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[The hypothesis of immune testing of partners--coordinated adaptations and changes in mating preferences].

Reproductive isolation plays the key role in speciation. According to the prevailing ideas, the main speciation mechanism is gradual accumulation of genetic differences in isolated populations (allopatric phase of speciation) based on mutations, selection, and genetic drift. In this case, reproductive isolation emerges as an occasional byproduct of adaptation to different conditions (ecological speciation) or accumulation of random changes in the gene pool resulting from long-term isolation. Pure sympatric speciation assumes isolation as a direct product of selection (divergent or disruptive selection) that favors individuals selectively mating with their likes. A third possibility is substantiated below. We believe that isolation can be a regular and determined product rather than occasional byproduct of divergence. It can rely on the friend/foe discrimination mechanisms, some of which can be "immune-based" and compare the partner's and own properties (signaling molecules, pheromones, and other antigens in a broad sense). Antigens of the major histocompatibility complex (MHC) can play a substantial role in such testing of potential mates.

Adaptation, Psychological↗

Natural selection at work.

Although much is known about the genetic basis of reproductive isolation between species, little is understood about its underlying evolutionary causes. A study of two very closely related, but reproductively isolated, plant species has provided some valuable insights.

Plant Physiological Phenomena↗

Experimental evidence of hybrid breakdown between genetically distinct populations of Echinostoma caproni.

In this paper we investigate whether the assortative mating between individuals of the same genetic entity (i.e. coming from the same geographical area) of Echinostoma caproni, reported in a previous paper, may be explained by a post-zygotic isolating mechanism. The fecundity of the adults of 2 parental genetic entities and of their hybrids (i.e. F1, F2, F3) was quantified through 3 successive generations. Whereas the number of eggs released by F1 hybrids is similar to that of the mid-parent, that of recombinant hybrids (F2-3 hybrids) is significantly lower than that of F1 and that of the mid-parent. Since these results seem to demonstrate hybrid breakdown, 2 important factors maintaining reproductive isolation, i.e. pre-mating reproductive isolation and low hybrid fecundity, may influence the evolution of E. caproni.

Analysis of Variance↗

Population structure attributable to reproductive time: isolation by time and adaptation by time.

Many populations are composed of a mixture of individuals that reproduce at different times, and these times are often heritable. Under these conditions, gene flow should be limited between early and late reproducers, even within populations having a unimodal temporal distribution of reproductive activity. This temporal restriction on gene flow might be called "isolation by time" (IBT) to acknowledge its analogy with isolation by distance (IBD). IBD and IBT are not exactly equivalent, however, owing to differences between dispersal in space and dispersal in time. We review empirical studies of natural populations that provide evidence for IBT based on heritabilities of reproductive time and on molecular genetic differences associated with reproductive time. When IBT is present, variation in selection through the reproductive season may lead to adaptive temporal variation in phenotypic traits [adaptation by time (ABT)]. We introduce a novel theoretical model that shows how ABT increases as (i) selection on the trait increases; (ii) environmental influences on reproductive time decrease; (iii) the heritability of reproductive time increases; and (iv) the temporal distribution of reproductive activity becomes increasingly uniform. We then review empirical studies of natural populations that provide evidence for ABT by documenting adaptive temporal clines in phenotypic traits. The best evidence for IBT and ABT currently comes from salmonid fishes and flowering plants, but we expect that future work will show these processes are more widespread.

Adaptation, Physiological↗

Genetics of flower size and nectar volume in Petunia pollination syndromes.

The two related Petunia species, P. axillaris and P. integrifolia, are sympatric at various locations in South America but do not hybridise. Divergent pollinator preferences are believed to be in part responsible for their reproductive isolation. The volume of nectar produced and several components of flower morphology might contribute to pollinator-dependant reproductive isolation. In this study, we aimed to identify the genetic changes underlying the quantitative differences observed between these two Petunia species in flower size and nectar volume. We mapped quantitative trait loci (QTL) responsible for the different phenotypes of P. axillaris and P. integrifolia in an inter-specific backcross population. QTL of small to moderate effect control the differences in flower size and volume of nectar. In addition, we observed strong suppression of meiotic recombination in Petunia, even between closely related species, which precluded a fine resolution of QTL mapping. Thus, our data suggest that flower size and nectar volume are highly polygenic. They are likely to have evolved gradually through pollinator-mediated adaptation or reinforcement, and are not likely to have been primary factors in early steps of pollinator isolation of P. axillaris and P. integrifolia.

Crosses, Genetic↗

Effects of a bovine herpesvirus-1 isolate on reproductive function in heifers: classification as a type-2 (infectious pustular vulvovaginitis) virus by restriction endonuclease analysis of viral DNA.

