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Genetic differentiation and reproductive isolation in a Cyprinodon fish species flock from Laguna Chichancanab, Mexico.

The Cyprinodon species flock from Laguna Chichancanab, aged 8000 years, provides another potential case of sympatric speciation. The flock consists of seven morphologically distinct species, each within partially different trophic niches, and a group of specimens which cannot unequivocally be assigned to one of these species. Genetic analyses, based on mtDNA and five microsatellite loci, revealed significant genetic differentiation of one species, C. maya, from other members of the species flock, providing strong evidence for reproductive isolation. For the remaining members of the flock significant genetic structuring was detected, with some evidence of gene flow with the most abundant species C. beltrani. These analyses suggest that speciation proceeds with ongoing hybridisation, and further suggest that the morphologically unidentifiable specimens found in the lake are probably hybrids. I propose that in the Cyprinodon species flock besides disruptive selection sexual selection plays an important part in achieving and maintaining reproductive isolation.

Animals↗

Allopolyploidy in bryophytes: Multiple origins of Plagiomnium medium.

Bryophytes are thought to be unique among land plants in lacking the important evolutionary process of allopolyploidy, which involves interspecific hybridization and chromosome doubling. Electrophoretic data show, however, that the polyploid moss Plagiomnium medium is an allopolyploid derivative of Plagiomnium ellipticum and Plagiomnium insigne, that P. medium has originated more than once from these progenitors, and that cross-fertilization results in interlocus genetic recombination. Evidence from restriction fragment length polymorphisms in chloroplast DNA implicates P. insigne as the female parent in interspecific hybridizations with P. ellipticum. Contrary to prevailing views, it appears that those evolutionary processes responsible for genetic differentiation and speciation in other land plants occur in the bryophytes as well.

Journal Article↗

Spore fitness components do not differ between diploid and allotetraploid species of Dryopteris (Dryopteridaceae).

BACKGROUND AND AIMS: Although allopolyploidy is a prevalent speciation mechanism in plants, its adaptive consequences are poorly understood. In addition, the effects of allopolyploidy per se (i.e. hybridization and chromosome doubling) can be confounded with those of subsequent evolutionary divergence between allopolyploids and related diploids. This report assesses whether fern species with the same ploidy level or the same altitudinal distribution have similar germination responses to temperature. The effects of polyploidy on spore abortion and spore size are also investigated, since both traits may have adaptive consequences. METHODS: Three allotetraploid (Dryopteris corleyi, D. filix-mas and D. guanchica) and three related diploid taxa (D. aemula, D. affinis ssp. affinis and D. oreades) were studied. Spores were collected from 24 populations in northern Spain. Four spore traits were determined: abortion percentage, size, germination time and germination percentage. Six incubation temperatures were tested: 8, 15, 20, 25 and 32 degrees C, and alternating 8/15 degrees C. KEY RESULTS: Allotetraploids had bigger spores than diploid progenitors, whereas spore abortion percentages were generally similar. Germination times decreased with increasing temperatures in a wide range of temperatures (8-25 degrees C), although final germination percentages were similar among species irrespective of their ploidy level. Only at low temperature (8 degrees C) did two allotetraploid species reach higher germination percentages than diploid parents. Allotetraploids showed faster germination rates, which would probably give them a competitive advantage over diploid parents. Germination behaviour was not correlated with altitudinal distribution of species. CONCLUSIONS: The results of this study suggest that (i) relative fitness of allopolyploids at sporogenesis does not differ from that of diploid parents and (ii) neither does allopolyploidization involve a change in the success of spore germination.

Dryopteris↗

Nest architecture and genetic differentiation in a species complex of Australian stingless bees.

We investigated the taxonomic significance of nest shape and its putative role in speciation in Trigona (Heterotrigona) carbonaria and T. (H.) hockingsi, two sibling species of stingless bee species from eastern Australia. These species are primarily distinguished by their nest architecture, as in all other respects they are nearly identical. We genotyped 130 colonies from six locations in Queensland at 13 microsatellite loci together with 106 additional colonies from six other Indo-Pacific Trigona species. Whether they were present in allopatry or in sympatry, colonies that displayed the T. carbonaria or the T. hockingsi nest architecture could be unambiguously differentiated at the genetic level. However, T. hockingsi colonies were classifiable into two highly differentiated paraphyletic and geographically separate populations, one in northern and one in southern Queensland. These two populations probably belong to two distinct species, T. hockingsi and T. davenporti nov. sp. Our results suggest that nest architecture characters are relevant but not sufficient criteria to identify species in this group. Consequently, modifications of nest architecture are probably not of prime importance in the speciation process of Australian stingless bees, although nest architecture differences probably result from relatively simple mechanisms. The rare interspecific hybrid colonies detected did not display a nest with an intermediate form between T. hockingsi and T. carbonaria.

