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.     

Genetic diversity in farmer grown spring barley material from Kyrgyzstan

Barley (Hordeum vulgare L.) is one of the economically important cereal crops grown in the highlands of the Kyrgyz Republic. In marginal agricultural areas of the country barley is considered as irreplaceable and very suitable fodder crop for livestock. In this study the genetic diversity of currently grown farmers’ spring barley material from Naryn and Issik-Kul provinces is described. In order to capture maximum diversity present in farmer field’s different morphological types were collected. The 22 spring barley accessions collected in 2008 were described morphologically and analysed using fourteen simple sequences repeat (SSR) markers. Most material was two-rowed barley, 78 %, whereas six-rowed barley was only 22 %. The spike colours present were light yellow, dark yellow, brown and violet. The material varied also for such characters as length of spike and number of kernels per spike. For the SSR loci 129 alleles were detected with an average of 9.3 alleles per locus and a genetic diversity with an average of h = 0.721 per locus. Based on analysis of molecular variance the 71.3 % of genetic diversity accounted for within accession variation, whereas 27.4 % for among accession variation. The variation between provinces was not significantly different. A group of two-rowed barley was identified that was more closely related to six-rowed barley than to the rest of the two-rowed barley material. Two-rowed material currently grown in Kyrgyzstan is more diverse than six-rowed barley (h = 0.65, A = 8.8 vs. h = 0.59, A = 3.1). From this study we could conclude that the materials nowadays grown by farmers are mixtures. Since there are no geographic differentiation between these mixtures suggests that materials is frequently inter-exchanged between farmers and are most probably not landrace material. The question concerning the origin of these mixtures remains unsolved and further investigation will be carried out to identify sources of variation in farmers’ field.     

Haplotype variations of gene Ppd-D1 in Aegilops tauschii and their implications on wheat origin

The Ppd-D1 controlling photoperiod response is an important gene for wheat adaptation since it affects heading time. In the present study, three haplotypes, i.e. haplotype I without deletion, haplotype II with a 24?bp deletion, and haplotype III with two deletions of 24 and 15?bp, were identified in the upstream of the coding region in 80 Ae. tauschii accessions. The haplotype distribution was related to subspecies taxon. All typical ssp. tauschii accessions had haplotype I, whereas all ssp. strangulata had haplotype III. The three haplotypes were observed in Ae. tauschii with morphologically intermediate forms between the two typical subspecies. Present results supported that ssp. strangulata or intermediate form was the D-genome donor of common wheat since only haplotype III were found in wheat. Moreover, a 16?bp deletion in exon 8 of gene Ppd-D1 exists in common wheat. However, none of Ae. tauschii accessions analyzed had the 16?bp deletion.     

Detection of genetic integrity of conserved maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) germplasm in genebanks using SNP markers

Twenty maize landrace accessions regenerated and conserved in five maize genebanks were investigated for genetic integrity using 1,150 Single Nucleotide Polymorphisms (SNPs) and 235 SNP haplotypes. The genetic diversity of three accessions changed significantly in terms of the average number of alleles per locus. Ten out of twenty accessions had significantly different SNP allelic frequencies, either after regeneration or in the same accession held in different genebanks. The proportion of loci with significant changes in SNP allelic frequency was very low (37/1,150). Changes in the major allelic frequency (MAF) for the majority of SNP loci (60.2–75.2%) were less than 0.05. For SNP haplotypes, the genetic diversity of four accessions changed significantly in terms of average number of haplotype alleles and polymorphic information content (PIC) per locus. The proportion of SNP haplotype alleles lost in the later generations ranged between 0 and 22.6%, and at the same time 0–19.9% of the SNP haplotype alleles appeared in later generations, however, these were absent in the earlier generations. Dynamic changes in genetic integrity, in terms of presence and absence of genes (alleles), by both SNP and SNP haplotype analysis were detected during regeneration. A suboptimum number of ears harvested in one generation can be combined with those from another, repeated regeneration to capture the diversity of the previous generation. Use of molecular markers during regeneration of accessions can help in understanding the extent of genetic integrity of the maize accessions in ex situ genebanks and in recommending the best practice for maintaining the original genetic diversity of the genebank accessions.     