A bovine herpesvirus-1 (BHV-1) isolate (FI) from an aborted fetus was used to infect 9 heifers at various stages of gestation. Two heifers were inoculated IV on postbreeding day (PBD) 1, 7, or 14, and 3 heifers were inoculated in the sixth month of pregnancy. Plasma progesterone assays were used to monitor corpus luteum function in heifers inoculated during early pregnancy. Low progesterone values and infertility were seen in the 2 heifers inoculated on PBD 1. Luteal function remained normal in heifers inoculated on PBD 7 or 14. These 4 heifers inoculated on PBD 7 or 14 carried their fetuses to term, and their calves were free of BHV-1 infection at birth. Three heifers inoculated during the sixth month of pregnancy also carried their fetuses to term. Two calves were born alive, and BHV-1 was not isolated from nasal swab samples of either calf; the third calf was stillborn. Virus was not isolated from the stillborn calf's tissues, but BHV-1 was isolated from the placenta. Lesions were not detected in several tissues examined by light microscopy, and BHV-1 antigen was not detected by immunohistochemical examination of paraffin sections. Restriction endonuclease analysis of viral DNA was used to compare the FI virus to other BHV-1 isolates (Colorado-1, Iowa, and K22). On the basis of restriction endonuclease analysis, the FI isolate should be classified as a type-2 (infectious pustular vulvovaginitis) virus, specifically subtype a.

Animals↗

A comprehensive study of genic variation in natural populations of Drosophila melanogaster. VII. Varying rates of genic divergence as revealed by two-dimensional electrophoresis.

Four sibling species from the melanogaster subgroup (Drosophila melanogaster, D. simulans, D. sechellia, and D. mauritiana) were studied for genetic divergence, by high-resolution two-dimensional protein electrophoresis (2DE) coupled with ultrasensitive silver staining. A total of eight tissues from larval and adult developmental stages representing both gonadal (germ-line) and nongonadal (somatic) tissues were analyzed for protein divergence between species. Close to 400 polypeptides (protein spots) were scored from each tissue and species, and protein divergence was measured on the basis of qualitative differences (presence/absence) of protein spots in pairwise species comparisons. The observed levels of genic divergence varied among tissues and among species. When larval hemolymph proteins (which are known to be highly polymorphic) were excluded, there was no evidence to suggest that either the larval or adult-stage proteins, as a whole, are more diverged than the other; variation between different tissues rather than between developmental stages appears to be the most significant factor affecting genetic divergence between species. The reproductive tissue (testis and accessory gland) showed more divergence than did the nonreproductive tissue; D. melanogaster testis (from both larvae and adult males) showed the highest level of divergence. In view of the previous observation that D. simulans, D. mauritiana, and D. sechellia show similar but significantly less reproductive isolation from each other than from D. melanogaster, the present results suggest a correlation between the levels of reproductive-tract-protein divergence and the degree of reproductive isolation in these species.

Animals↗

Speciation by postzygotic isolation: forces, genes and molecules.

New species arise as reproductive isolation evolves between diverging populations. Here we review recent work in the genetics of postzygotic reproductive isolation-the sterility and inviability of species hybrids. Over the last few years, research has taken two new directions. First, we have begun to learn a good deal about the population genetic forces driving the evolution of postzygotic isolation. It has, for instance, become increasingly clear that conflict-driven processes, like sexual selection and meiotic drive, may contribute to the evolution of hybrid sterility. Second, we have begun to learn something about the identity and molecular characteristics of the actual genes causing hybrid problems. Although molecular genetic data are limited, early findings suggest that "speciation genes" correspond to loci having normal functions within species and that these loci sometimes diverge as a consequence of evolution in gene regulation.

Animals↗

Fine-scale genetic structure, estuarine colonization and incipient speciation in the marine silverside fish Odontesthes argentinensis.

The identification of incipient ecological species represents an opportunity to investigate current evolutionary process where adaptive divergence and reproductive isolation are associated. In this study we analysed the genetic structure of marine and estuarine populations of the silverside fish Odontesthes argentinensis using nine microsatellite loci and 396 bp of the mitochondrial DNA (mtDNA) control region. Our main objective was to investigate the relationship among estuarine colonization, divergent selection and speciation in silversides. Significant genetic structure was detected among all marine and estuarine populations. Despite the low phylogeographic structure in mtDNA haplotypes, there was clear signal of local radiations of haplotypes in more ancient populations. Divergence among marine populations was interpreted as a combined result of homing behaviour, isolation by distance and drift. On the other hand, ecological shifts due to the colonization of estuarine habitats seem to have promoted rapid adaptive divergence and reproductive isolation in estuarine populations, which were considered as incipient ecological species. This conclusion is supported by the existence of a set of environmental factors required for successful reproduction of estuarine ecotypes. The pattern of genetic structure indicates that phenotypic and reproductive divergence evolved in the face of potential gene flow between populations. We suggest that the 'divergence-with-gene-flow' model of speciation may account for the diversification of estuarine populations. The approach used can potentially identify 'incipient estuarine species', being relevant to the investigation of the evolutionary relationships of silversides in several coastal regions of the world.

