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Diversification of the North American shrub genus Ceanothus (Rhamnaceae): conflicting phylogenies from nuclear ribosomal DNA and chloroplast DNA.

Ceanothus comprises ∼55 morphologically and ecologically diverse species of woody perennials endemic to North America. Interpretations of the natural history of Ceanothus have served as a general model of evolution for woody perennials with simple entomophilous pollination systems, but these interpretations lacked explicit phylogenetic context. We used cladistic analysis of sequences of the chloroplast-encoded matK and the internal transcribed spacers (ITS) and 5.8S coding region of nuclear ribosomal DNA (nrDNA) to reconstruct the phylogeny of Ceanothus. The nuclear and organellar phylogenies exhibited very low levels of both topological and character congruence. Subgenera Ceanothus and Cerastes are monophyletic sister taxa in both phylogenies, but both data sets suffer from a lack of resolution below the level of subgenus. Lack of taxonomic congruence between the two data sets may be a result of introgression and/or lineage sorting. The ITS tree was accepted as the better estimate of a species phylogeny for Ceanothus, on the assumption that nuclear markers are less prone to introgression. Three of five polytypic species in the ITS data set were paraphyletic, and four of six polytypic species in the matK data set were paraphyletic. This study demonstrates the degree to which matched independent data sets can produce conflicting summaries of evolutionary history.

Journal Article↗

Intersectional gene flow between insular endemics of Ilex (Aquifoliaceae) on the Bonin Islands and the Ryukyu Islands.

Hybridization and introgression play important roles in plant evolution, and their occurrence on the oceanic islands provides good examples of plant speciation and diversification. Restriction fragment length polymorphisms (RFLPs) and trnL (UAA) 3'exon-trnF (GAA) intergenic spacer (IGS) sequences of chloroplast DNA (cpDNA), and the sequences of internal transcribed spacer (ITS) of nuclear ribosomal DNA were examined to investigate the occurrence of gene transfer in Ilex species on the Bonin Islands and the Ryukyu Islands in Japan. A gene phylogeny for the plastid genome is in agreement with the morphologically based taxonomy, whereas the nuclear genome phylogeny clusters putatively unrelated endemics both on the Bonin and the Ryukyu Islands. Intersectional hybridization and nuclear gene flow were independently observed in insular endemics of Ilex on both sets of islands without evidence of plastid introgression. Gene flow observed in these island systems can be explained by ecological features of insular endemics, i.e., limits of distribution range or sympatric distribution in a small land area.

Journal Article↗

A biogeographic genetic approach for testing the role of reinforcement: the case of Drosophila pseudoobscura and D. persimilis.

The role of reinforcement in speciation can be explained by two distinct models. In model I, two diverged populations hybridize and produce fertile hybrids that successfully backcross (hybridization with gene flow). In model II, two populations hybridize but succeeding backcrosses are unproductive (hybridization without gene flow). Using Drosophila persimilis and D. pseudoobscura, we have tested model I by comparing the extent of heterospecific introgression in sympatric versus allopatric populations. We show that certain expectations of this particular model of reinforcement, which is based on hybridization and gene flow between divergent populations after secondary contact, are not realized in these two species. The evidence consists of the similarity of genetic distances as well as proportions of unique/rare alleles between sympatric and allopatric heterospecific populations and a negative correlation between genetic distance and geographical distance between heterospecific populations, which suggests ecological differentiation. This approach in quantifying differential gene flow has important consequences to studies that compare sympatric and allopatric isolation using genetic distance. Following model I, one would expect a pattern of higher prezygotic isolation in sympatric species compared to allopatric species of the same genetic distance simply as a result of an underestimation of genetic distance due to introgression between sympatric populations. We suggest more parsimonious explanations such as reinforcement without genetic exchange (model II) and ecological differentiation, which require high levels of preexisting reproductive isolation between populations.

Animals↗

Routes of origin of two recently evolved hybrid taxa: Senecio vulgaris var. hibernicus and York radiate groundsel (Asteraceae).

