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Isozymic differentiation of two sibling species of Squatina (chondrichthyes) in South Brazil.

Two sympatric morphotypes of the nominal species Squatina argentina from South Brazil are compared by the frequencies of 25 isozymes distributed over 14 presumptive loci. The Nei's Genetic Identity index obtained in a comparison of the populations is of 0.71. The loci EST-2 and EST-4 are fixed at different isozymes for each population, indicating a reproductive isolation between them. The most significant morphological and meristical differences between the morphotypes are the presence or absence of dorsal spines and the dental formula. The agreement between these characteristics and the genetical data indicates that the two morphotypes belong to different species.

Animals

A threshold model for punctuated gradualism.

A model for continuous and discontinuous evolutionary change is proposed that accommodates both punctuated equilibrium and phyletic gradualism. Natural selection operating on a gaussian distribution of phenotypes subjected to severe stress will result in a skewing of the normal distribution in the direction of the favoured phenotype. If a reproductive isolation threshold is imposed upon this model, rapid crossing of the threshold by large numbers of individuals within the skewed distribution will result in the sudden achievement of a critical descendant population mass. Mild directional selection however, will result in gentle shifts of the gaussian mean thus leading us to suggest that stasis and incipient speciatior are extremes of the same spectrum, defined by the rates of mean-shift in response to varying severities of selection pressure. Both punctuated and gradualistic evolution as accommodated in our model may further be described within Thom's cusp catastrophe theory.

Biological Evolution

Radiation of chromosome shuffles.

Rock wallabies, Petrogale, exhibit chromosome diversity that is exceptional in marsupials, with 20 distinct chromosome races being recognized. Many of the karyotypic changes identified within Petrogale appear to be recent, although the rate of chromosome evolution varies between taxa. While the patchy distribution of Petrogale and their social structure would facilitate the fixation of novel rearrangements, these factors alone do not explain the pattern of chromosome evolution shown in this group. The chromosome changes that have come to characterize each taxon may offer selective advantages in the particular areas occupied, or it may be that these rearrangements play an important role in reproductive isolation. In Petrogale, the taxa with the largest number of chromosome rearrangements are those that are sympatric, or have multiple zones of parapatry, with other members of the genus. Male hybrids from a variety of chromosomal admixtures were found to be sterile, but with those heterozygous for the least complex rearrangements being least affected. As expected, equivalent female hybrids were less severely affected. Chromosomal and genic changes both appear important in these processes.

Animals

De novo Genes in Plants: Origins, Mechanisms, and Functional Implications.

De novo genes originate from previously non-coding genomic regions. They provide an important source of lineage-specific innovation. In plants, these genes may contribute to adaptation, trait diversity and crop evolution. This review summarizes recent progress in plant de novo gene research. It first discusses major routes of gene birth, including transcription-first, open reading frame (ORF)-first and concurrent models. It also examines how nascent loci acquire regulatory control and enter existing biological networks. The review then summarizes their evolutionary features, including weak early constraint, rapid molecular change, restricted expression and structural refinement. It further discusses plant de novo genes involved in stress responses, seed germination, kernel dehydration, subspecies divergence, reproductive isolation and floral scent diversification. Current methods for identifying de novo genes remain limited by rapid sequence evolution, genome annotation quality, polyploidy and transposable elements. Whole-genome synteny alignment, multi-omics evidence and machine-learning approaches can improve candidate discovery. However, each method has important limitations. Finally, this review highlights key future questions in functional validation, latent coding potential in long non-coding RNAs, epigenetic activation, regulatory-network integration and crop improvement. These perspectives clarify how de novo genes shape plant adaptation and how they may be used in precision breeding and synthetic biology.

adaptive evolution

The Gibbons speciation mechanism.

A mechanism of sympatric speciation first proposed by Gibbons (1979, Am. Nat. 114, 719-741) is analyzed and submitted to computer simulation. It is found that, in its original form, the mechanism does not bring about reproductive isolation, but that with relatively minor modification, it may be made to work as claimed.

Animals

The inheritance of acquired epigenetic variations.