Animals↗

Inferring the history of speciation from multilocus DNA sequence data: the case of Drosophila pseudoobscura and close relatives.

The divergence of Drosophila pseudoobscura from its close relatives, D. persimilis and D. pseudoobscura bogotana, was examined using the pattern of DNA sequence variation in a common set of 50 inbred lines at 11 loci from diverse locations in the genome. Drosophila pseudoobscura and D. persimilis show a marked excess of low-frequency variation across loci, consistent with a model of recent population expansion in both species. The different loci vary considerably, both in polymorphism levels and in the levels of polymorphisms that are shared by different species pairs. A major question we address is whether these patterns of shared variation are best explained by gene flow or by persistence since common ancestry. A new test of gene flow, based on patterns of linkage disequilibrium, is developed. The results from these, and other tests, support a model in which D. pseudoobscura and D. persimilis have exchanged genes at some loci. However, the pattern of variation suggests that most gene flow, although occurring after speciation began, was not recent. There is less evidence of gene flow between D. pseudoobscura and D. p. bogotana. The results are compared with recent work on the genomic locations of genes that contribute to reproductive isolation between D. pseudoobscura and D. persimilis. We show that there is a good correspondence between the genomic regions associated with reproductive isolation and the regions that show little or no evidence of gene flow.

Animals↗

The ecological and evolutionary consequences of sperm chemoattraction.

Chemical communication between sperm and egg is a critical factor mediating sexual reproduction. Sperm attractants may be significant evolutionarily for maintaining species barriers, and important ecologically for increasing gamete encounters. Still unresolved, however, are the functional consequences of these dissolved signal molecules. Here, we provide experimental evidence that sperm chemoattraction directly affects the magnitude of fertilization success. The recent discovery of L-tryptophan as a potent attractant to red abalone (Haliotis rufescens) sperm affords the opportunity to quantify how navigation affects gamete interactions. Sperm behavioral responses to manipulations of the natural tryptophan gradient around individual eggs reveals that both chemotaxis and chemokinesis significantly promote contacts. Our results show further that attractant release by means of diffusion effectively doubles the target size of red abalone eggs, which in turn significantly increases fertilization success. Although long theorized as potential barriers to hybridization, species-specific sperm attractants in red and green (Haliotis fulgens) abalone are only minor contributors to maintaining reproductive isolation. Because abalone typically live in dense, multispecies aggregations, chemically mediated navigation would prevent sperm from pointlessly tracking heterospecific eggs. Thus, even though reproductive isolation fundamentally resides at the level of membrane recognition proteins, species-specific sperm attractants may evolve to locate the right target within mixed gamete suspensions of closely related species.

Animals↗

Recombination and the divergence of hybridizing species.

The interplay between hybridization and recombination can have a dramatic effect on the likelihood of speciation or persistence of incompletely isolated species. Many models have suggested recombination can oppose speciation, and several recent empirical investigations suggest that reductions in recombination between various components of reproductive isolation and/or adaptation can allow species to persist in the presence of gene flow. In this article, we discuss these ideas in relation to speciation models, phylogenetic analyses, and species concepts. In particular, we revisit genetic architectures and population mechanisms that create genetic correlations and facilitate divergence in the face of gene flow. Linkage among genes contributing to adaptation or reproductive isolation due to chromosomal rearrangements as well as pleiotropy or proximity of loci can greatly increase the odds of species divergence or persistence. Finally, we recommend recombination to be a focus of inquiry when studying the origins of biological diversity.

Biological Evolution↗

Does competitive divergence occur if assortative mating is costly?

Most models of sympatric speciation have assumed that assortative mating has no costs. A few studies, however, have shown that the costs for being choosy can prevent such speciation. Here, we investigate the role of the strength of assortment and of the costs for being choosy for a simple genetic model of a single ('magic') trait that mediates both intraspecific competition for a continuum of resources and assortative mating, which is induced by choosy females who preferentially mate with males of similar phenotype. Choosiness may be costly if it is difficult to find a mating partner. Such magic trait models are considered to be most conducive of sympatric speciation. We consider a sexually reproducing population of haploid individuals that is density regulated. The trait is determined by a single locus with multiple alleles. The strength of stabilizing selection (caused by a unimodal resource distribution), the strength of competition, the degree of assortment and the costs for being choosy are independent parameters. We investigate analytically and numerically how these parameters determine the equilibrium and stability structure. In particular, we identify conditions under which no polymorphism at all is maintained as well as conditions under which strong competitive divergence occurs, or the population even splits into two reproductively isolated classes of highly diverse phenotypes. If costs are absent or moderate, genetic variability tends to be minimized at intermediate strengths of assortment, and reproductively isolated classes of phenotypes are a likely result of evolution only for intermediate or strong competition and for very strong assortment. The likelihood of divergence depends relatively weakly on the costs as long as they are not high. With high costs, however, increasingly strong assortment rapidly depletes all genetic variation, and strong competitive divergence is prevented.