Animals↗

Refinement of a chimpanzee pericentric inversion breakpoint to a segmental duplication cluster.

BACKGROUND: Pericentric inversions are the most common euchromatic chromosomal differences among humans and the great apes. The human and chimpanzee karyotype differs by nine such events, in addition to several constitutive heterochromatic increases and one chromosomal fusion event. Reproductive isolation and subsequent speciation are thought to be the potential result of pericentric inversions, as reproductive boundaries form as a result of hybrid sterility. RESULTS: Here we employed a comparative fluorescence in situ hybridization approach, using probes selected from a combination of physical mapping, genomic sequence, and segmental duplication analyses to narrow the breakpoint interval of a pericentric inversion in chimpanzee involving the orthologous human 15q11-q13 region. We have refined the inversion breakpoint of this chimpanzee-specific rearrangement to a 600 kilobase (kb) interval of the human genome consisting of entirely duplicated material. Detailed analysis of the underlying sequence indicated that this region comprises multiple segmental duplications, including a previously characterized duplication of the alpha7 neuronal nicotinic acetylcholine receptor subunit gene (CHRNA7) in 15q13.3 and several Golgin-linked-to-PML, or LCR15, duplications. CONCLUSIONS: We conclude that, on the basis of experimental data excluding the CHRNA7 duplicon as the site of inversion, and sequence analysis of regional duplications, the most likely rearrangement site is within a GLP/LCR15 duplicon. This study further exemplifies the genomic plasticity due to the presence of segmental duplications and highlights their importance for a complete understanding of genome evolution.

Animals↗

The genetic basis of prezygotic reproductive isolation between Drosophila santomea and D. yakuba due to mating preference.

Sexual isolating mechanisms that act before fertilization are often considered the most important genetic barriers leading to speciation in animals. While progress has been made toward understanding the genetic basis of the postzygotic isolating mechanisms of hybrid sterility and inviability, little is known about the genetic basis of prezygotic sexual isolation. Here, we map quantitative trait loci (QTL) contributing to prezygotic reproductive isolation between the sibling species Drosophila santomea and D. yakuba. We mapped at least three QTL affecting discrimination of D. santomea females against D. yakuba males: one X-linked and one autosomal QTL affected the likelihood of copulation, and a second X chromosome QTL affected copulation latency. Three autosomal QTL also affected mating success of D. yakuba males with D. santomea. No epistasis was detected between QTL affecting sexual isolation. The QTL do not overlap between males and females and are not disproportionately concentrated on the X chromosome. There was some overlap in map locations of QTL affecting sexual isolation between D. santomea and D. yakuba with QTL affecting sexual isolation between D. simulans and D. mauritiana and with QTL affecting differences in pigmentation between D. santomea and D. yakuba. Future high-resolution mapping and, ultimately, positional cloning, will reveal whether these traits do indeed have a common genetic basis.

Animals↗

Localization of the U2 linkage group of horses to ECA 3 using chromosome painting.

The U2 linkage group of horses includes the genes albumin (ALB), vitamin D binding protein (GC), mitochondrial glutamate oxaloacetate transaminase 2 (GOT2), and haptoglobin (HP) which are found on two human chromosomes, namely, 4 (HSA 4) and 16 (HSA 16). Likewise these genes are also found on two different chromosomes in mice, rats, and cattle. Chromosome painting demonstrated that only horse chromosome 3 (ECA 3) hybridized with whole chromosome paints for both HSA 4 and HSA 16. This indicated that the equine U2 linkage group occurs on ECA 3, spanning the centromere. This technique will be useful to study the chromosome rearrangements associated with speciation of the genus Equus.

Albumins↗

Chromosomal inversions and the reproductive isolation of species.

Recent genetic studies have suggested that many genes contribute to differences between closely related species that prevent gene exchange, particularly hybrid male sterility and female species preferences. We have examined the genetic basis of hybrid sterility and female species preferences in Drosophila pseudoobscura and Drosophila persimilis, two occasionally hybridizing North American species. Contrary to findings in other species groups, very few regions of the genome were associated with these characters, and these regions are associated also with fixed arrangement differences (inversions) between these species. From our results, we propose a preliminary genic model whereby inversions may contribute to the speciation process, thereby explaining the abundance of arrangement differences between closely related species that co-occur geographically. We suggest that inversions create linkage groups that cause sterility to persist between hybridizing taxa. The maintenance of this sterility allows the species to persist in the face of gene flow longer than without such inversions, and natural selection will have a greater opportunity to decrease the frequency of interspecies matings.

Animals↗

On the relative roles of faster-X evolution and dominance in the establishment of intrinsic postzygotic isolating barriers.

