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The heterochromatin of grasshoppers from the Caledia captiva species complex. I. Sequence evolution and conservation in a highly repeated DNA family.

The restriction enzyme TaqI digests 0.2% of the genomic DNA from the grasshopper Caledia captiva to a family of sequences 168 bp in length (length of consensus sequence). The sequence variation of this "Taq family" of repeat units was examined among four races from C. captiva to assay the pattern of evolution within this highly repeated DNA. The Taq-family repeats are located in C-banded heterochromatin on at least one member of each homologous pair of chromosomes; the locations range from centromeric to telomeric. Thirty-nine cloned repeats isolated from two population 1A individuals along with 11 clones from seven populations taken from three of the races demonstrated sequence variation at 72 positions. Pairwise comparisons of the cloned repeats, both within an individual and between different races, indicate that levels of intraspecific divergence, as measured by reproductive incompatibility, do not correlate with sequence divergence among the 168-bp repeats. A number of subsequences within the repeat remain unchanged among all 50 clones; the longest of these is 18 bp. That the same 18-bp subsequence is present in all clones examined is a finding that departs significantly (P less than 0.01) from what would be expected to occur at random. Two other cloned repeats, from a reproductively isolated race of C. captiva, have sequences that show 56% identity with this 18-bp conserved region. An analysis showed that the frequency of occurrence of an RsaI recognition site within the 168-bp repeat in the entire Taq family agreed with that found in the cloned sequences. These data, along with a partial sequence for the entire Taq family obtained by sequencing uncloned repeats, suggest that the consensus sequence from the cloned copies is representative of this highly repeated family and is not a biased sample resulting from the cloning procedure. The 18-bp conserved sequence is part of a 42-bp sequence that possesses dyad symmetry typical of protein-binding sites. We speculate that this may be significant in the evolution of the Taq family of sequences.

Animals↗

Crossing relationships among ancient and experimental sunflower hybrid lineages.

Reproductive barrier formation between newly derived hybrid taxa and their parental species represents a major evolutionary hurdle. Here, I examine the development of a sterility barrier during hybrid speciation by examining the fertility of progeny from all combinations of crosses involving three experimentally synthesized sunflower hybrid lineages, their natural hybrid counterpart, Helianthus anomalus, and their parents, H. annuus and H. petiolaris. Crosses between the parental species and H. anomalus generated almost completely sterile offspring (pollen viability < 5%; seed set < 1%). A fairly strong sterility barrier also has developed between three hybrid lineages and both parental species (pollen viability 11.1-41.6%; seed set 0.84-20.1%). In contrast, the three hybrid lineages are almost fully interfertile (pollen viabilities 83.1-88.6%; seed set 72.1-75.3%), as predicted by molecular mapping studies that indicate they have converged on a similar set of gene combinations and chromosomal rearrangements. A modest decline in compability is observed in crosses between the three hybrid lineages and H. anomalus (pollen viabilities 64.1-70.7%; seed set 37-43%), a result that agrees well with prior data demonstrating significant congruence between the genomes of the natural and experimental hybrid lineages. These observations not only indicate that reproductive isolation can arise as a by-product of fertility selection in hybrid populations, but also testify to the repeatability of this mode of speciation.

Breeding↗

Population structure of Hydropsyche exocellata. Genetic homogeneity in a zone of fragmented distribution.

Using four allozymic loci, we analysed the genetic structure and differentiation of Hydropsyche exocellata (Trichoptera; Hydropsychidae) larvae in seven sites along the Upper Loire River (France). The genetic differentiation is low for this species despite distribution patchiness, and only due to one locus in one study site. No trend relating genetic differentiation to geographic distance (isolation by distance) was observed. Significant deviations from Hardy-Weinberg equilibrium were observed at most sites and loci. The population genetics analysis of an amphibiotic insect such as H. exocellata in light of current ecological and population genetics knowledge is discussed following two main factors: 1) sampling of partially reproductively isolated, genetically distinct subpopulations, and 2) form of selection. Despite knowledge on several aspects of the ecology of H. exocellata in the River Loire, numerous points limit genetic data interpretation. These points are underlined and discussed with reference to studies on other amphibiotic insects.

Animals↗

Parasitic exploitation as an engine of diversity.