Genetic diversity in Hordeum agriocrithon E. Åberg, six-rowed barley with brittle rachis, from Tibet

Twenty-two accessions of H. agriocrithon from Tibet were investigated for morphological and physiological characters, and genotypes combined alleles at the tightly linked Est1, Est2 and Est4 complex locus encoding esterase isozymes. H. agriocrithon was similar in plant type to Tibetan cultivated barley, but different from H. spontaneum. The accessions of H. agriocrithon were classified by their spike-awn types into vars. eu-agriocrithon, dawoense and paradoxon, with one exceptional accession of the awnless naked type similar to var. nuditonsum of cultivated barley. They were characterized by spring growth habit, susceptibility to Japanese race I of powdery mildew, and either of the same three esterase genotypes as those of cultivated barley from Tibet, Bhutan and Ladakh. The similarity suggested that the esterase complex locus on chromosome 3H is closely linked with some QTL for adaptation to Tibetan conditions. On the origin of H. agriocrithon, it might be concluded that natural hybridization between H. spontaneum and six-rowed barley occurred in northern parts of Afghanistan, Pakistan or India, and seeds of their segregants mixed into barley or wheat were brought up to Tibet, resulting six-rowed barley with brittle rachis.     

Molecular characterization and diversity of puroindoline b-2 variants in cultivated and wild diploid wheat

Cloning and phylogenetic analysis of puroindoline b-2 variants in common wheat (Triticum aestivum L.) and its relatives would advance the understanding of the genetic diversity and evolution of puroindoline b-2 gene in common wheat and its related species. In the present study, common wheat (AABBDD) and four related species, including T. urartu (A^uA^u), Aegilops speltoides (SS), Ae. tauschii (DD), and T. turgidum (AABB) were sampled for the presence of novel alleles at Pinb2v-A1, Pinb2v-B1/Pinb2v-S1 and Pinb2v-D1 loci corresponding to common wheat puroindoline b-2 variants. Nine new alleles were identified at these loci, designated Pinb2v-A1a through Pinb2v-A1c, Pinb2v-S1a through Pinb2v-S1e, and Pinb2v-D1a. Alignment of puroindoline variants or alleles from common wheat and its relatives indicated that all alleles in diploid wheats are attributed to single nucleotide substitution when compared with puroindoline b-2 variants in polyploids. Deduced amino acid sequences showed that all three alleles at Pinb2v-A1 locus and four alleles (Pinb2v-S1a, Pinb2v-S1b, Pinb2v-S1c and Pinb2v-S1e) at the Pinb2v-S1 locus could not be normally translated due to the presence of premature stop codons, whereas Pinb2v-D1a at the Pinb2v-D1 locus and Pinb2v-S1d at the Pinb2v-S1 locus could be normally translated, possibly suggesting that the puroindoline b-2 variant in Ae. tauschii was more highly conserved than those in T. urartu and Ae. speltoides. Meanwhile, puroindoline b-2 variant could be normally translated in all of the durum and common wheat cultivars surveyed. None of the puroindoline b-2 alleles previously identified in durum and common wheat were found in the diploid genome donors examined here, even though a greater diversity of alleles were found in diploid wheat compared to polyploid wheat. These results likely reflect the evolutionary history of tetraploid and hexaploid wheats, although it may be that puroindoline b-2 variant alleles have been selected for stability and functionality in common and durum wheat. This study provides a survey of puroindoline b-2 variants in common wheat and its relatives, and provides useful information for understanding the genetic diversity of puroindoline-like genes and their duplication events in wheat.     

