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Evolutionary divergence and possible incipient speciation in post-glacial populations of a cosmopolitan aquatic plant.

Habitat configuration is expected to have a major influence on genetic exchange and evolutionary divergence among populations. Aquatic organisms occur in two fundamentally different habitat types, the sea and freshwater lakes, making them excellent models to study the contrasting effects of continuity vs. isolation on genetic divergence. We compared the divergence in post-glacial populations of a cosmopolitan aquatic plant, the pondweed Potamogeton pectinatus that simultaneously occurs in freshwater lakes and coastal marine sites. Relative levels of gene flow were inferred in 12 lake and 14 Baltic Sea populations in northern Germany using nine highly polymorphic microsatellite markers developed for P. pectinatus. We found highly significant isolation-by-distance in both habitat types (P < 0.001). Genetic differentiation increased approximately 2.5-times faster among freshwater populations compared with those from the Baltic Sea. As different levels of genetic drift or population history cannot explain these differences, higher population connectivity in the sea relative to freshwater populations is the most likely source of contrasting evolutionary divergence. These findings are consistent with the notion that freshwater angiosperms are more conducive to allopatric speciation than their life-history counterparts in the sea, the relative species poor seagrasses. Surprisingly, population pairs from different habitat types revealed almost maximal genetic divergence expected for complete reproductive isolation, regardless of their respective geographical distance. Hence, the barrier to gene flow between lake and sea habitat types cannot be due to dispersal limitation. We may thus have identified a case of rapid incipient speciation in post-glacial populations of a widespread aquatic plant.

Baltic States↗

[Descent of ova and embryos in cows superovulated with serum gonadotropin].

The descent and localization of eggs and embryos in individual segments of the reproductive tract of superovulated cows were studied in this work. For the induction of superovulation, serum gonadotropin (PMSG, Ivanovice in Haná) at a dose of 2,500-3,500 I.U. was used, in combination with 0.5 mg of Cloprostenol (Oestrophan, Spofa), administered 48 hours after gonadotropin treatment. The start of superovulation fell on days 9 to 12 of the sexual cycle and was conditioned by the presence of the corpus luteum (CL). After the onset of the heat, 2-3 inseminations were carried out using fresh semen. Donor cows were slaughtered 3, 4, 5, 6, and 7 days after the second insemination and isolated reproductive organs (Fig. 1) were divided into five segments (two on oviducts and three on uterine horns) by the applied ligature. In laboratory conditions superovulation response was determined accurately, the volume of ovaries was assessed according to water displacement and the segments of oviducts and uterus were rinsed with TCM 199 or PBS supplemented with FCS. 3, 4, 5, 6, and 7 days after insemination (Tab. I). 18.1 (+/- 3.55), 12.4 (+/- 0.91), 19.2 (+/- 2.86), 20 and 23 (+/- 2.44) CL on average were recorded, which corresponded to the ovulation of 64, 50, 56, 71 and 72 percent of stimulated follicles (Fig. 2). Within 3 to 7 days after insemination nearly triple enlargement of ovaries was also observed (Tab. I, Fig. 3). During the lavage of individual segments of the tubular reproductive tract, 38 per cent of eggs and embryos were detected in the uterus as early as 3 days after insemination (Tab. II). Unfertilized eggs and degenerated embryos were found in the 2nd and 3rd uterine segment, embryos at the stage of 8-16 blastomeres were localized in the 1st and 2nd segment of the uterus. Four days after insemination (Tab. III), about 64 per cent of eggs and embryos at the stage up to 16 blastomeres were found in the uterus, but embryos up to 32 blastomeres were still flushed out of the oviduct. On day 5 after insemination, 92 per cent of eggs and embryos were released into the uterus, being localized mostly in the cranial and medial part of the uterus (Tab. IV). 7.5 per cent of recovered eggs and embryos at the stage of early or compacted morulae were still detected in the oviducts.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Hybrids of two closely related tropical sea urchins (genus Echinometra): evidence against postzygotic isolating mechanisms.

