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Environmental change and rates of evolution: the phylogeographic pattern within the hartebeest complex as related to climatic variation.

Global climate fluctuated considerably throughout the Pliocene-Pleistocene period, influencing the evolutionary history of a wide array of species. Using the phylogeographic patterns within the hartebeest (Alcelaphus buselaphus (Pallas, 1766)) complex, we evaluated the evolutionary consequences of such environmental change for a typical large mammal ranging on the African savannah. Our results, as generated from two mitochondrial DNA markers (the D-loop and cytochrome b), suggest an origin of the hartebeest in eastern Africa from where the species has colonized other parts of the continent. Phylogenetic analyses revealed an early diversification into southern and northern hartebeest lineages, an event that may be related to the formation of the Rift Valley lakes. The northern lineage has further diverged into eastern and western lineages, most probably as a result of the expanding central African rainforest belt and subsequent contraction of savannah habitats during a period of global warming. The diversification events appear to have coincided with major climatic changes and are highly correlated in time. These observations strongly suggest that large-scale climatic fluctuations have been a major determinant for the species' evolutionary history and that hartebeest evolution has mainly taken place in isolated yet environmentally favourable refugia during periods of global warming. Indications of sudden population expansion for two putative ancestral hartebeest populations provide further support for a refugia-based explanation of the diversification events. Reciprocal monophyly between southern and northern lineages may suggest that reproductive barriers exist and that the hartebeest complex comprises two different species.

Africa↗

Efficiency of gamete usage in nature: sperm storage, fertilization and polyspermy.

Gamete production for both males and females can be energetically expensive such that selection should maximize fertilization opportunities while minimizing fertilization costs. In laboratory studies of Drosophila reproduction, however, the failure of eggs to yield adult progeny can be quite high, suggesting that female control over gamete utilization is surprisingly inefficient. We examined gamete utilization in D. pseudoobscura from nature and compared our observations to those for laboratory populations. In natural populations 100% of oviposited eggs effectively produce adult progeny, and fertilization is exclusively monospermic, indicating that in nature, D. pseudoobscura females maintain a very strict control over their reproduction such that gamete usage is extremely efficient. The potential reasons for the inefficient gamete utilization in the laboratory, as well as the potential impact on laboratory studies of sperm competition, sexual conflict, and the evolution of reproductive barriers are discussed. Furthermore, in this sperm-heteromorphic species, our observations show definitively that in nature, as well as in the laboratory, only the long sperm morph participates in fertilization.

Animals↗

Phenotypic differentiation and pre-mating isolation between allopatric populations of Girardinichthys multiradiatus.

Sexual selection may lead to reproductive isolation between populations through divergence in female mate choice, and population differentiation driven by female mate choice is expected to produce pre- but not post-mating isolation. We tested these hypotheses by looking at whether allopatric populations of the Amarillo (Girardinichthys multiradiatus), a sexually dimorphic viviparous fish with effective female choice, (i) have undergone phenotypical differentiation that may be attributed to divergence in female mate choice, and (ii) are already separated by pre- and/or post-mating reproductive barriers. We found substantial divergence in morphological traits which are the target of female mate choice, and in male courtship behaviour. Strong female preferences for homogametic males indicate substantial and symmetric pre-copulatory isolation, but the few successful heterogametic crosses produced in confinement yielded litters of the same size as those produced in homogametic matings, suggesting that post-copulatory isolation between populations is non-existent or weak. It appears that the studied populations have undergone incipient speciation with a pattern that is consistent with speciation driven by sexual selection, yet further work should assess whether divergence in female preferences has promoted male phenotypic differentiation or whether variation in male attributes has driven divergence in female preferences.

Animals↗

Genetic polymorphism and taxonomic infrastructure of the Pleurotus eryngii species-complex as determined by RAPD analysis, isozyme profiles and ecomorphological characters.

