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Origin of reproductive isolation in the absence of apparent genic differentiation in a geographic isolate of Drosophila pseudoobscura.

F(1) males obtained from the cross of D. pseudoobscura females from Bogotá (Colombia) x males of this species from mainland, i.e. populations from various locations in the United States and from Guatemala, are sterile. This sterility is due to genes located on the X chromosome and the autosomes; the Y chromosome is not involved. The percentage of sterile males in backcrosses can be explained by assuming an interaction between two loci on the Bogotá X chromosome and probably two loci, one each on two of the mainland autosomes. The role of founder events, inbreeding and geographic isolation in the development of reproductive isolation and the magnitude of gene differences responsible for the origin of reproductive isolation is discussed. It is concluded that founder events, inbreeding and geographic isolation play a major role in the development of reproductive isolation and that major adaptive incorporation of new alleles at a large number of structural loci is not necessary for the origin of reproductive isolation.

Animals

Microorganism mediated reproductive isolation in flour beetles (genus Tribolium).

Reproductive isolation is induced by microorganisms in diverse geographic strains of the flour beetle Tribolium confusum (Coleoptera:Tenebrionidae). The incompatibility between populations is due to nongenetic cytoplasmically inherited factors. Males of infected strains produce no progeny when crossed with females of noninfected strains; however, they produce "normal" numbers of progeny when crossed with infected females. Males from noninfected strains show no reproductive isolation. Infected strains of T. confusum can be cured when tetracycline or other antibiotics are added to the flour medium. "Cured" strains become partially reproductively isolated from all noncured strains including the source strain

Animals

Evidence of parental care as a newly identified reproductive isolating barrier.

Variation in behavior can contribute to reproductive isolation by preventing gene flow among populations. Here, we tested the novel hypothesis that parental care, when dysregulated, can function as a reproductive isolating mechanism in three-spined stickleback fish (Gasterosteus aculeatus). In the typical "common" stickleback ecotype, males provide care to their offspring by fanning with their pectoral fins and defending their nest. In contrast, a divergent "white" stickleback ecotype has evolutionarily lost care and disperses embryos into the surrounding environment. We examined how paternal care from common, white, and F1 hybrid fathers influenced offspring survival. We detected no intrinsic incompatibilities in embryos, but F1 hybrid fathers exhibited dysregulated parental care that coincided with decreased survival of parented offspring. The increased offspring mortality may be explained by further dysregulation of feeding and parenting circuitry, as F2 hybrids exhibited significantly higher rates of post-fertilization filial cannibalism than common or white fathers. Additionally, despite strong divergence in nesting-building and courtship behavior, F1 hybrids achieved mating success at a similar rate as male commons and whites, suggesting that prezygotic barriers against hybrids may be weak. The observed postzygotic isolation, and potentially weak prezygotic isolation in lab-based studies, suggest that hybridization is likely occurring at low rates in the wild. Population genetic analysis supported this, as low proportions of putative hybrids were detected in sympatric sites. Together, these results may help explain why genetic divergence between these phenotypically distinct ecotypes is low and provide evidence that dysregulated parental behavior can act as a newly discovered postzygotic reproductive isolating barrier.

Reproductive isolation

Reproductive Isolation due to Divergent Ecological Selection Is Accompanied by Vast Genomic Instability in Experimentally Evolved Yeast Populations.

Populations evolving independently in divergent environments accumulate genetic differences and potentially evolve reproductive isolation as a by-product of divergence. The speed and mechanisms underlying this process are difficult to investigate because we rarely get the opportunity to witness them in natural settings, and histories of selection and gene flow between populations are often unknown. Here, we experimentally evolved yeast for 1000 generations of evolution in both divergent and parallel environments. At regular time points during experimental evolution, we made crosses between parallel- and divergent-evolving populations to measure postzygotic reproductive isolation (gamete viability). We used whole genome population sequencing to determine the mutational load, the number and types of structural variation, and other genomic features of the parent, F1 and F2 intraspecific hybrids. We found evidence for large-scale phenotypic and genome-wide differentiation in response to divergent laboratory selection. Divergent-selected populations produced hybrids with reduced gamete viability-a classic signature of postzygotic reproductive isolation in the form of hybrid breakdown. Parallel-selected populations, on the other hand, remained more reproductively compatible (with exceptions). We found that F2 hybrid genomes contained vast genomic instability, that is, new structural variants (especially insertions, deletions and interchromosomal translocations) that were not observed in parent and F1 genomes, which is likely a result of chromosome missegregation and recombination errors in hybrid meiosis. Our results provide phenotypic and genomic evidence that partial reproductive isolation evolved due to adaptation to divergent environments, consistent with predictions of ecological speciation theory.

