Understanding the cytological diploidization mechanism of polyploid wild wheats

The allohexaploid Aegilops species (2n = 6x = 42), Ae. neglecta 6x (UUXtXtNN), Ae. juvenalis (DcDcXcXcUU), and Ae. vavilovii (DcDcXcXcSsSs) regularly form bivalents at metaphase I. However, in Ae. crassa 6x (DcDcXcXcDD) 0.27 quadrivalents per cell were observed probably as a consequence of the partial homology displayed by the D and Dc genomes. Likewise, the synthetic amphiploid Ae. ventricosa-Secale cereale (DDNNRR) is fertile and displays a diploid-like behavior at metaphase I, despite its recent origin. The pattern of synapsis at late zygotene and pachytene in the natural and artificial allohexaploids was analyzed by whole-mount surface-spreading of synaptonemal complexes under an electron microscope. It revealed that chromosomes were mostly associated as bivalents in all cases, the mean of multivalents per nucleus ranging from 0.17 (Ae. neglecta 6x) to 1.03 (Ae. crassa 6x) in the natural species and 1.05 in the Ae. ventricosa-S. cereale amphiploid. It can be concluded that the mechanism controlling bivalent formation in these species and also in the synthetic amphiploid acts mainly at zygotene by restricting synapsis to homologous chromosomes, but also acts at pachytene by preventing chiasma formation in the homoeologous associations. These observations are discussed in relation to the origin and evolution of the mechanism of diploidization in the allopolyploid species of the Poaceae family.     

The evolution of polyploid wheats: identification of the A genome donor species.

Cytogenetic work has shown that the tetraploid wheats, Triticum turgidum and T. timopheevii, and the hexaploid wheat T. aestivum have one pair of A genomes, whereas hexaploid T. zhukovskyi has two. Variation in 16 repeated nucleotide sequences was used to identify sources of the A genomes. The A genomes of T. turgidum, T. timopheevii, and T. aestivum were shown to be contributed by T. urartu. Little divergence in the repeated nucleotide sequences was detected in the A genomes of these species from the genome of T. urartu. In T. zhukovskyi one A genome was contributed by T. urartu and the other was contributed by T. monococcum. It is concluded that T. zhukovskyi originated from hybridization of T. timopheevii with T. monococcum. The repeated nucleotide sequence profiles in the A genomes of T. zhukovskyi showed reduced correspondence with those in the genomes of both ancestral species, T. urartu and T. monococcum. This differentiation is attributed to heterogenetic chromosome pairing and segregation among chromosomes of the two A genomes in T. zhukovskyi.     

Heterochromatin differentiation and phylogenetic relationship of the A genomes in diploid and polyploid wheats

Summary Heterochromatin differentiation, including band size, sites, and Giemsa staining intensity, was analyzed by the HKG (HCl-KOH-Giemsa) banding technique in the A genomes of 21 diploid (Triticum urartu, T. boeoticum and T. monococcum), 13 tetraploid (T. araraticum, T. timopheevi, T. dicoccoides and T. turgidum var. Dicoccon, Polonicum), and 7 cultivars of hexaploid (T. aestivum) wheats from different germplasm collections. Among wild and cultivated diploid taxa, heterochromatin was located mainly at centromeric regions, but the size and staining intensity were distinct and some accessions' genomes had interstitial and telomeric bands. Among wild and cultivated polyploid wheats, heterochromatin exhibited bifurcated differentiation. Heterochromatinization occurred in chromosomes 4A^t and 7A^t and in smaller amounts in 2A^t, 3A^t, 5A^t, and 6A^t within the genomes of the tetraploid Timopheevi group (T. araraticum, and T. timopheevi) and vice versa within those of the Emmer group (T. dicoccoides and T. turgidum). Similar divergence patterns occurred among chromosome 4A^a and 7A^a of cultivars of hexaploid wheat (T. aestivum). These dynamic processes could be related to geographic distribution and to natural and artifical selection. Comparison of the A genomes of diploid wheats with those of polyploid wheats shows that the A genomes in existing diploid wheats could not be the direct donors of those in polyploid wheats, but that the extant taxa of diploids and polyploids probably have a common origin and share a common A-genomelike ancestor.Contribution of the College of Agricultural Sciences, Texas Tech Univ. Journal No. T-4-233.     

The genetic origin of spelt and related wheats

Summary The genetic theory ofMcFadden andSears regarding the origin of spelt and related wheats is defended. This theory postulates thatT. spelta arose fromT. dicoccum (orT. dicoccoides) x Ae. squarrosa in the natural area of the latter. This hybrid became a component of a mixed crop of emmer and einkorn, but only a fractional one, and this medley was brought to southwestern Germany and northern Switzerland, where conditions particularly favorable to spelt caused it to emerge as a major crop.