There is evidence that the functional history of a gene in one generation can influence its expression in the next. In somatic cells, changes in gene activity are frequently associated with changes in the pattern of methylation of the cytosines in DNA; these methylation patterns are stably inherited. Recent work suggests that information about patterns of methylation and other epigenetic states can also be transmitted from parents to offspring. This evidence is the basis of a model for the inheritance of acquired epigenetic variations. According to the model, an environmental stimulus can induce heritable chromatin modifications which are very specific and predictable, and might result in an adaptive response to the stimulus. This type of response probably has most significance for adaptive evolution in organisms such as fungi and plants, which lack distinct segregation of the soma and germ line. However, in all organisms, the accumulation of specific and random chromatin modifications in the germ line may be important in speciation, because these modifications could lead to reproductive isolation between populations. Heritable chromatin variations may also alter the frequency and distribution of classical mutations and meiotic recombination. Therefore, inherited epigenetic changes in the structure of chromatin can influence neo-Darwinian evolution as well as cause a type of "Lamarckian" inheritance.

Base Sequence

Allogeneic cellular reactions between intra-specific types of a solitary ascidian, Halocynthia roretzi.

Coelomic cells from a solitary ascidian, Halocynthia roretzi, exibit a nonphagocytic cellular reaction against coelomocytes from different species and another individuals of the same species. The reaction was denoted contact reaction. While xenogeneic contact reactions were always observed, allogeneic reactions were observed in most but not all combinations of individuals. Three variant types of H. roretzi (Type A, B, C) inhabit the coast of northern Japan. They are reproductively isolated under natural conditions. Non-reactive combinations exist between the different types, in all possible combinations. The paterns and frequencies of reactivities between different variants are almost the same to those observed within a single type. The results of the alloreactivity suggest that the three types of H. roretzi became separate from each other very recently and still remain intra-specific variants.

Alleles

Genic differentiation and origin of Robertsonian populations of the house mouse (Mus musculus domesticus Rutty).

This paper examines the relation between chromosomal and nuclear-gene divergence in 28 wild populations of the house mouse semi-species, Mus musculus domesticus, in Western Europe and North Africa. Besides describing the karyotypes of 15 of these populations and comparing them to those of 13 populations for which such information was already known, it reports the results of an electrophoretic survey of proteins encoded by 34 nuclear loci in all 28 populations. Karyotypic variation in this taxon involves only centric (or Robertsonian) fusions which often differ in arm combination and number between chromosomal races. The electrophoretic analysis showed that the amount of genic variation within Robertsonian (Rb) populations was similar to that for all-acrocentric populations, i.e. bearing the standard karyotype. Moreover, divergence between the two types of populations was extremely low. These results imply that centric fusions in mice have not modified either the level or the nature of genic variability. The genetic similarity between Rb and all-acrocentric populations is not attributed to the persistence of gene flow, since multiple fusions cause marked reproductive isolation. Rather, we attribute this extreme similarity to the very recent origin of chromosomal races in Europe. Furthermore, genic diversity measures suggest that geographically separated Rb populations have in situ and independent origins. Thus, Rb translocations are probably not unique events, but originated repeatedly. Two models are presented to explain how the rapid fixation of a series of chromosomal rearrangements can occur in a population without lowering variability in the nuclear genes. The first model assumes that chromosomal mutation rates are between 10(-3) and 10(-4) and that populations underwent a series of transient bottlenecks in which the effective population size did not fall below 35. In the second model, genic variability is restored following severe bottlenecks, through gene flow and recombination.

Animals

The divergence of a polygenic system subject to stabilizing selection, mutation and drift.

Polygenic variation can be maintained by a balance between mutation and stabilizing selection. When the alleles responsible for variation are rare, many classes of equilibria may be stable. The rate at which drift causes shifts between equilibria is investigated by integrating the gene frequency distribution W2N II (pq)4N mu-1. This integral can be found exactly, by numerical integration, or can be approximated by assuming that the full distribution of allele frequencies is approximately Gaussian. These methods are checked against simulations. Over a wide range of population sizes, drift will keep the population near an equilibrium which minimizes the genetic variance and the deviation from the selective optimum. Shifts between equilibria in this class occur at an appreciable rate if the product of population size and selection on each locus is small (Ns alpha 2 less than 10). The Gaussian approximation is accurate even when the underlying distribution is strongly skewed. Reproductive isolation evolves as populations shift to new combinations of alleles: however, this process is slow, approaching the neutral rate (approximately mu) in small populations.

Alleles

Mapping and characterization of a 'speciation gene' in Drosophila.