Assessing Genetic Diversity of Chinese Cultivated Barley by STS Markers

To assess the genetic diversity among China’s cultivated barley, sequence tagged site (STS) marker analysis was carried out to characterize 109 morphologically distinctive accessions originating from five Chinese eco-geographical zones. Fourteen polymorphic STS markers representing at least one in each chromosome were chosen for the analysis. The 14 STS markers revealed a total of 47 alleles, with an average of 3.36 alleles per locus (range 2–8). The proportion of polymorphic loci per population averaged 0.84 (range 0.71–1.00); the mean gene diversity averaged 0.39 (range 0.28–0.49). The means of P and H_e were highest in the Yangtze reaches and Southern zone (P = 1.00; H_e = 0.46) and lowest (P = 0.71 He = 0.28) in the Yellow river reaches zone. The STS diversity in different zones is quite different from the morphology diversity. The STS variation was partitioned into 17% among the zone and 83% within the zone. Both cluster and principal coordinate analyses clearly separated the accessions into a dispersed group (mostly two-rowed barley with a lower mean GS value) and a concentrated group (mostly six-rowed barley with a higher mean GS value) according to the spike characteristic with only a few exceptions. The accessions from the Qinghai-Tibet plateau formed a distinctive subgroup and can be distinguished from the concentrated group. The role of Tibet in the origin and evolution of cultivated barley has been discussed.     

Novel Ha Locus,Pina‐D1c/Pinb‐D1h,Affects Soft White Spring Wheat Milling and Baking

Wheat (Triticum aestivum L.) grain hardness is controlled by the Hardness locus on chromosome 5D which consists of the linked genes Puroindoline a and b (Pina and Pinb, respectively). The Ha locus haplotype, Pina‐D1a/Pinb‐D1a, is found in all soft hexaploid wheats. While Pin diversity is low among soft wheats, several novel Ha haplotypes were reported among synthetic hexaploid wheats created using the D genome donor, Aegilops tauschii. One haplotype, Pina‐D1c/Pinb‐D1h, confers a soft phenotype with increased grain hardness over Pina‐D1a/Pinb‐D1a wheats. Here, the Pina‐D1c/Pinb‐D1h haplotype was backcrossed into the soft white spring wheat cultivars ‘Vanna’ and ‘Alpowa’. Then the effect of the two haplotypes on soft wheat milling and baking quality was compared. The effects of the Pina‐D1c/Pinb‐D1h Ha locus haplotype were similar in both the Vanna and Alpowa backgrounds. The Pina‐D1c/Pinb‐D1h lines had significantly more large and fewer small flour particles in both backgrounds and 1.51% higher flour yield in the Alpowa background. The Pina‐D1c/Pinb‐D1h haplotype group was not associated with any consistent differences in solvent retention capacities or sugar snap cookie quality parameters. The results indicate that the Pina‐D1c/Pinb‐D1h haplotype could be used to modify soft wheat milling properties without substantial effects on baking quality.     

Novel and ancient HMW glutenin genes from Aegilops tauschii and their phylogenetic positions

A pair of novel high-molecular-weight glutenin subunits (HMW-GS) 1Dx3.1^t and 1Dy11*^t were revealed and characterized from Aegilops tauschii Coss. subspecies tauschii accession AS60. SDS-PAGE band of 1Dx3.1^t was between those of 1Dx2 and 1Dx3, while 1Dy11*^t was between 1Dy11 and 1Dy12. The lengths of 1Dx3.1 ^t and 1Dy11* ^t were 2,514?bp and 1,968?bp, encoding 836 and 654 amino acid residues, respectively. Their authenticity was confirmed by successful expression of the coding regions in Escherichia coli. Network analysis indicated that 1Dx3.1 ^t together with other five rare alleles only detected in Asia common wheat populations represented the ancestral sequences in Glu-D1 locus. Neighbor-joining tree analysis of previously cloned x-type and y-type alleles in the Glu-D1 locus supported the hypothesis that more than one Ae. tauschii genotypes were involved in the origin of hexaploid wheat and that different Ae. tauschii accessions contributed the D genome to common wheat and Ae. cylindrical Host, respectively. An Ae. tauschii accession with 1Dx3.1 ^t or a closely related allele probably have involved in the origin of common wheat. Since accession AS60 used in this study belonged to typical ssp. tauschii, present results suggested the possibility that ssp. tauschii was involved in the evolution of common wheat.     