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Zusammenfassung Die genetische Theorie vonMcFadden undSears über den Ursprung des Spelzes und verwandter Weizen wird unterstützt. Nach dieser Theorie istT. spelta aus der KreuzungT. dicoccum (oderT. dicoccoides) x Ae. squarrosa im natürlichen Areal vonAe. squarrosa entstanden. Diese Hybride wurde ein, allerdings nur in Spuren vorhandener Bestandteil einer aus Emmer und Einkorn bestehenden Mischkultur, und dieses Gemisch gelangte nach Südwestdeutschland und der nördlichen Schweiz, wo für den Spelz besonders günstige Bedingungen zu seiner Entwicklung als selbständige Kulturpflanze führten.

     

The hybrid origin of the polyploid liverwort Pellia borealis

Isozyme markers were used to investigate the origin of the polyploid liverwort, Pellia borealis (gametophytic n=18), which was believed to represent an autopolyploid form of Pellia epiphylla (n=9). Enzyme variation was studied in four taxa: polyploid P. borealis, two recently discovered sibling species of P. epiphylla complex, and the closely related P. neesiana (n=9). Gametophytes of the polyploid showed a complex electrophoretic phenotype for three diagnostic enzymes (DIA1, MPI1 and ACO) in contrast to simple pattern in all haploid taxa. It was postulated that the pattern found in the polyploid represents a fixed heterozygous phenotype resulting from allopolyploidy. Alleles present in the polyploid were found (with only one exception) in the two sibling species of the P. epiphylla complex, suggesting that they are the parents of the allopolyploid. Pellia neesiana was excluded as a donor of either of the genomes. Variation in the polyploid suggests at least three separate origins of P. borealis.     

Studies on maternal inheritance in polyploid wheats with cytoplasmic DNAs as genetic markers

Summary Restriction fragment patterns of DNA fragments obtained after EcoRI cleavage of chloroplastic (cp) and mitochondrial (mt) DNAs isolated from different wheat species were compared. T. aestivum, T. timopheevi, Ae. speltoides, Ae. sharonensis and T. urartu gave species specific mt DNA patterns. Consequently, the cytoplasmic genomes of wheat cannot have originated from contemporary Ae. speltoides, Ae. sharonensis and T. urartu species. It is shown that cp and mt DNAs of Ae. ventricosa, a tetraploid used to transfer eyespot resistance into T. aestivum, contains cp and mt DNAs differing from DNAs isolated from T. aestivum and other wheats. In contrast, the cytoplasmic DNAs of Ae. ventricosa and Ae. squarrosa reveal an important homology, suggesting that Ae. squarrosa was the female parent of Ae. ventricosa. Disomic addition lines (T. aestivum — Ae. ventricosa) in both Ae. ventricosa cytoplasm and T. aestivum cytoplasm contained cytoplasmic DNAs identical to those of the maternal parent. Restriction patterns of the cp and mt DNAs isolated from eight lines of Triticale differing in their cytoplasm have been compared to those of the maternal parent. A strict maternal inheritance has been observed in each case.     

Biochemical data bearing on the relationship between the genome of Triticum urartu and the A and B genomes of the polyploid wheats

To determine whether the Triticum urartu genome is more closely related to the A or B genome of the polyploid wheats, the amino acid sequence of its purothionin was compared to the amino acid sequences of the purothionins in Triticum monococcum, Triticum turgidum, and Triticum aestivum. The residue sequence of the purothionin from T. urartu differs by five and six amino acid substitutions respectively from the alpha 1 and alpha 2 forms coded for by genes in the B and D genomes, and is identical to the beta form specified by a gene in the A genome. Therefore, the T. urartu purothionin is either coded by a gene in the A genome or a chromosome set highly homologous to it. The results demonstrate that at least a portion of the T. urartu and T. monococcum genomes is homologous and probably identical. A variety of other studies have also shown that T. urartu is very closely related to T. monococcum and, in all likelihood, also possesses the A genome. Therefore, it could be argued that either T. urartu and T. monococcum are the same species or that T. urartu rather than T. monococcum is the source of the A genome in T. turgidum and T. aestivum. Except for Johnson's results, our data and that of others suggest a revised origin of polyploid wheats. Specifically, the list of six putative B genome donor species is reduced to five, all members of the Sitopsis section of the genus Aegilops.     

Hydatidiform mole: genetic origin in polyploid conceptuses

Summary The origin of 29 polyploid conceptuses with villous cystic swelling was examined. One tetraploid specimen showed one maternal and three paternal contributions to the genome. Informative cytogenetic markers in 24 triploids were consistent with fertilization by dispermy. Analysis of cytogenetic markers indicated that polyspermy may account for essentially all polyploid conceptions with an excess of paternal to maternal chromosome complements. The origin of the genome was primarily demonstrated by the study of cytogenetic markers, while HLA determination and enzyme analysis were less informative.     