Evolution, Molecular↗

The effect of isolation on reproduction and growth of Pseudosuccinea columella (Pulmonata: Lymnaeidae): a snail-conditioned water experiment.

A snail-conditioned water experiment was conducted in Pseudosuccinea columella to test the possible role of a chemical interaction between snails on the diminished growth and fecundity rates found for snails raised in pairs compared to those raised in complete isolation. The results permit to discard the hypothesis of an inhibition of growth and reproduction between snails due to factors released into the water.

Animals↗

Gametic incompatibilities between races of Drosophila melanogaster.

Reproductive-isolating mechanisms between nascent species may involve sperm-egg recognition and have been best described in externally fertilizing organisms where such recognition is essential in preventing undesirable fertilizations. However, reproductive barriers in internally fertilizing species differ in significant ways, and a direct role for sperm-egg interactions has yet to be demonstrated. Females of many strains of Drosophila melanogaster from Zimbabwe, Africa, do not mate readily with cosmopolitan males. This polymorphism in mate choice is postulated to represent incipient speciation. We now report that, in one direction, crosses between the above populations produce far fewer offspring than reciprocal crosses due to a lower rate of egg hatch. We established that egg inviability in these crosses was due to defects in fertilization. Thus, even in taxa with internal fertilization, gametic incompatibility may be a mechanism relevant to reproductive isolation during incipient speciation.

Animals↗

Parallel speciation: a key to sympatric divergence.

Until recently, our view of speciation was that reproductive isolation evolves during long periods of allopatry through the accumulation of genetic differences that result in genetic incompatibility. This view now contrasts with new findings that characters affecting reproductive isolation can diverge rapidly in sympatry as a result of natural selection. Recent studies combining research on phylogeny and ecology of natural populations cast new light on patterns, timescales and mechanisms, and emphasize the role of ecological factors in speciation. Studies of parallel speciation provide a strong case for sympatric speciation and for natural selection generating reproductive barriers.

Journal Article↗

Inter-locus antagonistic coevolution as an engine of speciation: assessment with hemiclonal analysis.

One of Ernst Mayr's legacies is the consensus that the allopatry model is the predominant mode of speciation in most sexually reproducing lineages. In this model, reproductive isolation develops as a pleiotropic byproduct of the genetic divergence that develops among physically isolated populations. Presently, there is no consensus concerning which, if any, evolutionary process is primarily responsible for driving the specific genetic divergence that leads to reproductive isolation. Here, we focus on the hypothesis that inter-locus antagonistic coevolution drives rapid genetic divergence among allopatric populations and thereby acts as an important "engine" of speciation. We assert that only data from studies of experimental evolution, rather than descriptive patterns of molecular evolution, can provide definitive evidence for this hypothesis. We describe and use an experimental approach, called hemiclonal analysis, that can be used in the Drosophila melanogaster laboratory model system to simultaneously screen nearly the entire genome for both standing genetic variation within a population and the net-selection gradient acting on the variation. Hemiclonal analysis has four stages: (i) creation of a laboratory "island population"; (ii) cytogenetic cloning of nearly genome-wide haplotypes to construct hemiclones; (iii) measurement of additive genetic variation among hemiclones; and (iv) measurement of the selection gradient acting on phenotypic variation among hemiclones. We apply hemiclonal analysis to test the hypothesis that there is ongoing antagonistic coevolution between the sexes in the D. melanogaster laboratory model system and then discuss the relevance of this analysis to natural systems.

Animals↗

[Characteristics of lactobacillus species strains isolated from reproductive organs in various clinical cases].

The study was aimed at detection of presence of Lactobacillus species bacteria in 167 samples of material derived from vagina of patients with cervix uteri (22 samples), chronic vaginosis (100 samples) and from women after treatment of inflammatory diseases (45 samples). Out of 100 isolated strains of Lactobacillus only 13 belonged to L. acidophilus. They were mainly present (10 strains) in patients after effective treatment of inflammatory diseases of reproductive organs. Remaining strains belonged to L. jensenii (18 strains) and Lactobacillus sp. (69 strains) and did not cleave glycogen. This was detected by quantitative determinations of lactic acid in cultures of these bacteria in the fluid media containing 0.25% of glycogen. It seems that treatment of bacterial infections of reproductive organs should be concentrated on not only elimination of pathogenic bacteria, but also on noneffective strains from Lactobacillus species (strains unable to cleave glycogen with production of lactic acid).

Chronic Disease↗