The possible pathways of origin of two recently arisen introgressant forms of Senecio vulgaris (i.e., var. hibernicus and York radiate groundsel) were investigated in experimental crosses between tetraploid S. vulgaris var. vulgaris and the normally diploid S. squalidus. Comparison of the morphology of synthesized hybrid progeny with established taxa, by discriminant function analysis, revealed that fertile hybrid offspring similar in morphology to S. vulgaris var. hibernicus and York radiate groundsel could be synthesized: (1) following formation of genomically stable diploid gametes by the triploid hybrid; (2) through the production of unreduced gametes by diploid S. squalidus; and (3) when a tetraploid form of S. squalidus acted as one of the parents. It was evident that hybrid offspring similar in morphology to the two introgressant taxa were more often produced in backcrosses to S. vulgaris than in segregating F2 or F3 generations (53% as opposed to 36%), and that fertile hybrid progeny were formed within two generations. Because hybridization between S. vulgaris and S. squalidus occurs regularly, although at very low frequency, in natural mixed populations in the British Isles, there is the potential for multiple origins to occur in the wild of both S. vulgaris var. hibernicus and York radiate groundsel.

Journal Article↗

Identification and characterization of wheat-wheatgrass translocation lines and localization of barley yellow dwarf virus resistance.

Stable introgression of agronomically important traits into crop plants through wide crossing often requires the generation and identification of translocation lines. However, the low efficiency of identifying lines containing translocations is a significant limitation in utilizing valuable alien chromatin-derived traits. Selection of putative wheatgrass-wheat translocation lines based on segregation ratios of progeny from gamma-irradiated seed using a standard phenotypic analysis resulted in a low 4% success rate of identifying barley yellow dwarf virus (BYDV) resistant and susceptible translocation lines. However, 58% of the susceptible progeny of this irradiated seed contained a Thinopyrum intermedium chromosome-specific repetitive sequence, which indicated that gamma-irradiation-induced translocations occurred at high rate. Restriction fragment length polymorphism (RFLP) analysis of susceptible lines containing alien chromatin, their resistant sister lines and other resistant lines showed that more than one third of the progeny of gamma-irradiated double monosomic seeds contained wheatgrass-wheat translocations. Genomic in situ hybridization (GISH) analysis of selected lines confirmed that these were wheatgrass-wheat translocation lines. This approach of initially identifying BYDV susceptible deletion lines using an alien chromosome-specific repetitive sequence followed by RFLP analysis of their resistant sister lines efficiently identified resistant translocation lines and localized the BYDV resistance to the distal end of the introgressed Th. intermedium chromosome.

Chromatin↗

The effects of hybridization in plants on secondary chemistry: implications for the ecology and evolution of plant-herbivore interactions.

Natural hybridization is a frequent phenomenon in plants. It can lead to the formation of new species, facilitate introgression of plant traits, and affect the interactions between plants and their biotic and abiotic environments. An important consequence of hybridization is the generation of qualitative and quantitative variation in secondary chemistry. Using the literature and my own results, I review the effects of hybridization on plant secondary chemistry, the mechanisms that generate patterns of chemical variation, and the possible consequences of this variation for plants and herbivores. Hybrids are immensely variable. Qualitatively, hybrids may express all of the secondary chemicals of the parental taxa, may fail to express certain parental chemicals, or may express novel chemicals that are absent in each parent. Quantitatively, concentrations of parental chemicals may vary markedly among hybrids. There are five primary factors that contribute to variation: parental taxa, hybrid class (F(1), F(2), etc.), ploidy level, chemical class, and the genetics of expression (dominance, recessive vs. additive inheritance). This variation is likely to affect the process of chemical diversification, the potential for introgression, the likelihood that hybrids will facilitate host shifts by herbivores, and the conditions that might lead to enhanced hybrid susceptibility and lower fitness.

Journal Article↗

[Cytogenetic analysis of hybrids resistant to yellow rust and powdery mildew obtained by crossing common wheat (Triticum aestivum L., AABBDD) with wheat of the Timopheevi group (AtAtGG)].