The Pleurotus eryngii species-complex includes populations of choice edible mushrooms, growing in the greater Mediterranean area in close association with different genera of plants of the family Apiaceae. Their distinct host-specialization served as the principal criterion for the discrimination of several taxa; however, the genetic relationships among the various P. eryngii ecotypes remain ambiguous. In the present study, 46 Pleurotus strains with a wide range of geographical origins were isolated from Eryngium spp., Ferula communis, Cachrys ferulacea, Thapsia garganica and Elaeoselinum asclepium subsp. asclepium, and were subjected to isozyme and random amplified polymorphic DNA-PCR (RAPD) analysis. The 16 enzyme activities tested were controlled by 28 loci, 11 of which were monomorphic. Host-exclusive zymograms for the Aph (acid phosphatase) and Phe-1 (dopa-phenoloxidase) loci were obtained from Pleurotus strains associated with C. ferulacea. Allele frequencies, genetic diversity and mean diversity were high for isolates from Eryngium spp. and Ferula communis. In RAPD analysis, the use of five primers allowed the production of 45 (out of 48) polymorphic bands, while four molecular markers specific for the identification of Pleurotus strains growing on E. asclepium subsp. asclepium and C. ferulacea were obtained. The Pleurotus strains produced 35 distinct electrophoretic types and 42 RAPD patterns, which independently permitted the separation of the fungal populations into five clusters in accordance with their host-specificity. In addition, the evaluation of the principal ecological and morphological characters provided further evidence for discriminating between P. nebrodensis growing on C. ferulacea and the rest of the host-associated populations. The latter represent taxa at the varietal level: P. eryngii var. eryngii, P. eryngii var. ferulae and P. eryngii var. elaeoselini. The position of taxa of dubious validity, such as P. hadamardii and P. fossulatus, is discussed in relation to the new findings. All Mediterranean Pleurotus populations growing on umbellifers seem to have recently diverged through a sympatric speciation process, that is based on both intrinsic reproductive barriers and extrinsic ecogeographical factors.

Apiaceae↗

Species preferentiality of the pollen tube attractant derived from the synergid cell of Torenia fournieri.

The synergid cell of Torenia fournieri attracts pollen tubes by a diffusible but yet unknown chemical attractant. Here we investigated the species difference of the attractant using five closely related species in two genera, namely T. fournieri, Torenia baillonii, Torenia concolor, Lindernia (Vandellia) crustacea, and Lindernia micrantha. These five species have an exserted embryo sac, and ablation experiments confirmed that their synergid cells attracted the pollen tube. When ovules of T. fournieri and one of the other species were cultivated together with pollen tubes of each species, pollen tubes were significantly more attracted to synergid cells of the corresponding species. The attraction was not affected by the close proximity of embryo sacs of different species. This suggests that the attractant is a species-preferential molecule that is likely synthesized in the synergid cell. The calcium ion, long considered a potential attractant, could not serve as the sole attractant in these species, because elevation of the calcium ion concentration did not affect the observed attraction. In vivo crossing experiments also showed that the attraction of the pollen tube to the embryo sac was impaired when pollen tubes of different species arrived around the embryo sac, suggesting that the species preferentiality of the attractant may serve as a reproductive barrier in the final step of directional control of the pollen tube.

Biological Factors↗

Diversity and natural hybridization in a highly endemic species of Petunia (Solanaceae): a molecular and ecological analysis.

Intrinsic reproductive barriers among the species of Petunia are weak and genetic isolation is obtained mainly by geographical separation and ecological diversification. The Serra do Sudeste region in the extreme south of Brazil is one of the centres of diversity of this genus and is characterized by the presence of species with different pollination syndromes. Petunia exserta is known only from four sandstone towers in a restricted area of this region (about 500 km(2)) and is characterized by its differentiated habitat (shelters in the sandstone towers) and by its floral characteristics adapted to ornithophily. In towers where this species is sympatric with the sphingophilous Petunia axillaris, phylogenetically close to P. exserta, we found plants with intermediate floral morphology, suggesting hybridization between them. To test this hypothesis and to better understand its consequences we analysed the sequences of the plastid trnH-psbA, trnS-trnG and psbB-psbH intergenic spacers in 121 individuals sampled all over the P. exserta distribution. The joint analysis of the three markers revealed 13 haplotypes and the network showed two main genetic clades, which probably represent the original gene pool of the two species in the region. In general, individuals of a given population presented the same haplotype, independently of phenotype, corroborating the hybridization hypothesis. Field observations suggest that hummingbirds are responsible for the interspecific gene flow. Analysis of molecular variance revealed high interpopulational diversity among the towers. The low gene flow between populations is possibly related to the autochoric seed dispersion system.

Ecology↗

Mole rat spalax ehrenbergi: mating behavior and its evolutionary significance.