A series of cross-fertilization experiments were conducted with two unnamed, sympatric species of sea urchins in the Echinometra mathaei species complex, Echinometra sp. A (Ea) and Echinometra sp. C (Ec). Heterogametic fertilization success was high when eggs of Ec and sperm of Ea were involved, and low with eggs of Ea and sperm of Ec. Hybrids produced from crosses in either direction developed normally to sexually mature adults; Ea x Ea were largest in test size, followed by Ec (ova) x Ea (sperm), Ea (ova) x Ec (sperm), and Ec x Ec, respectively. Color patterns of the hybrids were closer to the maternal coloration, whereas other characters such as relative test dimensions and spine lengths, morphology of tubefoot and gonad spicules, and gamete sizes were intermediate. Fertilization rates in F1 backcrosses were high, minimizing the possibility that hybrid infertility is a postzygotic mechanism of reproductive isolation. On the other hand, intensive surveys failed to find individuals with hybrid characteristics in the field, suggesting that natural hybridization between the two species is rare. Prezygotic isolating mechanisms, such as microhabitat separation and gamete incompatibility, at least between Ea eggs and Ec sperm, most likely maintain the genetic integrity of these two closely related species.

Animals↗

The evolution and maintenance of monoecy and dioecy in Sagittaria latifolia (Alismataceae).

Plant species rarely exhibit both monoecious and dioecious sexual systems. This limits opportunities to investigate the consequences of combined versus separate sex function on mating patterns and genetic variation and the analysis of factors responsible for the evolution and maintenance of the two sexual systems. Populations of the North American clonal aquatic Sagittaria latifolia are usually either monoecious or dioecious and often grow in close geographic proximity. We investigated mating patterns, genetic structure, and relationships between the two sexual systems using allozyme variation in populations from southern Ontario, Canada. As predicted, selfing rates in monoecious populations (n = 6, mean = 0.41) were significantly higher than in dioecious populations (n = 6, mean = 0.11). Moreover, marker-based estimates of inbreeding depression (delta) indicated strong selection against inbred offspring in both monoecious (mean delta = 0.83) and dioecious (mean delta = 0.84) populations. However, the difference in selfing rate between the sexual systems was not reflected in contrasting levels of genetic variation. Our surveys of 12 loci in 15 monoecious and 11 dioecious populations revealed no significant differences in the proportion of polymorphic loci (P), number of alleles per locus (A), or observed and expected heterozygosity (H(o) and H(e), respectively). Strong inbreeding depression favoring survival of outcrossed offspring may act to maintain similar levels of diversity between monoecious and dioecious populations. Despite geographical overlap between the two sexual systems in southern Ontario, a dendrogram of genetic relationships indicated two distinct clusters of populations largely corresponding to monoecious and dioecious populations. Reproductive isolation between monoecious and dioecious populations appears to be governed, in part, by observed differences in habitat and life history. We suggest that selfing and inbreeding depression in monoecious populations are important in the transition from monoecy to dioecy and that the maintenance of distinct sexual systems in S. latifolia is governed by interactions between ecology, life history, and mating.

Biological Evolution↗

Developmental isolation and subsequent adult behavior of Drosophila paulistorum. VI. Quantitative variation in cuticular hydrocarbons.

Our previous studies have demonstrated that long-term chemical contact with heterospecifics during development minimized unproductive heterospecific matings among the six Drosophila paulistorum semispecies. When socially isolated from conspecifics, discrimination significantly decreased so that more ultimately unproductive heterospecific matings occurred. Such results suggest that learning and social experiences play roles in mate recognition, using chemical information. In investigations into the development of discriminatory behavior in D. paulistorum, social experiences influence the production of cuticular hydrocarbons in both sexes among the six different semispecies of D. paulistorum. Produced by both sexes, 2-methyl triacontane has been detected from egg stages on, gradually increasing with age. However, 11-docosenyl acetate, a male pheromone, was not produced until early adult stages. There were consistent significant differences in quantities and ratios of each of these hydrocarbons among the six semispecies, contributing to their reproductive isolation. Furthermore, we found significant quantitative differences in hydrocarbons between socially isolated flies and communally raised ones: Socially isolated flies produced significantly greater quantities of hydrocarbons than communally raised flies, consistent with previous behavioral data, as isolated flies court more vigorously and often.