Polymorphism of high M r glutenin subunits in wild emmer Triticum turgidum subsp. dicoccoides: chromatographic,electrophoretic separations and PCR analysis of their encoding genes

Triticum turgidum subsp. dicoccoides (Körn. ex Asch. et Graebn.) Thell. (AABB), the immediate progenitor of tetraploid and hexaploid wheats, is a species characterised by a wide range of protein polymorphism and by high protein content. Surveys on polymorphism and genetic control of the high molecular weight glutenin subunits (HMW-GS) present in this species, in two forms x- and y-type at the Glu-A1 and Glu-B1 loci, are still considered useful, both to improve technological properties of breeding varieties and to study the genome evolutionary process in wheats. Comparative Sodium Dodecyl Sulphate Polyacrylamide Gel Electrophoretic and Reversed Phase High Performance Liquid Chromatographic analyses (SDS-PAGE, RP-HPLC) of the HMW-GS present in several accessions of T. turgidum subsp. dicoccoides allowed the detection of new alleles of Glu-A1 and Glu-B1 loci, with x- and y-type glutenin subunits, apparently similar to those present in cultivated wheats in molecular weight, but different in surface hydrophobicity. In addition, changes in the number of x- and y-type subunits at the glutenin loci were also ascertained. The y-type subunits at the Glu-A1 locus, which are never expressed in cultivated bread and durum wheats, and single y-type expressed glutenin subunits at the Glu-B1 locus were also identified in several accessions. DNA extracted from samples, differing in number or type of HMW-GS and corresponding to x- and y-type genes at Glu-1 loci, were amplified using specific primers, two of which were constructed within the transposon-like sequence of Chinese Spring DNA and analysed by polymerase chain reaction. The results showed this insertion in some accessions of T. turgidum subsp. dicoccoides and also the presence of silent Ax, Bx and By type genes. The usefulness for breeding of these comparative analyses carried out on different HMW-GS alleles detected in Triticum turgidum subsp. dicoccoides, is discussed.     

Genetic variation of high-molecular-weight glutenin subunit composition in Asian wheat

To clarify the genetic properties of the HMW glutenin subunit composition of Asian endemic wheats, SDS–PAGE analysis was conducted using 1,139 bread wheat accessions that were originally collected in Asia. The samples were divided into six regional groups, Western Asia, Caucasia, Central Asia, Afghanistan, Southern Asia, and Eastern Asia. The genotype Glu-A1c, Glu-B1b, and Glu-D1a encoding subunits null, 7+8, and 2+12 had an overall frequency of 55.2%. Thus, we conclude that it is the typical genotype of the HMW glutenin subunits that characterize Asian endemic wheat. The frequency of the typical Asian genotype was relatively high in the central belt of Asia (Western Asia, Afghanistan, and Eastern Asia) and low in the marginal regions (Caucasia, Central Asia, and Southern Asia). In Southern Asia, the frequency of Glu-B1i, which encodes subunit 17+18, was the highest at the Glu-B1 locus. In Caucasia and Central Asia, the frequency of Glu-D1d, which encodes subunit 5+10 (which is considered to be the most useful for making bread), was high. The level of genetic variation, as estimated using the frequencies of the various alleles, was relatively low in the central belt of Asia and high in the marginal regions. Among the three Glu-1 loci, the highest number of alleles was detected at the Glu-D1 locus. This result was caused by the presence of rare Asian specific alleles at the Glu-D1 locus, in which a newly found allele, Glu-D1bs, encoding subunit 2.1+12 was included.     