The modern theory of speciation assigns a prominent role to the recessivity of genetic incompatibilities in the two rules of speciation, namely Haldane's rule and the 'large X effect', and considers that the contribution of faster evolution of the X versus the autosomes to those patterns is generally of relatively minor importance. By extending Turelli and Orr's previous analysis of the model of two-locus Dobzhansky-Muller incompatibilities, I first show that when the X and the autosomes evolve at the same rate, the two dominance parameters involved in that model are not equally important for the declaration of a large X effect, but that the degree of recessivity of homozygous-homozygous incompatibilities is the major determinant for such a declaration. When the X evolves faster than the autosomes, the model obviously predicts that the importance of both dominance parameters will progressively vanish. It is then of importance to obtain estimates of the relative evolutionary rate of X-linked incompatibility loci. Several different procedures to obtain such estimates from the perspective of the large X effect are suggested. The application of the appropriate test to the only suitable data from Drosophila hybridizations so far available leads to the conclusion that the X actually evolves at least 2.5 times faster than the autosomes, as far as hybrid male sterility determinants are concerned, thus making dominance considerations absolutely irrelevant. Notwithstanding the necessity of further tests, the relative roles currently assigned to faster-X evolution and dominance in the theory of speciation should be revised, giving due prominence to faster-X evolution, at least for hybrid male sterility in the genus Drosophila.

Animals↗

Genetic basis of X-Y chromosome dissociation and male sterility in interspecific hybrids.

A high frequency of X-Y chromosome dissociation (95%) was found at first meiotic metaphase (MI) in spermatocytes of interspecific hybrids between laboratory mice, C57BL/6J (BL/6) and Mus spretus, compared with an X-Y dissociation frequency of only 3-4% in parental mice. The X-Y dissociation in F1 hybrids occurred before diakinesis rather than as a precocious dissociation at MI. The high X-Y dissociation was accompanied by spermatogenic breakdown after MI, resulting in male sterility. All F1 males were sterile and approximately half of the backcross males from fertile F1 females crossed with either BL/6 or M. spretus males were sterile. Male sterility was highly correlated with X-Y dissociation in both backcrosses. All of the mice with high X-Y dissociation were sterile and all of the males with low X-Y dissociation were fertile or subfertile. This correlation suggested that genetic divergence of the X-Y pairing region could contribute to the male sterile phenotype such that the BL/6 X chromosome would not pair with the M. spretus Y chromosome. The segregation of species-type alleles of amelogenin (Amelb and Amels), a distal X chromosome locus adjacent to the X-Y pairing region, was followed in backcross males that were analyzed for X-Y dissociation and sterility (we have used Amel as the designation for the mouse amelogenin locus; the current designation for this locus is Amg). A 95% concordance between Amelb with fertility and Amels with sterility was observed in backcrosses with BL/6, whereas the converse was observed in the backcross to M. spretus. These results imply that X-Y pairing plays an important role in male fertility and suggest that genetic divergence in X-Y pairing region between Mus species can contribute to the reproductive barriers between species and the process of speciation.

Animals↗

Amplification of telomeric DNA and the extent of karyotypic evolution.

The distribution of telomeric DNA in the genomes of the antelope ground squirrel, Ammospermophilus harrisii (family Sciuridae; 2n = 32) and the African black-footed cat, Felis nigripes (family Felidae; 2n = 38) were compared by fluorescence in situ hybridization (FISH) technique. These two mammalian species have the highest and the lowest amount of C-banded regions, respectively. FISH preparations with the human telomeric DNA probe showed that all C-banded segments in the A. harrisii chromosomes, except a few intercalary segments, were hybridizing with this DNA. F. nigripes showed hybridization only on the termini of each chromosome, and the C-banded regions did not hybridize with telomeric DNA on FISH analysis. The C-banded chromosomal arms in another rodent species, Peromyscus eremicus (family Cricetidae; 2n = 48), when hybridized with human telomeric DNA showed signals only in the termini of chromosomes but not in the heterochromatic arms. These observations indicate that not all C-banded regions in rodent species are telomeric DNA. The amplification of telomeric DNA in relation to speciation is discussed.

Animals↗

Study of the evolutionary relationships among Limonium species (Plumbaginaceae) using nuclear and cytoplasmic molecular markers.