The karyotypes of 47 hybrid lines obtained from crosses of common wheat Triticum aestivum L. (cv. Rodina and line 353) with Triticum timopheevii Zhuk, (AtAtGG) and related species T. militinae Zhuk. et Migusch. (AtAtGG) and T. kiharae Dorof. et Migusch. (AtAtGGDsqDsq) were analyzed by C-banding. Most lines were resistant to yellow rust and powdery mildew. The introgression of alien genetic material to the common wheat genome was realized via substitutions of complete At-, G-, and D-genome chromosomes, chromosome arms, or their fragments. The pattern of chromosome substitutions in resistant lines differed from that in introgressive hybrids selected for other traits. Substitutions of chromosomes 6G, 2At, 2G, and 5G were revealed in 31, 23, 18, and 13 lines, respectively. Substitutions of chromosomes 4G-, 4At, and 6At were not observed. In 15 lines, a 5BS. 5BL-5GL translocation was identified. High frequency of substitutions of chromosomes 2At, 2G, 5G, and 6G indicate that they may carry the resistance genes and that they are closely related to the respective homoeologous chromosomes of common wheat that determines their high compensation ability.

Fungi↗

Distribution of donor genome composition and its application in marker aided backcrossing.

Conditional distribution was derived for the genome composition given the marker genotype of individuals from a backcross population. Genome size from donor was predicted by the mean. The precision of the prediction, indicated by the correlation with the unknown actual genome size from donor, was formulated by the variance and expressed as a function of the marker density. Results indicated that although high correlation can be obtained with a moderate marker density, say 40 cM per marker, the best individual in a large population has to be identified through a map of high density, say 10 to 20 cM per marker. For the best use of marker information in a marker aided backcrossing program for gene introgression, a stepwise marker genotyping and individual selection were suggested. That is, a preliminary selection can be made among all individuals but only on a few markers for each individual, and a fine selection would then be made among a few individuals but on more markers for each individual. This would be a very efficient method to increase the selection intensity in a large population but with a manageable number of marker genotyping for a gene introgression experiment.

Crosses, Genetic↗

[Evaluation of phylogenetic relationships between five polyploid Aegilops L. species of the U-genome cluster by means of chromosomal analysis].

Four tetraploid (Aegilops ovata, Ae. biuncialis, Ae. columnaris, and Ae. triaristata) and one hexaploid (Ae. recta) species of the U-genome cluster were studied using C-banding technique. All species displayed broad C-banding polymorphism and high frequency of chromosomal rearrangements. Chromosomal rearrangements were represented by paracentric inversions and intragenomic and intergenomic translocations. We found that the processes of intraspecific divergence of Ae. ovata, Ae. biuncialis, and Ae. columnaris were probably associated with introgression of genetic material from other species. The results obtained confirmed that tetraploid species Ae. ovata and Ae. biuncialis occurred as a result of hybridization of a diploid Ae. umbellulata with Ae. comosa and Ae. heldreichii, respectively. The dissimilarity of the C-banding patterns of several chromosomes of these tetraploid species and their ancestral diploid forms indicated that chromosomal aberrations might have taken place during their speciation. Significant differences of karyotype structure, total amount and distribution of C-heterochromatin found between Ae. columnaris and Ae. triaristata, on the one hand, and Ae. ovata and Ae. biuncialis, on the other, evidenced in favor of different origin of these groups of species. In turn, similarity of the C-banding patterns of Ae. columnaris and Ae. triaristata chromosomes suggested that they were derived from a common ancestor. A diploid species Ae. umbellulata was the U-genome donor of Ae. columnaris and Ae. triaristata; however, the donor of the second genome of these species was not determined. We assumed that these tetraploid species occurred as a result of introgressive hybridization. Similarity of the C-banding patterns of chromosomes of Ae. recta and its parental species Ae. triaristata and Ae. uniaristata indicated that the formation of the hexaploid form was not associated with large modifications of the parental genomes.

Chromosome Aberrations↗

[About the possibility of hybridogenesis in the species origin of the midge Chironomus usenicus Loginova et Beljanina (Chironomidae, Diptera)].