Almost nothing is known about the identity of the genes causing reproductive isolation between species. As a first step towards molecular isolation of a 'speciation gene', I mapped and partly characterized a gene causing hybrid male sterility in Drosophila. This analysis shows that sterility of D. melanogaster males who carry the 'dot' fourth chromosome from D. simulans is due entirely to a very small region of the D. simulans chromosome (including only about 5 salivary gland bands or approximately 250 kb of DNA). Thus the hybrid sterility effect of the D. simulans fourth chromosome is almost surely due to a single gene of very large effect (here named hms, hybrid male sterile). Hms is zygotically acting, and the D. simulans allele of hms is completely recessive. Furthermore, complementation tests suggest that hms is not an allele of any known locus in D. melanogaster.

Alleles

The genetic basis of Haldane's rule.

'Haldane's rule', formulated by J. B. S. Haldane in 1922, states that: "When in the F1 offspring of two different animal races one sex is absent, rare, or sterile, that sex is the heterozygous [heterogametic] sex". His rule is now known to apply in mammals, lepidopterans, birds, orthopterans and dipterans. In Drosophila, for example, Bock cites 142 cases of interspecific hybridizations that produce one sterile and one fertile sex in the offspring, all but one of these crosses yielding sterile XY males and fertile XX females. Despite much speculation, however, the genetic basis of Haldane's rule remains unknown. Haldane himself rejected the simple explanation that males are innately more sensitive than females to the effects of hybridization because groups with heterogametic females (such as birds and butterflies) usually show female sterility in hybrids, so that heterogamety itself is the critical feature. He and others suggested that heterogametic infertility or inviability in hybrids arises by a genetic imbalance between X chromosomes and autosomes. An alternative explanation is that this syndrome is caused by a mismatch of X and Y chromosomes. Here I show that in the Drosophila melanogaster subgroup, Haldane's rule for fertility apparently arises from a genetic interaction between X and Y chromosomes and not from an imbalance between sex chromosomes and autosomes. This finding has important implications for understanding the evolution of interspecific reproductive isolation.

Animals

Bidirectional incompatibility between conspecific populations of Drosophila simulans.

Cytoplasmic incompatibility (CI) describes the phenomenon whereby eggs fertilized by sperm from insects infected with a rickettsial endosymbiont fail to hatch. Unidirectional CI between conspecific populations of insects is a well documented phenomenon. Bidirectional CI has, however, only been described in mosquito populations, and recently between closely related species of parasitic wasps, where it is of interest as both an unusual form of reproductive isolation and as a potential means of insect population suppression. Here we report on the first known example of bidirectional CI between conspecific populations of Drosophila simulans. Further, we show that defects as early as the first cleavage division are associated with CI. This observation suggests that the cellular basis of CI involves disruption of processes before or during zygote formation and that CI arises from defects in the structure and/or function of the sperm during fertilization.

Animals

Location of an X-linked factor causing sterility in male hybrids of Drosophila simulans and D. mauritiana.

We report the first mapping of a genetic factor responsible for reproductive isolation: a small segment of genome strongly affecting sperm motility in hybrids between the sibling species Drosophila simulans and D. mauritiana. Maximum-likelihood analysis of data from ten generations of backcrossing places this factor at 1.1 +/- 0.2 map units from the forked locus, at position 54.9 +/- 0.2 or 57.1 +/- 0.2 on the X chromosome.

Animals

Genetic analysis of X-linked sterility in hybrids between three sibling species of Drosophila.

Three morphological markers (yellow, miniature, and forked) are used to map the location of X-chromosome segments causing male sterility in Drosophila simulans/D. mauritiana and D. simulans/D. sechellia hybrids. In both hybridizations at least three sections of the chromosome contain genes with substantial effects on sterility. This represents the maximum genetic divergence detectable with the three markers, suggesting that the X chromosome contains many loci affecting postzygotic reproductive isolation. The tight linkage between some markers and "sterility loci" may be useful in localizing and later cloning genes important in speciation.

Animals

Fertility estimates in the Tunisian all-acrocentric and Robertsonian populations of the house mouse and their chromosomal hybrids.

The reproductive features of wild all-acrocentric and 2n = 22 Robertsonian (Rb) house mice (M. m. domesticus) from Tunisia were studied. The aim was to examine the possibility of a reproductive selective advantage associated with chromosomal change as well as to measure the effect of heterozygosity for a large number of Rb fusions on the fertility of hybrids. Results showed that litter sizes were significantly smaller in Rb than in all-acrocentric mice. This difference, which may represent a favourable demographic strategy related to the habitat segregation observed in the Tunisian mice, needs to be studied further. The F1 hybrids between the two chromosomal races showed a significantly reduced reproductive success and litter size (respectively, 53 per cent and 60 per cent less than either parental race). Analysis of the testicular histology of F1 and backcross males showed in some cases a breakdown of spermatogenesis. The degree of this disturbance was not related to the level of chromosomal heterozygosity suggesting that genetic incompatibilities between the two genomes might be involved. The strong reduction in fertility measured in these hybrids represents a reproductive isolating mechanism effectively reducing gene flow between the all-acrocentric and 22Rb mice populations of Tunisia.