Aging↗

Allozymic and chromosomal similarity in two Drosophila species.

D. setosimentum and ochrobasis are a pair of very close, partly sympatric species endemic to Hawaii island. Males of the two species differ strikingly in wing-pattern and there are altitudinal and breeding-site differences. Similarity indices have been calculated for both chromosomal (C) ald allozymic (A) variants. Within the main populations of each species both kinds of data give coefficients above 0.98. Interspecific comparison of the main populations shows 0.66 (C) and 0.79 (A). An isolated population of ochrobasis from Kohala Volcano (Ohu), when compared with setosimentum, shows 0.68 (C) and 0.98 (A). Chromosomes are thus much more sensitive than allozymes in distinguishing these species; the same is true in the case of D. silvestris and heteroneura from the same forests. These morphologically distinct species, when compared, show 0.96 (A). All four species appear to be very new in the historical sense. In one area, about 2% of wild-caught D. setosimentum/ochrobasis are interspecific hybrids although adequate samples indicate that the separate gene pools have not broken down. The specific names should be retained but the two entities are perhaps best described as quite advanced semispecies in which reproductive isolation in nature is now nearly complete.

Animals↗

[Hypothesis of "immunological testing" of partners--systems of "friend and foe" recognition in historical prospect].

The hypothesis of "autoimmune testing" of mating partners assumes formation of individual system of perception on the basis of an immunologic principle, i.e. by lifetime selection of "direct" or "reverse casts" of own key antigens or signal molecules. Such system provides the coordinated change of signaling systems and system of their perception at formation of new adaptations that leads to automatic formation of reproductive isolation within the limited number of generations. Presence of the "friend-foe" recognition systems practically in all living organisms assumes formation of potential mating partners "autoimmune testing" mechanisms at the earliest evolution stages. In this article we analyze possible mechanisms of a "friend and foe" discrimination with MHC-proteins and their homologues in historical prospect--from bacteria to the lower Chordates.

Animals↗

Interaction between natural and sexual selection during the evolution of mate recognition.

The interaction between natural and sexual selection is central to many theories of how mate choice and reproductive isolation evolve, but their joint effect on the evolution of mate recognition has not, to my knowledge, been investigated in an evolutionary experiment. Natural and sexual selection were manipulated in interspecific hybrid populations of Drosophila to determine their effects on the evolution of a mate recognition system comprised of cuticular hydrocarbons (CHCs). The effect of natural selection in isolation indicated that CHCs were costly for males and females to produce. The effect of sexual selection in isolation indicated that females preferred males with a particular CHC composition. However, the interaction between natural and sexual selection had a greater effect on the evolution of the mate recognition system than either process in isolation. When natural and sexual selection were permitted to operate in combination, male CHCs became exaggerated to a greater extent than in the presence of sexual selection alone, and female CHCs evolved against the direction of natural selection. This experiment demonstrated that the interaction between natural and sexual selection is critical in determining the direction and magnitude of the evolutionary response of the mate recognition system.

Animals↗

Evidence for inversion polymorphism related to sympatric host race formation in the apple maggot fly, Rhagoletis pomonella.