Assimilate and nitrogen remobilization of six-rowed and two-rowed winter barley under drought stress at different nitrogen fertilization

Field experiments were conducted to investigate the effects of irrigation regimes and N levels on assimilate remobilization of two barley cultivars (Yousef_six-rowed and Nimrouz_two-rowed) in 2011 and 2012. There were three levels of water regimes (full irrigation (I₁₀₀), 75% and 50% of I₁₀₀: I₇₅ and I₅₀, respectively) in 2011. Rain-fed treatment (I₀) was included in 2012. Three N levels (0, 60, and 120 kg ha^?1) were used. Grain yield and assimilate remobilization decreased by severe water stress (I₀), however, the reduction of them were intensified by N fertilizer application. The N remobilization was negatively affected by N fertilization and water stress. The two-rowed showed higher N remobilization (10.7%) and contribution of N remobilized to N content of grain (5.8%) than the six-rowed. The two-rowed cultivar showed significantly higher assimilate remobilization and grain yield than the six-rowed under I₅₀ (26.3% and 6.5%, respectively) and I₀ (48.7% and 17.1%, respectively), while the six-rowed had similar or higher performances in terms of these traits under I₁₀₀ and I₇₅. This study showed that optimizing irrigation and N rates (decrease N level with increasing water stress) and selection of the suitable cultivars (Nimrouz_two-rowed) might increase assimilate remobilization and consequently grain yield under drought stress conditions.     

Field-applying an inexpensive, 13C-depleted,labile carbon source to study in situ fate and short-term effects on soils

Background

Labile carbon (C_labile) limits soil microbial growth and is critical for soil functions like nitrogen (N) immobilization. Most experiments evaluating C_labile additions use laboratory incubations. We need to field-apply C_labile to fully understand its fate and effects on soils, especially at depth, but high cost and logistical difficulties hinder this approach.

Aims

Here, we evaluated the impact of adding an in situ pulse of an inexpensive and ¹³C-depleted source of C_labile—crude glycerol carbon (C_glyc), a by-product from biodiesel production—to agricultural soils under typical crop rotations in Iowa, USA.

Methods

We broadcast-applied C_glyc at three rates (0, 216, and 866 kg C ha⁻¹) in autumn after soybean harvest, tracked its fate, and measured its impact on soil C and N dynamics to four depths (0–5, 5–15, 15–30, and 30–45 cm). Nineteen days later, we measured C_glyc in microbial biomass carbon (MBC), salt-extractable organic C, and potentially mineralizable C pools. We paired these measurements with nitrate N (NO₃⁻–N) and potential net N mineralization to examine short-term effects on N cycling.

Results

C_glyc was found to at least 45-cm depth with the majority in MBC (18%–23% of total C_glyc added). The δ¹³C values of the other measured C pools were too variable to accurately track the C_labile fate. NO₃⁻–N was decreased by 13%–57% with the 216 and 866 kg C ha⁻¹ rates, respectively, and was strongly related to greater microbial uptake of C_glyc (i.e., immobilization via microbial biomass). Crude glycerol application had minor effects on soil pH—the greatest rate decreased pH 0.18 units compared to the control.

Conclusions

Overall, glycerol is an inexpensive and effective way to measure in situ, C_labile dynamics with soil depth—analogous to how mobile, dissolved organic C might behave in soils—and can be applied to rapidly immobilize NO₃⁻–N.     

Chloroplast Microsatellites to Investigate the Origin of Grapevine

The origin of the grapevine, Vitis vinifera ssp . sativa L., has been investigated with archaeobotanical–archaeological, cultural and historical data indicating a unique domestication centre located in the Caucasian and Middel-East regions about 6–7000 years ago, but, events leading to the domestication of this species are still an open issue. In this work, eight universal chloroplast microsatellites are used to assess genetic relationships among varieties selected as representatives of four distinct geographical groups from Middle-East to Western European regions. Results show that two out of the eight analysed chloroplast loci are polymorphic within the 142 individuals. Allele variants of the cpSSR loci combine in a total of six different haplotypes. The analysis of haplotypes distribution and haplotype diversity (HD) suggest that only three out of the six haplotypes are represented in the Caucasian and Middle-East samples, with 90% of individuals sharing the same haplotype. Moreover, the presence of all six haplotypes in the European accessions, with a high level of haplotype diversity, suggests varietal influx in these areas. Concerning the Western European varieties, especially in Spanish accessions, half of the individuals share haplotype VI which is completely absent in the Caucasian and Middle-East cultivars. This result opens the discussion about the existence of a unique and common domestication centre, located in the Caucasian and Middle-East area, for all the European cultivars. This work suggests the usefulness of chloroplast genome markers to provide information on haplotype distributions that could help to identify further geographical areas for grapevine varietal evolution.     