Analysis of Variance

Genetic variability of the interpulse interval of courtship song among some European populations of Drosophila melanogaster.

The interpulse interval of the courtship song of Drosophila melanogaster is a character which may play a significant role in mating success and reproductive isolation. Here we examine the variability of interpulse interval among replicated laboratory strains of D. melanogaster. There is no significant variation among populations of different geographical origin. This suggests that interpulse interval is subject to strong selection, as the populations are known to differ for other characters. One population, however, was sufficiently different to allow a genetic analysis. Reciprocal F1s and backcrosses implied that the variance was predominantly additive and autosomal. Possible sources of selection on interpulse interval are discussed.

Animals

Mitochondrial DNA differentiation among geographical populations of Pronolagus rupestris, Smith's red rock rabbit (Mammalia: Lagomorpha).

Geographical genetic population structure was determined for an endemic African leporid, Smith's red rock rabbit, Pronolagus rupestris. Restriction fragment length polymorphism analysis of mitochondrial DNA from 55 specimens revealed 32 distinct material lineages for the population sampled. The data show two major genetic assemblages separated by a mean sequence divergence of 7.94 per cent (+/- 1.40 per cent) and provide little support for the continued recognition of most of the described subspecies. The south-eastern assemblage is confined to the mountain ranges comprising the Great Escarpment of South Africa, while the north-western assemblage is not so tightly constrained. With the possible exception of elevation, no readily apparent ecological or topographical barrier could be identified which delimits the two mitochondrial clades. The sequence divergence separating the south-eastern and north western P. rupestris clades is high, and approximates the interspecific sequence divergences detected between P. rupestris and other Pronolagus species. The two P. rupestris clades are parapatric for part of their distribution, and the absence of shared mtDNA lineages is consistent with the hypothesis that the two populations are reproductively isolated from each other. We provisionally interpret this to reflect the presence of two hitherto undetected biological species in what has conventionally been recognized as a single taxon, P. rupestris.

Animals

[Original adaptive characters of intestinal Digenea of Sarpa salpa (Teleostei, Sparidae) and their interpretation in terms of evolution].

In the family Sparidae, the genus Sarpa is distinguished by a few characteristics: monospecificity, vegetarian diet and wide geographical distribution. The helminth fauna of Sarpa salpa is also very original. Indeed, the digenean parasites of this Teleostean fish are essentially classified into two families restricted to this fish. In the present paper, the author redescribes Mesometra orbicularis, M. brachycoelia, Centroderma spinosissima, Elstia stossichianum, Wardula capitellata (family Mesometridae) together with Robphildollfusium fractum (family Robphildollfusidae). Various original and yet unknown features are pointed out. Among these unusual structures, several correspond to adaptive characteristics favouring the settlement of the Digenean on the peculiar digestive gut wall of this herbivorous fish. Indeed, the intestinal mucous membrane of Sarpa salpa exhibits very few villi giving it an unusual smooth aspect. Therefore, the Mesometridae which always have just a single sucker (monostomatous) have selected a new kind of compensatory adhesive structure. Sometimes, the anterior end of the body becomes a sucker due to the particular distribution of the muscle strings; in other examples, the whole body becomes a sucker and its edges become considerably thinner to improve the tightness of the adhesive system. Other original anatomical features have been selected to allow survival in a medium rich in plant detritus. So, in the oral sucker crests ornemented by numerous sclerous denticles seem to act as a microfilter for the intestinal chyme in which plant fibres predominate. The original pharynx seems to act as a suction-force pump. The excretory system, which is of a reticular type, penetrates the whole parenchyma and this could be a response to huge intestinal fermentations. The Digenea of Sarpa salpa are not interpreted by the author as true parasites but as endocommensal symbionts. These inquiline species are not immunogenic, or at least only slightly so, since they do not feed upon the host itself but upon its intestinal chyme. In most cases this results in a high parasite density (post larvae and adults) together with a cohabitation of the various species along the various intestinal segments. Coexistence of several species, systematically very close, evidently raises the question of their reproductive isolation. The author proposes an answer founded upon data of allopatric speciation.

Adaptation, Physiological