserved segments, and (3) provide a corroborative support for the alpha-proteobacterial and cyanobacterial origins of those mitochondrial and chloroplast genes, respectively. Second, we compared the nucleotide sequences of wheat mitochondrial genes to those of rice and maize to determine the types and frequencies of base changes and indels occurred in cereal evolution. Our analyses showed that both the evolutionary speed, in terms of number of base substitutions per site, and the transition/transversion ratio of the cereal mitochondrial genes were less than two-fifths of those of the chloroplast genes. Eight mitochondrial gene groups differed in their evolutionary variability, RNA and Complex I (nad) genes being most stable whereas Complex V (atp) and ribosomal protein genes most variable. C-to-T transition was the most frequent type of base change; C-to-G and G-to-C transversions occurred at lower rates than all other changes. The excess of C-to-T transitions was attributed to C-to-U RNA editing that developed in early stage of vascular plant evolution. On the contrary, the editing of C residues at cereal T-to-C transition sites developed mostly during cereal divergence. Most indels were associated with short direct repeats, suggesting intra- and intermolecular recombination as an important mechanism for their origin. Most of the repeats associated with indels were di- or trinucleotides, although no preference was noticed for their sequences. The maize mt genome was characterized by a high incidence of indels, comparing to the wheat and rice mt genomes.

Flowering time diversification and dispersal in central Eurasian wild wheat Aegilops tauschii Coss.: genealogical and ecological framework.

Timing of flowering is a reproductive trait that has significant impact on fitness in plants. In contrast to recent advances in understanding the molecular basis of floral transition, few empirical studies have addressed questions concerning population processes of flowering time diversification within species. We analyzed chloroplast DNA genealogical structure of flowering time variation in central Eurasian wild wheat Aegilops tauschii Coss. using 200 accessions that represent the entire species range. Flowering time measured as days from germination to flowering varied from 144.0 to 190.0 days (average 161.3 days) among accessions in a common garden/greenhouse experiment. Subsequent genealogical and statistical analyses showed that (1) there exist significant longitudinal and latitudinal clines in flowering time at the species level, (2) the early-flowering phenotype evolved in two intraspecific lineages, (3) in Asia, winter temperature was an environmental factor that affected the longitudinal clinal pattern of flowering time variation, and (4) in Transcaucasus-Middle East, some latitudinal factors affected the geographic pattern of flowering time variation. On the basis of palaeoclimatic, biogeographic, and genetic evidence, the northern part of current species' range [which was within the temperate desert vegetation (TDV) zone at the Last Glacial Maximum] is hypothesized to have harbored species refugia. Postglacial southward dispersal from the TDV zone seems to have been driven by lineages that evolved short-flowering-time phenotypes through different genetic mechanisms in Transcaucasus-Middle East and Asia.

Population structure and genetic diversity of New World maize races assessed by DNA microsatellites.

Because of the economic importance of maize and its scientific importance as a model system for studies of domestication, its evolutionary history is of general interest. We analyzed the population genetic structure of maize races by genotyping 964 individual plants, representing almost the entire set of ∼350 races native to the Americas, with 96 microsatellites. Using Bayesian clustering, we detected four main clusters consisting of highland Mexican, northern United States (US), tropical lowland, and Andean races. Phylogenetic analysis indicated that the southwestern US was an intermediary stepping stone between Mexico and the northern US. Furthermore, southeastern US races appear to be of mixed northern flint and tropical lowland ancestry, while lowland middle South American races are of mixed Andean and tropical lowland ancestry. Several cases of post-Columbian movement of races were detected, most notably from the US to South America. Of the four main clusters, the highest genetic diversity occurs in highland Mexican races, while diversity is lowest in the Andes and northern US. Isolation by distance appears to be the main factor underlying the historical diversification of maize. We identify highland Mexico and the Andes as potential sources of genetic diversity underrepresented among elite lines used in maize breeding programs.

Natural variation for fertile triploid F1 hybrid formation in allohexaploid wheat speciation.

The tempo, mode, and geography of allopolyploid speciation are influenced by natural variation in the ability of parental species to express postzygotic reproductive phenotypes that affect hybrid fertility. To shed light on the impact of such natural variations, we used allohexaploid Triticum aestivum wheats' evolution as a model and analyzed the geographic and phylogenetic distributions of Aegilops tauschii (diploid progenitor) accessions involved in the expression of abnormality and fertility in triploid F(1) hybrids with Triticum turgidum (tetraploid progenitor). Artificial-cross experiments and chloroplast-DNA-based evolutionary analyses showed that hybrid-abnormality-causing accessions had limited geographic and phylogenetic distributions, indicative that postzygotic hybridization barriers are underdeveloped between these species. In contrast, accessions that are involved with fertile triploid F(1) hybrid formation have wide geographic and phylogenetic distributions, indicative of a deep evolutionary origin. Wide-spread hybrid-fertilizing accessions support the theory that T. aestivum speciation occurred at multiple sites within the species range of Ae. tauschii, in which existing conditions enabled natural hybridization with T. turgidum. Implications of our findings on how natural variation in the ability of Ae. tauschii to express those postzygotic reproductive phenotypes diversified and contributed to the speciation of T. aestivum are discussed.