Hybridogenesis as a possible way of speciation in Chironomidae was considered with special reference to the species Chironomus usenicus resulting from hybridization between C. plumosus and C. behningi. The three species had 2n = 8 and belonged to the thummi cytocomplex with chromosome arm combinations AB, CD, EF, and G. Arm G had a marker chromosome disk sequence (CDS) and was used to demonstrate the hybrid origin of C. usenicus. Most C. usenicus larvae were heterozygous in CDS of arm G. CDS use G2 proved to be identical to CDS beh G1 of C. behningi and CDS use G1, to CDS plu G1 of C. plumosus. It was assumed that C. usenicus results from hybridization between eurybiont C. plumosus and stenobiont C. behnigni at the boundary of their species areas, in freshwater or brackish water bodies of the southern Saratov oblast and northern Kazakhstan. Morphologically and karyotypically, the hybrid was probably similar to C. plumosus. Crosses with C. plumosus eliminated virtually all C. behningi chromosome sequences from the karyotype of the hybrid. Further chromosome divergence resulting in C. usenicus involved a number of chromosome rearrangements, including duplication of pericentric heterochromatin and other chromosome regions; inversion, which occurred in arm F (regions 13-16) and was fixed in the karyotype; and other paracentric inversions and deletions accumulated in heterozygote in the karyotype pool of the species. Since C. behningi was eliminated from the introgression zone and its species are reduced, the assimilation character was assumed for introgressive hybridization of C. behningi and C. plumosus.

Animals↗

Genetic verification of the role of CCK-AR in pancreatic proliferation and blood glucose and insulin regulation using a congenic rat carrying CCK-AR null allele.

A number of studies have indicated that cholecystokinin type A receptor (CCK-AR) plays a crucial role in postnatal pancreatic proliferation and blood glucose regulation through stimulating insulin secretion. The Otsuka Long-Evans Tokushima Fatty (OLETF) rat has been shown to possess poor pancreatic proliferation (PPP) capability after pancreatectomy (Px). Here we have constructed a congenic strain which introgressed an OLETF-derived 18.5 cM genomic fragment identified in our previous quantitative trait locus (QTL) analysis as a locus responsible for PPP into normoglycemic F344 genetic background The introgressed region includes CCK-AR null mutation. After Px, the congenic rat showed weak pancreatic proliferation equivalent to that of the OLETF rat. Furthermore, post-surgery non-fasting blood glucose levels for the congenic rats are significantly higher in comparison with the F344 rats. At 28 days after Px, the congenic rats also showed lower blood insulin levels than the F344 rats. These results further provide the genetic evidence that 1) CCK-AR is essential for pancreatic regeneration; 2) impaired pancreatic proliferation mediates the development of hyperglycemia.

Alleles↗

Allochronic differentiation among Daphnia species, hybrids and backcrosses: the importance of sexual reproduction for population dynamics and genetic architecture.

Seasonal dynamics of the abundance, sexual reproduction and genetic architecture in a Daphnia hyalina-galeata hybrid complex were studied in the large and deep Lake Constance. We found evidence for the occurrence of first and second order hybridization. Our study revealed strong differences between the parental species not only regarding their seasonal dynamics, genetic architecture and diversity, but also their sexual reproductive behaviour. The overwintering D. hyalina showed low genetic diversity, no genetic differentiation during the season, and reproduced sexually in autumn, whereas D. galeata reached higher levels of genetic diversity, reproduced sexually in early summer, and exhibited changes in genetic structure during the season, but was only present from spring to autumn. However, in both species sexual reproduction was a rare event, and daphnids, including hybrids, reproduced predominantly asexually. This allows long-term persistence of hybrids as well without continuing hybridization events. Within all variables studied, F1 and F2 hybrids showed an intermediate pattern, whereas proposed backcross hybrids were more similar to their respective parentals. These differences in phenotype as well as significant differences in pairwise Fst values between parentals suggest that gene flow seems to be relatively low in the Lake Constance hybrid system. We found evidence for unidirectional introgression by backcrossing from D. galeata to D. hyalina and found a decrease in at least one of the proposed introgressed alleles in the hyalina-backcross while the season progressed. Our findings suggest allochronic differentiation within this hybrid population and different microevolutionary trajectories of the parental species, which will be discussed in the light of the ongoing reoligotrophication process of Lake Constance.

Animals↗

Selection and dispersal in a multispecies oak hybrid zone.