Reproductive Isolation

Divergent selection on locomotor activity in Drosophila melanogaster. II. Test for reproductive isolation between selected lines.

Tests for reproductive isolation between lines selected for locomotor activity were performed. Three sets of selection lines were used, each consisting of lines selected for low and high locomotor activity from the same base population. Females preferred high-activity males in almost every case. However, in one of the sets temporary sexual isolation was found between flies of the high and low lines. This was accompanied in the low-activity females with a higher fertility when they were mated with their own males. After further selection the partial isolation disappeared.

Animals

Reproductive isolation with little genetic divergence in sympatric populations of brown trout (Salmo trutta).

Two reproductively isolated demes of brown trout coexist in a small Swedish mountain lake, Lake Bunnersjörna. We electrophoretically examined 102 specimens from that lake for 27 enzymes encoded by 54 loci. The two demes are fixed for different alleles at a lactate dehydrogenase locus (LDH-1); statistically significant allele frequency differences at five other loci further support the complete lack of gene flow between these demes. There are significant differences in growth rates between fish in the two demes, but no further morphological differentiation h-s been detected.--In light of these findings, the genetic distance between these populations is surprisingly small (Nei's I = 0.975). These demes represent one of the least genetically divergent, reproductively isolated sympatric pair of vertebrate populations that have been identified. The results are discussed from both an evolutionary and ecological perspective.

Alleles

Pleiotropic effects of environment-sensitive genes affecting fitness in relation to postmating reproductive isolation.

With regard to speciation in sexually reproducing organisms, some population geneticists continue to argue about the relative merits of sympatry versus allopatry. However, all workers seem quite comfortable with the conventional scenario depicting how reproductive isolation arises between subpopulations in the state of incipient speciation. This view according to which the evolution of reproductive isolation mainly results from some genetic divergence consecutive to a substantial restriction in gene flow is questioned here. A verbal model is described in which gene flow is no longer seen as being first interrupted by a mere physical barrier. The model is based on limited genetic changes at loci influencing fitness but it places two important constraints on the properties of the genetic elements involved in it. One of them is concerned with the environment-sensitivity of the mutations implicated in the process, and the other with their presumed pleiotropic action on a behavioural trait.

Animals

No character displacement for reproductive isolation between Drosophila bipectinata and Drosophila malerkotliana.

To test whether character displacement for reproductive isolation between Drosophila bipectinata and Drosophila malerkotliana exists, the degree of sexual isolation was measured between their sympatric and allopatric populations. Although the isolation indices vary in different crosses, the average isolation index for sympatric populations is very close to that for allopatric populations. This shows no difference in the degree of sexual isolation between sympatric and allopatric populations of D. bipectinata and D. malerkotliana. Thus there is no evidence for the existence of character displacement for sexual isolation between these two closely related sympatric species.

Animals

The genetics of Artemia salina. VII. Reproductive isolation.

Fifteen of 20 gonochoristic Artemia populations are crossfertile with diploid San Francisco shrimps, producing fertile F1 and viable F2 progeny. Partial sex linkage of white eye was observed and frequency of crossing over between the white and sex loci did not exceed the range of values observed in San Francisco shrimps. Possible mechanisms for wide dispersal of this diploid genotype are discussed. Five populations are reproductively isolated from San Francisco shrimps: Mono Lake, Hidalgo, Lake Urmia, San Bartolomeo, and Tunisia. The last two are inter-fertile.

Animals

Sex chromosome translocations in the evolution of reproductive isolation.