Evidence suggests that the apple maggot, Rhagoletis pomonella (Diptera: Tephritidae) is undergoing sympatric speciation (i.e., divergence without geographic isolation) in the process of shifting and adapting to a new host plant. Prior to the introduction of cultivated apples (Malus pumila) in North America, R. pomonella infested the fruit of native hawthorns (Crataegus spp.). However, sometime in the mid-1800s the fly formed a sympatric race on apple. The recently derived apple-infesting race shows consistent allele frequency differences from the hawthorn host race for six allozyme loci mapping to three different chromosomes. Alleles at all six of these allozymes correlate with the timing of adult eclosion, an event dependent on the duration of the overwintering pupal diapause. This timing difference differentially adapts the univoltine fly races to an approximately 3- to 4-week difference in the peak fruiting times of apple and hawthorn trees, partially reproductively isolating the host races. Here, we report finding substantial gametic disequilibrium among allozyme and complementary DNA (cDNA) markers encompassing the three chromosomal regions differentiating apple and hawthorn flies. The regions of disequilibrium extend well beyond the previously characterized six allozyme loci, covering substantial portions of chromosomes 1, 2, and 3 (haploid n = 6 in R. pomonella). Moreover, significant recombination heterogeneity and variation in gene order were observed among single-pair crosses for each of the three genomic regions, implying the existence of inversion polymorphism. We therefore have evidence that genes affecting diapause traits involved in host race formation reside within large complexes of rearranged genes. We explore whether these genomic regions (inversions) constitute coadapted gene complexes and discuss the implications of our findings for sympatric speciation in Rhagoletis.

Adaptation, Physiological↗

Sympatric speciation by sexual selection.

There is increasing evidence for the process of sympatric speciation, in which reproductive isolation of species occurs without physical isolation. Theoretical models have focused on disruptive natural selection as the crucial pressure for splitting a species. Here we report the theoretical finding that sympatric speciation may be caused by sexual selection even without disruptive natural selection. Specifically, we show that variation in a male secondary sexual character with two conspicuous extremes and the corresponding variance in female mating preference around no preference may jointly evolve into bimodal distributions with increasing modal divergence of the male and female traits, pulling a population apart into two prezygotically isolated populations. This mode of speciation, driven by two runaway processes in different directions, is promoted by an increase in the efficiency of females in discriminating among males or a decrease in the cost of male conspicuousness, indicating that sympatric speciation may occur more readily if barrier-free or predator-free conditions arise. Although even a slight cost of female preference would cancel the runaway process of sexual selection, it would not cancel the divergent runaway processes of sympatric speciation.

Animals↗

Characterization of genetic diversity in Dermacentor andersoni (Acari: Ixodidae) with body size and weight polymorphism.

Morphological and discrete genetic differences are found between geographically isolated, allopatric, tick populations. However, we have found differences in sympatric tick populations. Notable differences were found in the body size and weight of Dermacentor andersoni collected from a single location in Montana, USA. These ticks were separated in groups consisting of big (B) and small (S) individuals. The objectives of this study were: (a) to characterize genetic diversity in B and S D. andersoni individuals, (b) to evaluate transmissibility of the character associated with body size and weight, and (c) to correlate morphological differences with biological, physiological, and behavioral characteristics. We found extensive genetic variation in 16S rDNA and ITS2 loci in B and S ticks and demonstrated genetic differentiation between B and S individuals. We further provide some support for Mendelian autosomal dominant transmission of characters associated with tick body size and weight. The results reported herein show that B ticks have a better reproductive success than S ticks and suggest partial reproductive isolation of S ticks.

Animals↗

Smelling home can prevent dispersal of reef fish larvae.

Many marine fish and invertebrates show a dual life history where settled adults produce dispersing larvae. The planktonic nature of the early larval stages suggests a passive dispersal model where ocean currents would quickly cause panmixis over large spatial scales and prevent isolation of populations, a prerequisite for speciation. However, high biodiversity and species abundance in coral reefs contradict this panmixis hypothesis. Although ocean currents are a major force in larval dispersal, recent studies show far greater retention than predicted by advection models. We investigated the role of animal behavior in retention and homing of coral reef fish larvae resulting in two important discoveries: (i) Settling larvae are capable of olfactory discrimination and prefer the odor of their home reef, thereby demonstrating to us that nearby reefs smell different. (ii) Whereas one species showed panmixis as predicted from our advection model, another species showed significant genetic population substructure suggestive of strong homing. Thus, the smell of reefs could allow larvae to choose currents that return them to reefs in general and natal reefs in particular. As a consequence, reef populations can develop genetic differences that might lead to reproductive isolation.