Parasitic exploitation occurs within and between a wide variety of taxa in a plethora of diverse contexts. Theoretical and empirical analyses indicate that parasitic exploitation can generate substantial genetic and phenotypic polymorphism within species. Under some circumstances, parasitic exploitation may also be an important factor causing reproductive isolation and promoting speciation. Here we review research relevant to the relationship between parasitic exploitation, within species-polymorphism, and speciation in some of the major arenas in which such exploitation has been studied. This includes research on the vertebrate major histocompatibility loci, plant-pathogen interactions, the evolution of sexual reproduction, intragenomic conflict, sexual conflict, kin mimicry and social parasitism, tropical forest diversity and the evolution of language. We conclude by discussing some of the issues raised by comparing the effect of parasitic exploitation on polymorphism and speciation in different contexts.

Animals↗

Olfactory discrimination as an isolating mechanism in speciating mole rats.

Olfactory discrimination was tested in two chromosome forms of the speciating fossorial rodent, Spalax ehrenbergi, in Israel. Females of the chromosome forms 2n = 52 and 2n = 58 were tested for male odour discrimination, the source of odour being either case litter or urine. Estrous females of both forms preferred homochromosomal odurs, whereas diestrous females showed no discrimination. These results suggest that olfactory discrimination may serve as a reproductive isolating mechanism in the speciation of mole rats.

Animals↗

On the origin of species by sympatric speciation.

Understanding speciation is a fundamental biological problem. It is believed that many species originated through allopatric divergence, where new species arise from geographically isolated populations of the same ancestral species. In contrast, the possibility of sympatric speciation (in which new species arise without geographical isolation) has often been dismissed, partly because of theoretical difficulties. Most previous models analysing sympatric speciation concentrated on particular aspects of the problem while neglecting others. Here we present a model that integrates a novel combination of different features and show that sympatric speciation is a likely outcome of competition for resources. We use multilocus genetics to describe sexual reproduction in an individual-based model, and we consider the evolution of assortative mating (where individuals mate preferentially with like individuals) depending either on an ecological character affecting resource use or on a selectively neutral marker trait. In both cases, evolution of assortative mating often leads to reproductive isolation between ecologically diverging subpopulations. When assortative mating depends on a marker trait, and is therefore not directly linked to resource competition, speciation occurs when genetic drift breaks the linkage equilibrium between the marker and the ecological trait. Our theory conforms well with mounting empirical evidence for the sympatric origin of many species.

Animals↗

Do Wolbachia infections play a role in unidirectional incompatibilities in a field cricket hybrid zone?

Two closely related field crickets, Gryllus firmus and G. pennsylvanicus, hybridize along an extensive north-south zone in the eastern United States. Crosses between G. firmus males and G. pennsylvanicus females produce viable and fertile F1, but the reciprocal cross consistently fails to produce offspring. Wolbachia, a bacterial parasite of arthropods that causes unidirectional incompatibilities in a variety of insect species, has been suggested as the cause of the observed incompatibility between G. pennsylvanicus and G. firmus. We examine the presence/absence of Wolbachia strains, defined by sequencing the ftsZ gene, in four cricket populations from the north-eastern United States. Most G. firmus individuals are infected (100% in Guilford, Connecticut; 65% in Seaside Park, New Jersey) and > 95% of those infected harbour a single strain of Wolbachia. All individuals in G. pennsylvanicus populations (Ithaca, New York; Sharon, Connecticut) are infected; the majority of individuals carry a second strain of Wolbachia, but a significant fraction carry the same strain found commonly in G. firmus. The presence of an apparently identical Wolbachia strain in crickets of both species means that some crosses between G. pennsylvanicus males and G. firmus females should be compatible. We have no evidence of such compatibility. Furthermore, if Wolbachia infections are responsible for the observed incompatibility between species, then incompatibilities must also exist within G. pennsylvanicus, because this species harbours both Wolbachia strains. Although some single pair crosses within G. pennsylvanicus do fail to produce offspring, the proportion is lower than expected if Wolbachia were responsible. Therefore, Wolbachia is unlikely to be involved in reproductive isolation between the two cricket species.

Animals↗

Isozyme Variation among Biological Species in the Gibberella fujikuroi Species Complex (Fusarium Section Liseola).