Mating behavior of the subterranean mole rat, Spalax ehrenbergi, consists of three distinct stages-agonistic, courtship, and copulation. Spalax sexual behavior reflects certain cricetid affinities, some features general in rodents, and others presumably related to its subterranean, territorial life. Within four groups of Spalax ehrenbergi, each with different numbers of chromosomes, recently found in Israel, mating behavior seems to provide partial reproductive barriers. Selective matings between chromosome forms may complement a cytologic isolating mechanism to prevent widespread natural hybridization.

Animals↗

Comparative linkage map of the Solanum lycopersicoides and S. sitiens genomes and their differentiation from tomato.

The wild nightshades Solanum lycopersicoides and Solanum sitiens are closely affiliated with the tomatoes (Lycopersicon spp.). Intergeneric hybridization with cultivated tomato (Lycopersicon esculentum) is impeded by strong reproductive barriers including hybrid sterility and suppressed recombination. Conservation of genome structure between these nightshades and tomato was studied by construction of a genetic map from F2 S. sitiens x S. lycopersicoides and comparison with existing maps of tomato. Owing to self-incompatibility of the F1, two hybrid plants were crossed to obtain a population of 82 F2 individuals. Using 166 previously mapped RFLP markers and 5 restriction enzymes, 101 loci polymorphic in the S. sitiens x S. lycopersicoides population were identified. Analysis of linkage between the markers resulted in a map with 12 linkage groups covering 1192 cM and one unlinked marker. Recombination rates were similar to those observed in tomato; however, significant segregation distortion was observed for markers on 7 out of the 12 chromosomes. All chromosomes were colinear with the tomato map, except for chromosome 10, where a paracentric inversion on the long arm was detected. In this region, S. sitiens and S. lycopersicoides share the same chromosomal configuration previously reported for potato (S. tuberosum) and pepper (Capsicum), suggesting that of tomato is derived. The 10L inversion explains the lack of recombination detected among homeologous chromosomes of intergeneric hybrids in this region. On this basis, we recognize two principle genomes, designated L for the Lycopersicon spp., and S for S. lycopersicoides and S. sitiens, the first examples of structural differentiation between tomato and its cross-compatible wild relatives.

Chromosomes, Plant↗

Meiotic association between the XY chromosomes and unpaired autosomal elements as a cause of human male sterility.

Intimate association between autosomal translocation trivalents and XY bivalents at pachytene was observed in a majority of cells of two men ascertained through primary sterility and found to be heterozygous for a 14;21 Robertsonian translocation. The association, studied by light and electron microscopy of spread first spermatocytes, was between the unpaired short arms of the normal chromosomes of the translocation trivalent and the differential axes of the XY chromosomes. In a minority of cells, this contact was not established, or not maintained, as alternative combinations between the elements available for non-homologous pairing were realized. Following a suggestion of Lifschytz and Lindsley (1972), sterility in these patients was attributed to spermatogenic arrest caused by physical contact of sex chromosomes with autosomal material and consequent interference with the normal metabolism of the sex chromosomes. Autosomal aberrations and polymorphisms, which lead to the presence of unpaired segments at meiosis, may thus play a critical role in a general mechanism of chromosomally-derived male sterility. It is proposed that such a mechanism may also be instrumental in the initiation of reproductive barriers in nature.

Chromosomes, Human, 13-15↗

The chromosomes of Nectomys (Rodentia, Cricetidae) with 2n = 52, 2n = 56, and interspecific hybrids (2n = 54)

Karyologic comparisons of Nectomys with 2n = 52 and 2n = 56, captured in different regions of Brazil, revealed the existence of two presumptive tandem fusions. These differences proved to represent drastic reproductive barriers between these two karyologic groups, based on an analysis of spermatogenesis in captive-bred hybrids (2n = 54). In these hybrid specimens, cytologic and histologic analyses indicated severe meiotic arrest and the presence of two trivalents in the first meiotic division, in which 26 elements were frequently observed. These findings and the sterility of crosses between F1 males and females as well as backcrosses indicated that Nectomys populations with 2n = 52 and 2n = 56 should be considered different species and that the present taxonomic arrangement of this genus must be reviewed.

Animals↗

Chromosomal rearrangements and speciation of sportive lemurs (Lepilemur species).