Animal Migration↗

Rapidly evolving zona pellucida domain proteins are a major component of the vitelline envelope of abalone eggs.

Proteins harboring a zona pellucida (ZP) domain are prominent components of vertebrate egg coats. Although less well characterized, the egg coat of the non-vertebrate marine gastropod abalone (Haliotis spp.) is also known to contain a ZP domain protein, raising the possibility of a common molecular basis of metazoan egg coat structures. Egg coat proteins from vertebrate as well as non-vertebrate taxa have been shown to evolve under positive selection. Studied most extensively in the abalone system, coevolution between adaptively diverging egg coat and sperm proteins may contribute to the rapid development of reproductive isolation. Thus, identifying the pattern of evolution among egg coat proteins is important in understanding the role these genes may play in the speciation process. The purpose of the present study is to characterize the constituent proteins of the egg coat [vitelline envelope (VE)] of abalone eggs and to provide preliminary evidence regarding how selection has acted on VE proteins during abalone evolution. A proteomic approach is used to match tandem mass spectra of peptides from purified VE proteins with abalone ovary EST sequences, identifying 9 of 10 ZP domain proteins as components of the VE. Maximum likelihood models of codon evolution suggest positive selection has acted among a subset of amino acids for 6 of these genes. This work provides further evidence of the prominence of ZP proteins as constituents of the egg coat, as well as the prominent role of positive selection in diversification of these reproductive proteins.

Adaptation, Biological↗

Positive selection on an acrosomal sperm protein, M7 lysin, in three species of the mussel genus Mytilus.

Marine invertebrate sperm proteins are particularly interesting because they are characterized by positive selection and are likely to be involved in prezyogotic isolation and, thus, speciation. Here, we present the first survey of interspecific and intraspecific variation of a bivalve sperm protein among a group of species that regularly hybridize in nature. M7 lysin is found in sperm acrosomes of mussels and dissolves the egg vitelline coat, permitting fertilization. We sequenced multiple alleles of the mature protein-coding region of M7 lysin from allopatric populations of mussels in the Mytilus edulis species group (M. edulis, M. galloprovincialis, and M. trossulus). A significant McDonald-Kreitman test showed an excess of fixed amino acid replacing substitutions between species, consistent with positive selection. In addition, Kolmogorov-Smirnov tests showed significant heterogeneity in polymorphism to divergence ratios for both synonymous variation and combined synonymous and nonsynonymous variation within M. galloprovincialis. These results indicate that there has been adaptive evolution at M7 lysin and, furthermore, show that positive selection on sperm proteins can occur even when postzygotic reproductive isolation is incomplete.

Acrosome Reaction↗

Surname as 'cancer risk' in extreme isolates: example from the island of Lastovo, Croatia.

The aim of this study was to analyze whether there are surnames which appear more frequently among the ancestors of cancer cases in a small isolate, in comparison to the ancestral surnames of the healthy controls, using the classic case-control design. The chosen setting was the island of Lastovo, Croatia, located more than 100 kilometers from the nearest coastal region. The period of study was 1970-1995, during which a total of 76 cancer cases were recorded in a population of approximately 800. The comparison of surname frequencies was performed in current and in five ancestral generations. The leading hypothesis was that, if inbreeding and common ancestry contributed to the development of the disease, then those phenomena should be reflected in increasing frequency of some surnames among ancestors, identifying the 'hidden' consanguinity, or 'following' cancer-promoting genes on the Y-chromosome. The results imply that there are surnames representing a classic "risk" for cancer, but also those "protecting" from its development, which all underscores the importance of founder effect and genetic predisposition to the disease in a small, reproductively isolated population. All of the results become more evident and increasingly significant when analyzed in more distant ancestral generations.

Case-Control Studies↗

Interactions among quantitative traits in the course of sympatric speciation.