Isozyme phenotypes were determined for 101 strains of Gibberella fujikuroi and 2 strains of Gibberella nygamai that represent seven biological species (mating populations) isolated from a variety of plant hosts in dispersed geographic locations. Fourteen enzymes were resolved in one or more of three buffer systems. Two of the enzymes, arylesterase and acid phosphatase, were polymorphic within two or more biological species and are suitable for intraspecific studies of population variation. Six enzymes, alcohol dehydrogenase, aspartate aminotransferase, glucose-6-phosphate dehydrogenase, mannitol dehydrogenase, phosphoglucomutase, and phosphogluconate dehydrogenase, were monomorphic in all of the isolates examined. The remaining six enzymes, fumarase, glucose phosphate isomerase, glutamate dehydrogenase (NADP), isocitrate dehydrogenase (NADP), malate dehydrogenase, and triose-phosphate isomerase, could potentially be used to distinguish the different biological species. Mating populations C and D are the most similar, since the mating population C isolates examined had the same isozyme phenotype as did a subset of the isolates in mating population D. Mating population E is the least similar to the other taxa examined. Unique isozyme phenotypes are present but are composed of banding patterns shared among the biological species. This finding supports the hypothesis that these biological species, with the possible exception of mating populations C and D, are reproductively isolated from one another and that no significant gene flow is occurring between them. Isozyme analysis is a useful method to distinguish these closely related biological species. Examination of isozyme phenotypes is more rapid than the present technique, which is based on sexual crosses; can be applied to strains that are not sexually fertile; and is more sensitive than traditional morphological characters, which cannot distinguish more than three or four morphological groups among the seven biological species. While emphasizing the discreteness of the mating populations as biological entities, our isozyme data also reaffirm the close genetic relationship among these groups.

Journal Article↗

Population genetic differentiation in taxa of Lotus (Fabaceae: Loteae) endemic to the Gran Canarian pine forest.

A survey of allozyme variation at 17 loci in 14 populations representing four taxonomically problematic Gran Canarian pine forest endemic taxa of Lotus (L. genistoides, L. holosericeus, L. spartioides and some taxonomically uncertain populations collected under the designation Lotus sp.) was conducted to examine their diversification and systematic relationships. All groups exhibited high values of genetic variation, although inbreeding was common within populations. Considerable among-population genetic homogeneity was detected, as inferred from low values of Gst within each of the groups. The high population sizes of these taxa and a lack of evidence for isolation by distance or genetic bottlenecks indicate that diversity has accumulated over a long period of environmental stability. The association of high genetic distances with low linear distances, and the substantial increase in the values of Gst when the taxa considered were merged in different combinations hint at an incipient (yet probably taxonomically insufficient) reproductive isolation. The genetic similarity between L. genistoides, L. holosericeus and L. spartioides, together with the different behaviour of the populations collected under the designation Lotus sp., may have important implications for the restructuring of the taxonomy of this group when the ongoing morphological studies are completed.

Atlantic Islands↗

Kodamaea kakaduensis and Candida tolerans, two new ascomycetous yeast species from Australian Hibiscus flowers.

Two new yeast species were isolated from flowers of Hibiscus species in Eastern and Northern Australia. Kodamaea kakaduensis is heterothallic, haploid, and similar to other Kodamaea species and to Candida restingae. Buds are often produced on short protuberances, and a true mycelium is formed. The new species differs from others by the assimilation of trehalose, melezitose, and xylitol, and is reproductively isolated. The cells of Candida tolerans are small and a pseudomycelium is formed. The carbon and nitrogen assimilation pattern is reminiscent of that of Zygosaccharomyces rouxii but the two are not closely related. Sequences of the D1/D2 domain of large subunit ribosomal DNA confirm the membership of K. kakaduensis in the genus Kodamaea and indicate that C. tolerans belongs to the Clavispora-Metschnikowia clade, with a moderate relatedness to Candida mogii. The type strains are: K. kakaduensis, UWO(PS)98-119.2 (h+, holotype, CBS 8611) and UWO(PS)98-117.1 (h-, isotype, CBS 8612); and C. tolerans, UWO(PS)98-115.5 (CBS 8613).

Ascomycota↗

Comparative biology of two populations of Lutzomyia umbratilis (Diptera: Psychodidae) of Central Amazonia, Brazil, under laboratory conditions.