The four western North American red oak species (Quercus wislizeni, Q. parvula, Q. agrifolia, and Q. kelloggii) are known to produce hybrid products in all interspecific combinations. However, it is unknown whether hybrids are transitory resulting from interspecific gene flow or whether they are maintained through extrinsic selection. Here, we examine cryptic hybrid structure in Q. wislizeni through a broad region including contact and isolation from three other western North American red oaks using amplified fragment length polymorphism molecular markers. All four species were simultaneously detected in the genetic background of individuals morphologically assigned to Q. wislizeni, although the contribution of Q. kelloggii was minor. In some cases, introgression was detected well outside the region of sympatry with one of the parental species. Molecular structure at the individual level indicated this was due to long-distance pollen dispersal and not to local extinction of parental species. Species admixture proportions were correlated with climatic variables and greater proportions of Q. agrifolia and Q. parvula were present in the genetic background of Q. wislizeni in sites with cooler and more humid summers, corresponding with habitat preferences of the parental species. Partial Mantel tests indicated that climate was more important than distance from pollen source in this association. Despite high levels of introgression, species integrity was maintained in some populations in close proximity to the other species, providing further support to environmental selection in determining population genetic structure. Thus, the contribution of species mixtures to population genetic structure varies across the landscape according to availability of pollen, but more importantly to varying environmental selection pressures that produce a complex pattern of hybrid and pure gene pools.

California↗

[Polymorphism of Phaseolus vulgaris var. aborigineus (fabaceae). Evidences of natural hibridation].

A polymorphic population of Phaseolus vulgaris var. aborigineus growing at the Northwest of Argentina was studied. In order to know the origin of this polymorphism, some plants belonging to the var. aborigineus, other plants showing floral dimorphism and other individuals with particular characters were collected. Their seeds, obtained after field-work treatments of autogamy and free pollination, were sown in a greenhouse, isolated of the access of pollinators. The growth of each plant was followed until its fructification, and the number of plants that died due to infections was recorded. The number of plants that flowered and fructified was registered in order to study their reproductive success. The floral, fruit and seed qualitative and quantitative characters were documented. With the results obtained, the authors concluded that those individuals that showed floral dimorphism are probably a result of hybridization and introgression between the var. aborigineus and "old cultivars". This hypothesis is supported by the presence of divergent segregation, observed in the offspring of the plants with this segregation. Other crops should allow the genic flow between the parental entities, with the consequence of the establishment of an hybrid population coexistent with their ancestors. Perhaps, as a result of introgression, the stabilized lines exhibit characters different from their parental varieties. The results of autopollination and free pollination in those individuals assigned to var. aborigineus, showed that free pollination brings a great genetic plasticity, because next generations can persist and resist infections. The offspring of the F1 was followed. The plants that belonged to var. aborigineus, product of free pollination, exhibited fast growth and were healthy, while the descendant of the individuals with the floral dimorphism showed characteristics that allowed to conclude the possible existence of degeneration of the hybrid progeny; this characteristics were: curled radicles with cotyledons that never emerge, plantule's apex that soon die with the following development of branches from the cotyledon's axil, and death after some weeks. This degeneration indicates that an unwanted gene flow in the area could lead to a decline in the wild bean population. The vigor, high reproductive success and resistance to illnesses of the individuals corresponding to the var. vulgaris, whose progenitor was treated for free fecundation, and the offspring of the plants with cultivated characteristics, are indicative of the necessity of preserving this germplasm to evaluate its agronomic potential to brief term. The DNA analyses already initiated, will allow the confirmation of the hypotheses outlined in this work.

Argentina↗

[Intraspecific structure and reproductive relationships between Elymus mutabilis and E. transbaicalensis (Poaceae) in southern Siberia from the viewpoint of taxonomical genetics].

The relationships between StH-genomic species Elymus mutabilis (Drob.) Tzvel. and E. transbaicalensis (Nevski) Tzvel. were examined using biosystematic and biochemical (SDS electrophoresis of endosperm proteins) methods. The results showed the following. (1) There is interspecific morphological discreteness that permits assigning most individuals from the wild to a definite species. However, self-fertile intermediate individuals of introgressive origin were found. (2) Both taxa exhibited marked specificity of electrophoretic characteristics of endosperm proteins. (3) The species form two separate recombination gene pools (RGPs). Thus, E. mutabilis and E. transbaicalensis should be regarded as two distinct species. Nevertheless, there is some genetic exchange and unidirectional gene flow between the RGPs, especially in secondary (successive) habitats where mixed populations are observed. Two RGPs form the single introgressive gene pool (IGP), which includes other StH-genomic species of Northern Eurasia.