Sympatric speciation, the origin of two or more species from a single local population, has almost certainly been involved in formation of several species flocks, and may be fairly common in nature. The most straightforward scenario for sympatric speciation requires disruptive selection favouring two substantially different phenotypes, and consists of the evolution of reproductive isolation between them followed by the elimination of all intermediate phenotypes. Here we use the hypergeometric phenotypic model to show that sympatric speciation is possible even when fitness and mate choice depend on different quantitative traits, so that speciation must involve formation of covariance between these traits. The increase in the number of variable loci affecting fitness facilitates sympatric speciation, whereas the increase in the number of variable loci affecting mate choice has the opposite effect. These predictions may enable more cases of sympatric speciation to be identified.

Animals↗

Contrasting hybridization rates between sympatric three-spined sticklebacks highlight the fragility of reproductive barriers between evolutionarily young species.

Three-spined sticklebacks (Gasterosteus aculeatus) are a powerful evolutionary model system due to the rapid and repeated phenotypic divergence of freshwater forms from a marine ancestor throughout the Northern Hemisphere. Many of these recently derived populations are found in overlapping habitats, yet are reproductively isolated from each other. This scenario provides excellent opportunities to investigate the mechanisms driving speciation in natural populations. Genetically distinguishing between such recently derived species, however, can create difficulties in exploring the ecological and genetic factors defining species boundaries, an essential component to our understanding of speciation. We overcame these limitations and increased the power of analyses by selecting highly discriminatory markers from the battery of genetic markers now available. Using species diagnostic molecular profiles, we quantified levels of hybridization and introgression within three sympatric species pairs of three-spined stickleback. Sticklebacks within Priest and Paxton lakes exhibit a low level of natural hybridization and provide support for the role of reinforcement in maintaining distinct species in sympatry. In contrast, our study provides further evidence for a continued breakdown of the Enos Lake species pair into a hybrid swarm, with biased introgression of the 'limnetic' species into that of the 'benthic'; a situation that highlights the delicate balance between persistence and breakdown of reproductive barriers between young species. A similar strategy utilizing the stickleback microsatellite resource can also be applied to answer an array of biological questions in other species' pair systems in this geographically widespread and phenotypically diverse model organism.

Animals↗

Comparison of the evolutionary dynamics of symbiotic and housekeeping loci: a case for the genetic coherence of rhizobial lineages.

In prokaryotes, lateral gene transfer across chromosomal lineages may be mediated by plasmids, phages, transposable elements, and other accessory DNA elements. However, the importance of such transfer and the evolutionary forces that may restrict gene exchange remain largely unexplored in native settings. In this study, tests of phylogenetic congruence are employed to explore the range of horizontal transfer of symbiotic (sym) loci among distinct chromosomal lineages of native rhizobia, the nitrogen-fixing symbiont of legumes. Rhizobial strains isolated from nodules of several host plant genera were sequenced at three loci: symbiotic nodulation genes (nodB and nodC), the chromosomal housekeeping locus glutamine synthetase II (GSII), and a portion of the 16S rRNA gene. Molecular phylogenetic analysis shows that each locus generally subdivides strains into the same major groups, which correspond to the genera Rhizobium, Sinorhizobium, and Mesorhizobium. This broad phylogenetic congruence indicates a lack of lateral transfer across major chromosomal subdivisions, and it contrasts with previous studies of agricultural populations showing broad transfer of sym loci across divergent chromosomal lineages. A general correspondence of the three rhizobial genera with major legume groups suggests that host plant associations may be important in the differentiation of rhizobial nod and chromosomal loci and may restrict lateral transfer among strains. The second major result is a significant incongruence of nod and GSII phylogenies within rhizobial subdivisions, which strongly suggests horizontal transfer of nod genes among congenerics. This combined evidence for lateral gene transfer within, but not between, genetic subdivisions supports the view that rhizobial genera are "reproductively isolated" and diverge independently. Differences across rhizobial genera in the specificity of host associations imply that the evolutionary dynamics of the symbiosis vary considerably across lineages in native settings.

Base Sequence↗