Lutzomyia umbratilis is the main vector of cutaneous leishmaniasis due to Leishmania guyanensis in northern South America. It has been found naturally infected with this species of Leishmania only east of the Rio Negro and north of the Rio Amazonas. However, populations of this sand fly species are also present in areas south of the Amazon river system, which may act as a geographical barrier to the Leishmania guyanensis cycle. With the aim of looking for possible biological differences between populations of L. umbratilis from each side of this river system, their biology in the laboratory was investigated. Progenitors collected on tree bases in Manaus and Manacapuru (east and west, respectively, of the Rio Negro) were reared in the laboratory. Results from observations of the life cycle, fecundity, fertility, and adult longevity at 27 degrees C and 92% RH were analyzed by descriptive statistics and z, t, U, and chi2 tests. Although the Manaus and Manacapuru colonies showed a longer developmental time than most Lutzomyia species reared at similar temperatures, length of time of egg and 4th instar larva of the two populations differed significantly (p < 0.01). Females of the latter retained significantly (p < 0.001) less mature oocytes, and the general productivity (% adults from a known number of eggs) of the colony was significantly (p < 0.01) higher than that of the former. These results show that the L. umbratilis population of Manaus is more productive, and thus a better candidate for future mass-rearing attempts. The two populations differ in their life cycle, fecundity, fertility, adult longevity, and emergence. These differences may reflect some divergence of intrinsic biological features evolved as a result of their geographical isolation by the Rio Negro. It is expected that further investigations on morphometry, cuticular hydrocarbon, isoenzyme, molecular and chromossomal analyses, infection, and cross-mating experiments with these and other allopatric populations of both margins of the Amazon river system will help reveal whether or not L. umbratilis has genetically diverged into two or more reproductively isolated populations of vectors or non-vectors of Leishmania guyanensis.

Animals↗

Wolbachia and reproductive conflict in Exorista sorbillans.

Many arthropods harbour endosymbiotic bacteria of the genus Wolbachia. These endosymbionts are transmitted vertically from one generation to the next and are obligatory in several Dipterans that have been studied to date. These bacteria induce an array of reproductive isolation mechanisms that are implicated in pest management to evolutionary biology of respective hosts. The uzifly, Exorista sorbillans, a tachinid endoparasitoid of the silkworm, Bombyx mori L. (Lepidoptera: Bombycidae), causes enormous losses to the silk industry; now it is known that it harbours Wolbachia endobacteria. The elimination of Wolbachia by antibiotics interrupts embryogenesis and causes various reproductive conflicts such as (1) a reduction of fecundity of uninfected female, (2) cytoplasmic incompatibility in the uninfected females crossed with infected males, (3) genomic incompatibility in crosses between males and females from uninfected population, and (4) sex-ratio distortion in uninfected females irrespective of the presence of Wolbachia in males. These results suggest that the relationship of Wolbachia with its uzifly host is one of mutual symbiosis as it controls the reproductive physiology of its host.

Analysis of Variance↗

Hidden evolution: progress and limitations in detecting multifarious natural selection.

From illustrative examples of research on the best-studied group of species to date, Drosophila melanogaster and its closest relatives, we argue that selection is multifarious, but often hidden. Selective fixation of new, highly advantageous alleles is the most parsimonious explanation for a typical pattern of molecular variation observed in genomic regions characterized by very low recombination: drastically reduced DNA sequence variation within species and typical levels of sequence divergence among species. At the same time, the identity of the gene (or genes) influenced by selection is not just difficult to discern; it may be impossible. Studies of the genetic basis of reproductive isolation demonstrate that, although the D. melanogaster complex species appear virtually identical, dozens of currently unidentified genes contribute to hybrid sterility. We argue that these findings are best explained by selectively-driven functional divergence and demonstrate the multifarious nature of selection. Although multifarious selection certainly occurs, the exact characters responsible for differences in survival and reproductive success are unknown. We do not see these inherent limits as a cause for despair or a problem for evolutionary biology. Instead, we hope to raise awareness of these complexities of evolution by highlighting both the progress and the limitations of characterizing multifarious natural selection.

Alcohol Dehydrogenase↗

Sexual isolation in Drosophila melanogaster: a possible case of incipient speciation.

It is generally believed that Drosophila melanogaster has no closely related species with which it can produce the viable and fertile hybrids that are essential for the genetic analysis of speciation. Following the recent report of molecular differentiation between a Zimbabwe, Africa, population and two United States populations, we provide evidence that strong sexual isolation exists between the D. melanogaster population in Zimbabwe and populations of other continents. In the presence of males of their own kind, females from most isofemale lines of Zimbabwe would not mate with males from elsewhere; the reciprocal mating is also significantly reduced, but to a lesser degree. The genes for sexual behaviors are apparently polymorphic in Zimbabwe and postmating reproductive isolation between this and other populations has not yet evolved. Whole chromosome substitutions indicate significant genetic contributions to male mating success by both major autosomes, whereas the X chromosome effect is too weak to measure. In addition, the relative mating success between hybrid and pure line males supports the interpretation of strong female choice. These observations suggest that we are seeing the early stages of speciation in this group and that it is driven by sexual selection. The genetic and molecular tractability of D. melanogaster offers great promise for the detailed analysis of this apparent case of incipient speciation.