Electrophoresis, Polyacrylamide Gel↗

Genetic basis of hybrid male sterility among three closely related species of Drosophila.

The genetic basis of hybrid male sterility among three closely related species, Drosophila bipectinata, D. parabipectinata and D. malerkotliana has been investigated by using backcross analysis methods. The role of Y chromosome, major hybrid sterility (MHS) genes (genetic factors) and cytoplasm (non-genetic factor) have been studied in the hybrids of these three species. In the species pair, bipectinata--parabipectinata, Y chromosome introgression of parabipectinata in the genomic background of bipectinata and the reciprocal Y chromosome introgression were unsuccessful as all males in second backcross generation were sterile. Neither MHS genes nor cytoplasm was found important for sterility. This suggests the involvement of X-Y, X-autosomes or polygenic interactions in hybrid male sterility. In bipectinata--malerkotliana and parabipectinata--malerkotliana species pairs, Y chromosome substitution in reciprocal crosses did not affect male fertility. Backcross analyses also show no involvement of MHS genes or cytoplasm in hybrid male sterility in these two species pairs. Therefore, X- autosome interaction or polygenic interaction is supposed to be involved in hybrid male sterility in these two species pairs. These findings also provide evidence that even in closely related species, genetic interactions underlying hybrid male sterility may vary.

Animals↗

[Sequence variation of TYR exon 1 and origin of pigs].

To investigate the origin and genetic diversity of domestic pigs, the porcine TYR exon 1 in 36 individuals from 12 Chinese indigenous breeds, three European breeds, eight Chinese wild boars and two Vietnamese wild boars was sequenced. Sequence analysis revealed six synonymous mutations, and all the sequences could be sorted into 4 haplotypes. Combining with the published sequences, we constructed a reduced median network (RM network), in which TYR*2 was a haplotype dominated by European domestic pigs and wild boars, plus only three chromosomes from Asian pigs. Most Asian domestic pigs and wild boars shared haplotype TYR*1, demonstrating that TYR*1 was an Asian specific haplotype. Meanwhile, some European domestic pigs and wild boars carried the haplotype TYR*1. TYR*3 and TYR*4 were two haplotypes with low frequencies, containing mainly Chinese indigenous pigs and Asian wild boars, plus some European domestic pigs. Independent domestication of pigs from Asia and Europe was supported by the pattern of RM network. The European commercial breeds had been suffered from introgression from Chinese pigs, and a few Chinese indigenous breeds and Japanese wild boars were also suffered from introgression from European pigs as well.

Animals↗

The evolution of California's wild radish has resulted in the extinction of its progenitors.

If two previously isolated taxa mutually assimilate through hybridization and subsequent biparental introgression, and if their introgressed descendants have the same or higher fitness than their parents, then gene flow should result in the local extinction of parental taxa via replacement by hybrid derivatives. These dramatic events may occur rapidly, even in a few generations. Given the speed at which such extinction by hybridization may occur, it may be difficult to identify that the process has occurred. Thus, documented instances of extinction by hybridization are rare, and especially so for cases in which both parents are replaced by the hybrid lineage. Here we report morphological and allozyme evidence for the local extinction of two Raphanus species in California via replacement by their hybrid-derived descendants. The results from a greenhouse experiment demonstrate that California wild radishes have a specific combination of traits from their progenitors, and comparison of our results to that of an earlier report indicate that pure parental types are no longer present in the wild. Our results also show the hybrid-derived lineage has transgressive fruit weight compared to its parents. Allozyme analysis demonstrates that California wild radishes are derived from hybridization between the putative parental species. However, that analysis also demonstrates that California wild radish has now become an evolutionary entity separate from both of its parents. We suggest that the aggressive colonizing behavior of the hybrid-derived lineage probably results from a novel combination of parental traits, rather than genetic variability of the population per se.

Biodiversity↗