Haldane's rule states that in organisms with differentiated sex chromosomes, hybrid sterility or inviability is generally expressed more frequently in the heterogametic sex. This observation has been variously explained as due to either genic or chromosomal imbalance. The fixation probabilities and mean times to fixation of sex-chromosome translocations of the type necessary to explain Haldane's rule on the basis of chromosomal imbalance have been estimated in small populations of Drosophila melanogaster. The fixation probability of an X chromosome carrying the long arm of the Y(X.Y(L)) is approximately 30% greater than expected under the assumption of no selection. No fitness differences associated with the attached Y(L) segment were detected. The fixation probability of a deficient Y chromosome is 300% greater than expected when the X chromosome contains the deleted portion of the Y. It is suggested that sex-chromosome translocations may play a role in the establishment of reproductive isolation.

Animals

Adding the heterochromatic YL arm to an X chromosome reduces reproductive fitnesses in Drosophila melanogaster: implications for the evolution of rDNA, heterochromatin, and reproductive isolation.

For X-Y exchange to be of importance in the coevolution of X and Y rDNA, there must be a mechanism to maintain cytologically normal X chromosomes in the face of continual infusions of X.YL chromosomes produced by X-Y exchanges. Replicated populations were founded with different frequencies of isogenic X and X.YL chromosomes. The X.YL chromosome declined in frequency over time in all lines. Relative fitnesses, estimated from chromosome frequency trajectories, were 0.40, 1.01, and 1.0 for X.YL/X.YL, X.YL/X, and X/X females and 0.75 and 1.0 for X.YL/Y and X/Y males, respectively. The equilibrium frequency for the X.YL chromosome due to the balance between X-Y exchange and selection was predicted to be 4-16 x 10(-4). The results strengthen the evidence for the involvement of X-Y exchange in the coevolution of X and Y rDNA arrays. Conditions for the evolution of reproductive isolation by sex-chromosome translocation are much less probable than previously supposed since the X.YL translocation chromosome is at a selective disadvantage to cytologically normal X chromosomes. Additional heterochromatin was not neutral but was only deleterious beyond a threshold, as one dose of the heterochromatic XL arm did not reduce female reproductive fitness, but two doses did.

Animals

Advances in the genetics of reproductive isolation in Drosophila.

Speciation genetics is defined as the study of genetic events and processes that differentiate the probabilities that genetic material from individual members of a population will co-occur in individuals of some future generation. It follows that phenotypic attributes that contribute to this differentiation of probabilities (e.g., mating preferences, sterility, or infertility of individuals from certain types of matings) constitute the phenotype of speciation, and genetic loci that may affect these phenotypic attributes can be considered as speciation genes. The literature on genetic differences between hybridizable species of Drosophila that are responsible for morphological differences, mating preferences, hybrid inviability, and hybrid sterility are reviewed with special reference to the species pair D. mojavensis - D. arizonensis. The case for the involvement of karyotypic changes in speciation in rodents is briefly discussed. It is concluded that no major advance has been made in the speciation genetics of Drosophila since Dobzhansky initiated the field 40 years ago. Yet, the identification of several gene loci that cause hybrid inviability or sterility may open the way to the understanding of reproductive isolation at the molecular level. It is not clear whether this approach will lead to general molecular mechanisms underlying the speciation process.

Animals

Microorganisms associated with chromosome destruction and reproductive isolation between two insect species.

Microorganisms have been implicated in causing cytoplasmic incompatibility in a variety of insect species, including mosquitoes, fruitflies, beetles and wasps. The effect is typically unidirectional: incompatible crosses produce no progeny or sterile males, whereas the reciprocal crosses produce normal progeny. The parasitic wasp Nasonia vitripennis is one of the few species in which the cytogenetic mechanism of incompatibility is known. In this species the paternal chromosome set forms a tangled mass in a fertilized egg and is eventually lost. Here we report that cytoplasmic microorganisms are associated with complete bidirectional incompatibility between N. vitripennis and a closely related sympatric species, N. giraulti. Microorganisms can be seen in the eggs of both species. Hybrid offspring are normally not produced in crosses between the two species, but do occur after elimination of the microorganisms by antibiotic treatment. A cytogenetic and genetic study shows that bidirectional interspecific incompatibility is due to improper condensation of the paternal chromosomes. Microorganism-mediated reproductive isolation is of interest because it could provide a rapid mode of speciation. The mechanism of incompatibility in Nasonia is also of interest as a potential tool for studying chromosome imprinting and chromosome condensation.