Polymorphism of high molecular weight glutenin subunits in three species of Aegilops

A collection of 136 accessions of Aegilops umbellulata (39), Ae. comosa (75) and Ae. markgrafii (22) was analysed for high-molecular-weight (HMW) glutenin subunits composition. The homogeneity of the accessions was studied and 55.1% of the collection was homogeneous for HMW glutenin subunits (29 Ae. umbellulata, 33 Ae. comosa and 14 Ae. markgrafii). The HMW glutenin subunits of Ae. umbellulata are encoded by the Glu-U1 locus; in Ae. comosa results showed that this proteins are encoded at the 1M chromosome, and the locus was named Glu-M1. In Ae. markgrafii it was assumed that HMW glutenin subunits were encoded by an homoeologous locus and it was named Glu-C1. All the accessions of Ae. umbellulata and Ae. markgrafii expressed both, x-type and y-type subunits. Among the Ae. comosa accessions, only one expressed an x-type subunit alone. All the accessions of Ae. umbellulata and some of Ae. comosa had x-type glutenins of higher molecular weights than those commonly present in bread wheat. A total of 8 alleles were detected at the Glu-U1 locus, 11 at the Glu-M1 and 4 at the Glu-C1. The new HMW glutenin variation found in this work suggests their possible utilisation in breeding for wheat quality.     

Genetic diversity and origin of cultivated potatoes based on plastid microsatellite polymorphism

The origin of cultivated potatoes has remaining questions. In this study, 237 accessions of all cultivated species and 155 accessions of wild species closely related to cultivated potatoes, including their putative ancestors, were analyzed using 15 plastid microsatellites (SSRs) to investigate genetic diversity and their relationships with the wild species. We here used polymorphic plastid SSRs we developed from potato plastid genome sequences as well as already known plastid SSR markers. All 15 loci were polymorphic and identified a total of 127 haplotypes. Dramatic decreases in levels of genetic diversity were revealed in landraces in comparison with wild ancestor species. The plastid SSR results showed a decrease in haplotype number and diversity from Peru to both north and south. Phylogenetic analysis revealed two distinct groups. One of them, group A, contained the majority of accessions of cultivated species of the Solanum tuberosum Andigenum group including all accessions of cultivated diploid and triploid cytotypes of this group (S. chaucha, S. phureja, and S. stenotomum by a former taxonomic system) and most of tetraploid accessions of the S. tuberosum Andigenum group (S. tuberosum subsp. andigenum), and the majority of accessions of wild ancestors from the northern members of the S. brevicaule complex. Another group B comprised most of the wild species accessions and almost exclusively hybrid cultivated species which have introgressed plastid genomes from the other wild gene pools. Lack of clustering of traditional cultivated species (as used above) support a revised group classification of S. tuberosum.     

Multiple domestications of the Mesoamerican gene pool of lima bean (Phaseolus lunatus L.): evidence from chloroplast DNA sequences

The Mesoamerican (MA) gene pool of lima bean (Phaseolus lunatus L.) is widely distributed from northern Mexico to northern Argentina in its wild forms and from the southern United States to the east coast of Brazil in its domesticated forms. This broad distribution and lack of wild accessions of the MA gene pool in many areas of its natural distribution has impeded determining its center of domestication and establishing whether it has a single or multiple centers. To answer these questions, we evaluated 262 accessions of P. lunatus using two intergenic spacers of chloroplast DNA: atpB-rbcL and trnL-trnF. The data were analyzed using a maximum likelihood tree (ML), a haplotype network and two estimators of genetic differentiation (N _ST and G _ST). Nucleotide diversity (π) and haplotype (Hd) were quantified to estimate the percentage of reduction in genetic diversity (%r) as a founder effect. The ML tree and haplotype network indicated the existence of three groups (AI, MI and MII), which was supported by the high values of N _ST (0.61–0.80). Values for %r were high (58.67–60.83 %). Existence of Mesoamerican and Andean gene pools was confirmed, with two genetically and geographically distinct groups (MI and MII) within the MA gene pool. We present the first evidence for multiple origins of domestication for the MA gene pool. For MI, we propose western central Mexico as the domestication area and between Guatemala and Costa Rica for MII. We observed a founder effect in the MA gene pool as a result of domestication.     