Theoretical configurations of meiotic chromosomes of potential hybrids between the different Lepilemur species were examined, and the classification of this genus was reviewed in the light of this information. Among the chromosomal rearrangements that occurred during the chromosomal evolution of the sportive lemurs, only those which would generate a pronounced reproductive barrier were considered in relation to the geographic distribution of this genus. The analysis showed that the pattern of geographic distribution is compatible with the inferred chronological occurrence of these chromosomal rearrangements in the phylogenetic tree of the genus Lepilemur.

Animals↗

Linkage maps of the dwarf and Normal lake whitefish (Coregonus clupeaformis) species complex and their hybrids reveal the genetic architecture of population divergence.

Elucidating the genetic architecture of population divergence may reveal the evolution of reproductive barriers and the genomic regions implicated in the process. We assembled genetic linkage maps for the dwarf and Normal lake whitefish species complex and their hybrids. A total of 877 AFLP loci and 30 microsatellites were positioned. The homology of mapped loci between families supported the existence of 34 linkage groups (of 40n expected) exhibiting 83% colinearity among linked loci between these two families. Classes of AFLP markers were not randomly distributed among linkage groups. Both AFLP and microsatellites exhibited deviations from Mendelian expectations, with 30.4% exhibiting significant segregation distortion across 28 linkage groups of the four linkage maps in both families (P < 0.00001). Eight loci distributed over seven homologous linkage groups were significantly distorted in both families and the level of distortion, when comparing homologous loci of the same phase between families, was correlated (Spearman R = 0.378, P = 0.0021). These results suggest that substantial divergence incurred during allopatric glacial separation and subsequent sympatric ecological specialization has resulted in several genomic regions that are no longer complementary between dwarf and Normal populations issued from different evolutionary glacial lineages.

Animals↗

The genetics of inviability and male sterility in hybrids between Anopheles gambiae and An. arabiensis.

Male hybrids between Anopheles gambiae and An. arabiensis suffer from hybrid sterility, and inviability effects are sometimes present as well. We examined the genetic basis of these reproductive barriers between the two species, using 21 microsatellite markers. Generally, recessive inviability effects were found on the X chromosome of gambiae that are incompatible with at least one factor on each arabiensis autosome. Inviability is complete when the gambiae and arabiensis inviability factors are hemi- or homozygous. Using a QTL mapping approach, regions that contribute to male hybrid sterility were also identified. The X chromosome has a disproportionately large effect on male hybrid sterility. Additionally, several moderate-to-large autosomal QTL were found in both species. The effect of these autosomal QTL is contingent upon the presence of an X chromosome from the other species. Substantial regions of the autosomes do not contribute markedly to male hybrid sterility. Finally, no evidence for epistatic interactions between conspecific sterility loci was found.

Animals↗

Genetic mapping of sexual isolation between E and Z pheromone strains of the european corn Borer (Ostrinia nubilalis).

The E and Z pheromone strains of the European corn borer (ECB) provide an exceptional model system for examining the genetic basis of sexual isolation. Differences at two major genes account for variation in female pheromone production and male behavioral response, components of the pheromone communication system known to be important for mate recognition and mate choice. Strains of ECB are morphologically indistinguishable, and surveys of allozyme and DNA sequence variation have revealed significant allele frequency differences at only a single sex-linked locus, Tpi. Here we present a detailed genetic linkage map of ECB using AFLP and microsatellite markers and map the factors responsible for pheromone production (Pher) and male response (Resp). Our map covers 1697 cM and identifies all 31 linkage groups in ECB. Both Resp and Tpi map to the Z (sex) chromosome, but the distance between these markers (>20 cM) argues against the hypothesis that patterns of variation at Tpi are explained by tight linkage to this "speciation gene." However, we show, through analysis of marker density, that Tpi is located in a region of low recombination and suggest that a second Z-linked reproductive barrier could be responsible for the origin and/or persistence of differentiation at Tpi.

Alleles↗

Patterns of allozyme variation in diploid and tetraploid Centaurea jacea at different spatial scales.