Pairing affinities of the B- and G-genome chromosomes of polyploid wheats with those of Aegilops speltoides.

Chromosome pairing at metaphase I was studied in different interspecific hybrids involving Aegilops speltoides (SS) and polyploid wheats Triticum timopheevii (AtAtGG), T. turgidum (AABB), and T. aestivum (AABBDD) to study the relationships between the S, G, and B genomes. Individual chromosomes and their arms were identified by means of C-banding. Pairing between chromosomes of the G and S genomes in T. timopheevii x Ae. speltoides (AtGS) hybrids reached a frequency much higher than pairing between chromosomes of the B and S genomes in T. turgidum x Ae. speltoides (ABS) hybrids and T. aestivum x Ae. speltoides (ABDS) hybrids, and pairing between B- and G-genome chromosomes in T. turgidum x T. timopheevii (AAtBG) hybrids or T. aestivum x T. timopheevii (AAtBGD) hybrids. These results support a higher degree of closeness of the G and S genomes to each other than to the B genome. Such relationships are consistent with independent origins of tetraploid wheats T. turgidum and T. timopheevii and with a more recent formation of the timopheevi lineage.     

Comparison of the primary structure ofwaxy proteins (granule-bound starch synthase) between polyploid wheats and related diploid species

Thewaxy proteins encoded by the genomes A, B, and D in polyploid wheats and related diploid species were isolated by SDS-PAGE. The N-terminal amino acid sequences of mature proteins and V8 protease-induced fragments were determined. A total of five amino acid substitutions was detected in these sequences, which represent about 10% of the whole sequences of thewaxy proteins. A comparison of these sequences in polyploid wheats with those in related diploid species revealed the following: (i)waxy proteins encoded by the A genome of polyploid wheats were identical to that ofTriticum monococcum, (ii) thewaxy protein encoded by the B genome ofT. turgidum was identical to that ofT. searsii, but differed from those ofT. speltoides andT. longissimum by one amino acid substitution, (iii) thewaxy protein encoded by the B genome ofT. aestivum differed from that encoded by the B genome ofT. turgidum by one amino acid substitution, and (iv) thewaxy protein encoded by the D genome ofT. aestivum was identical to that ofT. tauschii.     

A QTL located on chromosome 4A associated with dormancy in white- and red-grained wheats of diverse origin

Improved resistance to preharvest sprouting in modern bread wheat (Triticum aestivum. L.) can be achieved via the introgression of grain dormancy and would reduce both the incidence and severity of damage due to unfavourable weather at harvest. The dormancy phenotype is strongly influenced by environmental factors making selection difficult and time consuming and this trait an obvious candidate for marker assisted selection. A highly significant Quantitative Trait Locus (QTL) associated with grain dormancy and located on chromosome 4A was identified in three bread wheat genotypes, two white- and one red-grained, of diverse origin. Flanking SSR markers on either side of the putative dormancy gene were identified and validated in an additional population involving one of the dormant genotypes. Genotypes containing the 4A QTL varied in dormancy phenotype from dormant to intermediate dormant. Based on a comparison between dormant red- and white-grained genotypes, together with a white-grained mutant derived from the red-grained genotype, it is concluded that the 4A QTL is a critical component of dormancy; associated with at least an intermediate dormancy on its own and a dormant phenotype when combined with the R gene in the red-grained genotype and as yet unidentified gene(s) in the white-grained genotypes. These additional genes appeared to be different in AUS1408 and SW95-50213.     

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.     

A reconsideration of the domestication geography of tetraploid wheats

The domestication of tetraploid wheats started from their wild progenitor Triticum dicoccoides. In this paper, the geographical distribution of this progenitor is revised to include more sampling locations. The paper is based on a collection of wild and domesticated lines (226 accessions in total) analyzed by AFLP at 169 polymorphic loci. The collection includes the 69 wild lines considered by Mori et al. (2003) in their study on chloroplast DNA haplotypes of T. dicoccoides. The goal of the experiment was to reconsider which location thought to have generated the domesticated germplasm has the highest chance of being the actual site from which wild progenitors were sampled during domestication. Phylogenetic analysis of the nuclear AFLP databases indicates that two different genetic taxa of T. dicoccoides exist, the western one, colonizing Israel, Syria, Lebanon and Jordan, and the central-eastern one, which has been frequently sampled in Turkey and rarely in Iran and Iraq. It is the central-eastern race that played the role of the progenitor of the domesticated germplasm. This is supported by the cumulative results of the AFLP data from the collections of Ozkan et al. (2002) and of Mori et al. (2003), which indicate that the Turkish Karacadag population, intermixed with some Iraq-Iran lines, has a tree topology consistent with that of the progenitor of domesticated genotypes. The Turkish Kartal population belongs genetically to the central-eastern T. dicoccoides race but at the nuclear DNA level is less related to the domesticated gene pool. A general agreement between published work on tetraploid wheat domestication emerges from these results. A disagreement is nevertheless evident at the local geographical scale; the chloroplast DNA data indicate the Kartal mountains while AFLP fingerprinting points to the Karacadag Range as the putative site of tetraploid wheat domestication.     