Evaluation of geographic distribution of high-molecular-weight glutenin subunits (HMW-GS) in wild and cultivated Triticum species

Variation of high-molecular-weight glutenin subunit (HMW-GS) in 632 wild and cultivated Triticum accessions was investigated by sodium dodecyl sulfate polyacrylamide gel electrophoresis. A total of 11 alleles of HMW-GS in diploid species, 22 in tetraploid species, and 15 in hexaploid species were detected. Diploid species on Glu-1A locus and tetraploid species Glu-1B locus showed the highest diversity, respectively. Tetraploid species had the highest level of diversity on three Glu-1 loci, followed by hexaploid and diploid, based on Shannon’s information index, Nei’s genetic diversity, and percentage of polymorphic loci. Molecular variance analysis confirmed main variance of HMW-GS within species, regions, and locations, respectively. Variance among species and regions was enhanced gradually with the increase of ploidy. Significant non-random distributions between the phylogenic trees of HMW-GS and the locations of accessions were tested by GenGIS software, indicated that geographic factors played an important role along the different orientations in the spread of Triticum species. We found one original diversified center in diploid what located around Elazig, Malatya, Gaziantep, Urfa, and Kiziltepe in Turkey, and three diversified centers in tetraploid wheat, including Turkey–Armenia–Georgia–Iran, Portugal–Spain, and Ethiopia, respectively, and two diversified adjacent areas between Turkey and Switzerland and around Turkey, Georgia, and Armenia. The original center of diploid species located in southeast Turkey, where the unexpressed 1Ay subunit was mainly distributed in T. urartu, could be one of the candidate regions of polyploidization of Triticum L. The regional distribution of HMW-GS and species also provided geographic evidences for the existence of founder effect on the spread of Triticum species. The present study suggests that integrating genetic diversity with geographic characterization in Triticum could very useful for collection, conservation, and utilization, as well as for research microevolution and domestication.     

Evolution of tetraploid wheat based on variations in 5′ UTR regions of Ppd-A1: evidence of gene flow between emmer and timopheevi wheat

Previous study showed that tetraploid wheat was divided into two groups (Type AI and Type AII) based on sequences around Ppd-A1 gene (Takenaka and Kawahara in Theor Appl Genet 125(5):999–1014, 2012). That study focused on domesticated emmer wheat and used only 19 wild emmer wheats, so could not be clear the evolutional relationship between Type AI and Type AII. Here, a total of 669 accessions comprising 65 einkorn wheats, 185 wild emmer wheats, 107 hulled emmer wheats, 204 free-threshing (FT) emmer wheats, and 108 timopheevii wheats were studied by PCR assay and DNA sequencing for Type AI/AII. Type AII was an older type than Type AI because all einkorn accessions had Type AII. In wild emmer, Type AI was distributed in the northeast regions of its distribution and Type AII was found to be centered on Israel. A total of 37.4 % of hulled emmer accessions were Type AI, while 92.2 % of FT emmer accessions were Type AI. Differences in the proportion of Type AI/AII in domesticated emmer suggested a strong bottle-neck effect. We also found two MITE-like sequence deletion patterns from a part of Type AII accessions (dic-del and ara-del). Dic-del was found from only Israeli wild emmer accessions and ara-del was found from almost all timopheevii wheat accessions. Only three timopheevii accessions did not have ara-del, and one wild emmer accession and ten hulled emmer accessions had ara-del. These accessions suggested gene flow between emmer and timopheevii wheat.