The genus Limonium, due to the patchiness of the natural habitats of its species as well as the high frequency of hybridization and polyploidy and the possibility of reproduction by apomixis, provides an example of all the principal mechanisms of rapid speciation of plants. As an initial study of evolution in this genus, we have analyzed intra- and interspecific variability in 17 species from section Limonium, the largest in the genus, based on RFLPs of cpDNA and nuclear rDNA ITS sequences. In the cpDNA analysis, 21 restriction enzymes were used, resulting in 779 fragments, 490 of which were variable and 339 parsimony informative. L. furfuraceum exhibited two relatively divergent cpDNA haplotypes. The relationships found among the species based on cpDNA restriction fragments were coincident using different methods of phylogenetic analysis. Due to the presumed reticulate evolution in the genus Limonium, the comparison of these results with data from the nuclear DNA was necessary; ITS sequences were analyzed. The final alignment contained 488 characters, of which 198 were variable and 156 parsimony informative. Two relatively divergent ITS types were present at the intraindividual level in L. delicatulum, a triploid species. Each type was related to ITS from different groups of diploid Limonium species, one with a base haploid chromosome number n = 8 (represented by L. cossonianum) and the other with n = 9 (represented by L. minutum). The different phylogenetic inference methods used for the analysis of ITS sequences rendered very similar topologies. In general, the relationships among the species studied were coincident with those obtained with the chloroplast genome. Both nuclear and cytoplasmic markers support the polyphyly of section Limonium, with at least two species, L. narbonense and L. vulgare, clearly divergent from the rest. Moreover, the remaining subsections into which section Limonium is currently divided seem to be artificial.

Base Sequence↗

Lack of prezygotic isolation by assortative mating between the two cryptic species of the polytypic Schistosoma intercalatum taxon.

Ten mice were individually exposed to 30 male and 30 female cercariae of Schistosoma intercalatum from Cameroon and to 30 male and 30 female cercariae of S. intercalatum from Zaire. Ten weeks later, the nine surviving mice were killed, and a total of 263 adult schistosomes (228 pairs, 7 unpaired males, 28 unpaired females) were perfused. All worms were identified utilising the random amplified polymorphic DNA technique using primer OPG06 which enables the differentiation of the Cameroon and Zaire genotypes. The data demonstrate that there were no significant differences between the number of homospecific pairs observed compared with the expected number under the null hypothesis. Matings occurred in a random manner indicating that mate choice did not exist for the two genotypes of S. intercalatum. Hence there is no prezygotic isolation mechanism which would prevent the interaction of the two genotypes if they were sympatric. However, as they are allopatric, the possibility of the production of unfit hybrids does not arise. Consequently, the reinforcement of a prezygotic isolating mechanism does not play a significant role in the speciation process.

Animals↗

Female meiosis drives karyotypic evolution in mammals.

Speciation is often accompanied by changes in chromosomal number or form even though such changes significantly reduce the fertility of hybrid intermediates. We have addressed this evolutionary paradox by expanding the principle that nonrandom segregation of chromosomes takes place whenever human or mouse females are heterozygous carriers of Robertsonian translocations, a common form of chromosome rearrangement in mammals. Our analysis of 1170 mammalian karyotypes provides strong evidence that karyotypic evolution is driven by nonrandom segregation during female meiosis. The pertinent variable in this form of meiotic drive is the presence of differing numbers of centromeres on paired homologous chromosomes. This situation is encountered in all heterozygous carriers of Robertsonian translocations. Whenever paired chromosomes have different numbers of centromeres, the inherent asymmetry of female meiosis and the polarity of the meiotic spindle dictate that the partner with the greater number of centromeres will attach preferentially to the pole that is most efficient at capturing centromeres. This mechanism explains how chromosomal variants become fixed in populations, as well as why closely related species often appear to have evolved by directional adjustment of the karyotype toward or away from a particular chromosome form. If differences in the ability of particular DNA sequences or chromosomal regions to function as centromeres are also considered, nonrandom segregation is likely to affect karyotype evolution across a very broad phylogenetic range.

Animals↗

Tracing the evolution of brain and behavior using two related species of whiptail lizards: Cnemidophorus uniparens and Cnemidophorus inornatus.

Cnemidophorus whiptail lizards offer a unique opportunity to study behavioral and neural evolution because unlike most genera, ancestral and descendant species are still extant, and comparisons between species provide a window into correlated changes in biological organization through speciation. This review focuses on the all-female or parthenogenetic species Cnemidophorus uniparens (descendant species), which evolved through several hybridization events involving the sexually reproducing species Cnemidophorus inornatus (ancestral species). Data compiled over more than 2 decades include behavioral, endocrine, and neural differences between these two related species of whiptail lizards. For example, unlike females of the ancestral species, individuals of the descendant species display male-like mounting behavior (pseudocopulatory behavior) after ovulation. Pseudocopulatory behavior in the parthenogen is triggered by the progesterone surge after ovulation, and the behavioral capacity to respond to progesterone appears to be an ancestral trait that was inherited from C. inornatus males through the hybridization events. Interestingly, the regulation of sex steroid hormone receptor mRNA in brain areas critical for the expression of sociosexual behaviors differs between females of the two species and suggests that evolutionary changes in the regulation of gene expression could be a proximate mechanism that underlies the evolution of a novel social behavior in the parthenogen. Finally, because the sexual species is diploid, whereas the parthenogen is triploid, differences between the species could directly assess the effect of ploidy. The behavioral and neuroendocrinological data are pertinent for considering this possibility.

Animal Husbandry↗