A molecular reassessment of the Leptomyxid amoebae

Leptomyxid amoebae encompass a diverse assemblage of amoeboid protists that have been implicated as encephalitis-causing agents. This characteristic is attributed to recent studies identifying new members of the Leptomyxidae, in particular, Balamuthia mandrillaris, that cause the disease. Their morphologies range from limax to plasmodial, as well as reticulated and polyaxial. Although systematic studies have identified B. mandrillaris as a new member of the Leptomyxidae, its precise placement within the leptomyxids is uncertain. To further assess the taxonomic placement of Balamuthia among the leptomyxid amoebae and to determine whether the members of the Leptomyxida form a monophyletic assemblage, we have sequenced 16S-like rRNA genes from representatives of three leptomyxid families. Our phylogenetic analyses revealed that current members of the order Leptomyxida do not constitute a monophyletic assemblage. Our analyses clearly show that Gephyramoeba, as well as Balamuthia do not belong in the order Leptomyxida. We highlight where molecular data give differing insights than taxonomic schemes based on traditional characters.     

N-terminal sequence of omega-gliadins from Aegilops longissima. The origin of the genome of polyploid wheat

Using high-performance reversed phase liquid chromatography, the major components of omega-gliadins were isolated from four samples of Aegilops longissima. A high interspecific variability of Ae. longissima with regard to gliadin composition was demonstrated. The N-terminal sequences of omega-gliadins were determined. It was shown that omega-gliadins under study belong to the SRQ type earlier discovered in hexaploid wheat species and in Ae. squarrosa. It is supposed that this type of sequence is specific to the whole Aegilops genus. The N-terminal sequence of omega-gliadin of Ae. longissima was identified and its similarity to the alpha/beta-type sequence found in hexaploid wheat species was revealed. The data obtained are discussed in terms of the origin of polyploid wheat genomes.     

Nuclear and chloroplast DNA reassessment of the origin of Indian potato varieties and its implications for the origin of the early European potato

The modern cultivated potato was first recorded in Europe in 1562, but its area(s) of exportation has long been in dispute. Two competing hypotheses have proposed an Andean area (somewhere from upland Venezuela to northern Argentina) or a lowland south central Chilean area. Potato landraces from these two areas can be distinguished, although sometimes with difficulty, by (1) cytoplasmic sterility factors, (2) morphological traits, (3) daylength adaptation, (4) microsatellite markers, and (5) co-evolved chloroplast (cp) and mitochondria (mt) DNA. The Chilean introduction hypothesis originally was proposed because of similarities among Chilean landraces and modern European cultivars with respect to traits 2 and 3. Alternatively, the Andean introduction hypothesis suggests that (1) traits 2 and 3 of European potato evolved rapidly, in parallel, from Andean landraces to a Chilean type through selection following import to Europe, and (2) the worldwide late blight epidemics beginning in 1845 in the United Kingdom displaced most existing European cultivars and the potato was subsequently improved by importations of Chilean landraces. We reassess these two competing hypotheses with nuclear microsatellite and cpDNA analyses of (1) 32 Indian cultivars, some of which are thought to preserve putatively remnant populations of Andean landraces, (2) 12 Andean landraces, and (3) five Chilean landraces. Our microsatellite results cluster all Indian cultivars, including putatively remnant Andean landrace populations, with the Chilean landraces, and none with the old Andigenum landraces. Some of these Indian landraces, however, lack the cpDNA typical of Chilean landraces and advanced cultivars, indicating they likely are hybrids of Andean landraces with Chilean clones or more advanced cultivars. These results lead us to reexamine the hypothesis that early introductions of potato to Europe were solely from the Andes.     

A reassessment of the telomere hypothesis of senescence

According to the telomere hypothesis of senescence, the telomeric shortening that accompanies the replication of normal somatic cells acts as the mitotic clock that eventually results in their permanent exit from the cell cycle. Although evidence consistent with the telomere hypothesis continues to accumulate, on the basis of recent findings it is suggested that instead of a single clock mechanism there are multiple inducers of senescence.