The extent and spatial patterns of genetic variation at allozyme markers were investigated within and between diploid and autotetraploid knapweeds (Centaurea jacea L. sensu lato, Asteraceae) at contrasted geographic scales: (1) among populations sampled from a diploid-tetraploid contact zone in the northeastern part of the Belgian Ardennes, and (2) within mixed populations from that zone where diploids and tetraploids coexist. Our data were also compared with a published dataset by Sommer (1990) describing allozyme variation in separate diploid and tetraploid knapweeds populations collected throughout Europe. Genetic diversity was higher in tetraploids. In the Belgian Ardennes and within the mixed populations, both cytotypes had similar levels of spatial genetic structure, they were genetically differentiated, and their distributions of allele frequencies were not spatially correlated. In contrast, at the European scale, diploids and tetraploids did not show differentiated gene pools and presented a strong correlation between their patterns of spatial genetic variation. Numerical simulations showed that the striking difference in patterns observed at small and large geographic scales could be accounted for by a combination of (1) isolation by distance within cytotypes; and (2) partial reproductive barriers between cytotypes and/or recurrent formation of tetraploids. We suggest that this may explain the difficulty of the taxonomic treatment of knapweeds and of polyploid complexes in general.

Asteraceae↗

Evidence through crossmating experiments of a species complex in Anopheles pseudopunctipennis sensu lato: a primary malaria vector of the American continent.

Crossmating experiments were conducted to determine if postmating reproductive barriers are involved in the maintenance of genetic divergence among populations of Anopheles pseudopunctipennis sensu lato, a primary malaria vector of the American continent. Reciprocal crosses were conducted between colony and wild strains from Mexico, Bolivia, and Peru. Hybridization experiments revealed unidirectional male/female hybrid sterility in crosses between Mexican females and South American males. The data presented provide the first evidence that genetic differences exist among geographic strains of An. pseudopunctipennis in neotropical America. There is a consistent pattern suggesting the presence of at least two allopatric sibling species. One species occurs in central Mexico, the other in the South American Andean Cordillera.

Animals↗

Reproductive biology and the process of domestication of the columnar cactus Stenocereus Stellatus in Central Mexico.

Pollination biology, breeding system, and floral phenology of the columnar cactus Stenocereus stellatus were studied in wild, wild managed in situ and cultivated populations of central Mexico, in order to examine whether these aspects have been modified under domestication and whether they determine reproductive barriers between wild and manipulated individuals. Individuals of both wild and manipulated populations are self-incompatible, indicating that artificial selection has not modified the breeding system. Their pollination biology is also similar. Anthesis is mainly nocturnal, with a peak of nectar production between 0200 and 0400 when the stigma presents maximum turgidity. Nocturnal visitors are the effective pollinators. Nearly 75% of flowers exposed for nocturnal pollination set fruit, while none of the flowers exposed for diurnal pollination produced fruits. The bats Leptonycteris curasoae, L. nivalis, and Choeronycteris mexicana (Glossophaginae) are the most likely pollinators, and their time of foraging is synchronized with the time of nectar production and stigma receptivity in S. stellatus. Bats potentially move pollen over a considerable distance, so there is apparently no spatial isolation to prevent pollen exchange between wild and cultivated populations. Phenological studies showed that there are also no apparent temporal barriers. However, manual cross pollination failed between some domesticated and wild phenotypes, suggesting that gene flow between wild and cultivated populations might be limited by pollen incompatibility.

Journal Article↗

Hybridization, introgression, and linkage evolution.

Genetic mapping methods provide a unique opportunity to study the interactions of differentiated genes and genomes in a hybrid genetic background. After a brief discussion of theoretical and analytical concerns, we review the application of these methods to a wide range of evolutionary issues. Map-based studies of experimental hybrids indicate that most postzygotic reproductive barriers in plants are polygenic and that the expression of extreme or novel traits in segregating hybrids (transgressive segregation) results from the complementary action of divergent parental alleles. However, genetic studies of hybrid vigor do not concur in their interpretations of the relative roles of dominance, overdominance, and epistasis. Map-based studies of natural hybrids are much rarer, but the few existing studies confirm the polygenic basis of postzygotic barriers and demonstrate the utility of genetic linkage for detecting cryptic introgression. In addition, studies of experimental and natural hybrid lineages provide compelling evidence that homoploid hybrid speciation has occurred in nature, and that it represents a rapid and repeatable mode of speciation. Data further indicate that this mode is facilitated by strong fertility selection and high chromosomal mutation rates. We recommend that future studies of hybrid genomes focus on natural hybrids, not only because of the paucity of data in this area, but also because of the availability of highly recombinant hybrid genotypes in hybrid zones. Of particular value will be studies of long-lived or difficult-to-propagate organisms, which previously have not been amenable to genetic study.

Crosses, Genetic↗