Animals

Strong reproductive isolation between closely related tropical sea urchins (genus Echinometra).

Morphological, mitochondrial DNA, and single-copy nuclear DNA differences show that the tropical sea urchin Echinometra mathaei is composed of at least four independent gene pools. Evolutionary distance between species measured with restriction-site changes (for mitochondrial DNA) and thermal renaturation (for single-copy nuclear DNA) is 1%-3% nucleotide divergence. Thus these are the most closely related sea urchin species known. Despite this genetic similarity, strong blocks to interspecific fertilization exist in this genus. Between two Hawaiian species, few eggs are fertilized in hybrid crosses, even in the presence of excess sperm. Microscopic examination of such crosses shows that sperm attachment to heterologous eggs is inhibited. Measures of genetic distance between species can help reveal the tempo of speciation and allow comparisons of morphological, biochemical, and ecological characteristics to be made in an evolutionary framework. Our results show that strong reproductive isolation can evolve by changes in egg-sperm recognition without extensive genetic divergence between species. Such mechanisms are most easily studied in free-spawning animals such as sea urchins but as well may represent an important aspect of speciation in species with internal fertilization.

Animals

Cytogenetic components of reproductive isolation in Trimerotropis thalassica and T. occidentalis.

The grasshopper Trimerotropis thalassica (Bruner) has a diploid count of 2n=23 male (XO), 24 female (XX). The two largest autosomes pairs are regularly metacentric, a consequence of fixed pericentric inversions. The X-chromosome is also a fixed metacentric. The remaining nine pairs of autosmes are polymorphic for floating percentric inversions so that the complement consists of a mixture of telocentric and metacentric members. Trimerotropis occidentalis (Bruner) is polymorphic for comparable inversions in only two of its autosome pairs and has a telocentric X. It is however, unique among the species of the genus Trimerotropis in having only 21 chromosomes in its male diploid set in all the populations so far studied. A single male found in a mixed population of these two species at Jasper Ridge, Stanford University, was characterized by the count 2n=22 male. In both this respect of and in its phenotype it was intermediate in character, representing a natural F1 hybrid between the two species. Cytogenetic analysis of this hybrid male indicated that occidentalis is differentiated from thalassica only is respect of a single tandem translocation. This has involved two of the telocentric elements of thalassica which have fused into a single composite telocentric partly homologous with each of the smaller progenitors. Although potentially capable of forming a multiple of three, one or other of the progenitor chromosomes regularly fails to pair with the tandem product in the hybrid so that one or more univalents invariably occur. These, by lagging, prevent cytokinesis and subsequently lead to the formation of macrospermatids which inevitably produce a measure of sterility. It is argued that this sterility provides a basis of reproductive isolation.

Animals

Evolutionary genetics of the Drosophila montium subgroup. I. Reproductive isolations and the phylogeny.

Seventeen species of the Drosophila montium subgroup, originated from the Southeast Asia, were genetically examined to clarify the phylogenetic relationships. Among 272 interspecific crosses, 61 combinations were successful in mating and 39 combinations produced hybrid flies. These results enabled us to classify the subgroup into three species complexes: the kikkawai complex (6 species), the jambulina complex (4 species) and the auraria complex (7 species), which were very similar to that obtained by the electrophoretic classification (Ohnishi and Watanabe, 1984). Asymmetrical mating preference between species was found in the present experiment. They were applied to estimate the relative age of species according to the hypothesis proposed by Watanabe and Kawanishi (1979). The evolving order was as follows: kikkawai, leontia, pennae, lini-like, lini, bocki in the kikkawai complex, punjabiensis, punjabiensis-like, jambulina, barbarae in the jambulina complex, and quadraria, yuwanensis, rufa, subauraria, biauraria, triauraria, auraria in the auraria complex. Hybrid flies, if produced, were mostly fertile or partially fertile in both sexes (32/39) within species complex crosses. Therefore premating isolation played a more important role than postmating isolation in speciation of the D. montium subgroup.

Animals