Animals↗

Recent radiation of endemic Caribbean Drosophila of the dunni subgroup inferred from multilocus DNA sequence variation.

Studies of island endemism provide a unique opportunity to elucidate fundamental mechanisms of speciation. Here we examine intra- and interspecific DNA sequence variation at four unlinked genetic loci among populations of the Drosophila dunni subgroup to provide a detailed genealogical portrait of the process of speciation among these island endemic species. Our data indicate two major rounds of diversification that have shaped the D. dunni subgroup. The first occurred 1.6-2.6 million years ago and separated three major lineages, one in Puerto Rico and the Virgin Islands, a second in the northern Lesser Antilles and Barbados, and a third in St. Vincent and Grenada. A second round of diversification occurred in the last 96,000 years in the northern Lesser Antilles and Barbados. The four distinct species that resulted from this recent round of diversification maintain relatively high amounts of genetic variation, similar to that of a closely related mainland species, and share extensive ancestral polymorphism. These data suggest a minimal role for population bottlenecks linked to founder events in the history of the D. dunni subgroup. Further, the recent divergence of these island populations highlights the extremely rapid development of reproductive isolation and distinct patterns of abdominal pigmentation that has occurred in these species.

Animals↗

Experimental evolution of mating discrimination in budding yeast.

Assortative mating, when individuals of similar phenotypes mate, likely plays a key role in preventing gene flow during speciation. Reinforcement occurs when two previously geographically separated (allopatric) groups meet after having evolved partial postzygotic isolation; they are selected to evolve or enhance assortative mating to prevent costly intergroup matings that produce only maladaptive or sterile hybrids. Studies in Drosophila have shown that the genetic architectures of mating discrimination could differ significantly with or without reinforcement, suggesting that the evolution of assortative mating may be more complicated than expected. To study the evolution of assortative mating, we evolved mating discrimination in populations of the budding yeast, Saccharomyces cerevisiae. After 36 cycles of selection, these cells are five times more likely to mate with each other than to their ancestors, despite detectable one-way gene flow between the selected and reference populations. Several individual cultures evolved mating discrimination by changing their mating kinetics, with some mating more rapidly and others more slowly than the ancestral population. Genetic analysis indicates that multiple mutations have accumulated to produce the altered mating preference. Our results show that subtle details of mating behavior can play an important role in the evolution of reproductive isolation.

Biological Evolution↗

The Maynard Smith model of sympatric speciation.

The paper entitled "Sympatric speciation," which was published by John Maynard Smith in 1966, initiated the development of mathematical models aiming to identify the conditions for sympatric speciation. A part of that paper was devoted to a specific two-locus, two-allele model of sympatric speciation in a population occupying a two-niche system. Maynard Smith provided some initial numerical results on this model. Later, Dickinson and Antonovics (1973) and Caisse and Antonovics (1978) performed more extensive numerical studies on the model. Here, I report analytical results on the haploid version of the Maynard Smith model. I show how the conditions for sympatric and parapatric speciation and the levels of resulting genetic divergence and reproductive isolation are affected by the strength of disruptive selection and nonrandom mating, recombination rate, and the rates of male and female dispersal between the niches.

Animals↗

Leaky prezygotic isolation and porous genomes: rapid introgression of maternally inherited DNA.

Accurate phylogenies are crucial for understanding evolutionary processes, especially species diversification. It is commonly assumed that "good" species are sufficiently isolated genetically that gene genealogies represent accurate phylogenies. However, it is increasingly clear that good species may continue to exchange genetic material through hybridization (introgression). Many studies of closely related species reveal introgression of some genes without others, often with more rapid introgression of maternally inherited chloroplast or mitochondrial DNA (cpDNA, mtDNA). We seek a general explanation for this biased introgression using simple models of common reproductive isolating barriers (RIBs). We compare empirically informed models of prezygotic isolation (for pre- and postinsemination mechanisms of both female choice and male competition) with postzygotic isolation and demonstrate that rate of introgression depends critically upon type of RIB and mode of genetic inheritance (maternal versus biparental versus paternal). Our frequency-dependent prezygotic RIBs allow much more rapid introgression of biparentally and maternally inherited genes than do commonly modeled postzygotic RIBs (especially maternally inherited DNA). After considering the specific predictions in the context of empirical observations, we conclude that our model of prezygotie RIBs is a general explanation for biased introgression of maternally inherited genomic components. These findings suggest that we should use extreme caution when interpreting single gene genealogies as species phylogenies, especially for cpDNA and mtDNA.

Animals↗