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The last decade has brought renewed interest in the genetics of speciation, yielding a number of new models and empirical results. Defining speciation as 'the origin of reproductive isolation between two taxa', we review recent theoretical studies and relevant data, emphasizing the regular patterns seen among genetic analyses. Finally, we point out some important and tractable questions about speciation that have been neglected.
Restriction fragment length polymorphisms (RFLPs) of two major histocompatibility class II genes (P alpha 1 and Q beta) were studied in 13 populations of four chromosomal species (2n = 52, 54, 58 and 60) of the mole rat, Spalax ehrenbergi superspecies in Israel. A substantial frequency of allelic fragments was found in both genes for all populations, including a desert isolate. In the P alpha 1 gene, one allelic fragment is a result of a deletion mutation which is diagnostic of the 2n = 52 chromosomal species. All other ten allelic variants are the result of point mutations. All mutations are located in a short region flanking the 3' end of the gene. Based on Mhc polymorphisms we confirm earlier evidence that gene flow does not occur between the older chromosomal species (2n = 52, 54, 58), and that reproductive isolation decreases, progressively from the oldest to the youngest species (2n = 60).
Changes in isolation, inbreeding, population subdivision, and isonymous relationships are examined in six Quechua communities from the upper valley of the Ichu River in the Peruvian Central Andes (3700 m). All marriages registered between 1825 and 1914 in the Parish of Santa Ana were analyzed. The data (1680 marriages) were divided into 2 periods (1825-1870 and 1871-1914) and into the 6 villages that constitute the parish. Endogamy rates are between 81% and 100%, indicating high levels of reproductive isolation. The inbreeding indicated by isonymy (Ft, Fr, and Fn) is lower than in other mountain populations studied. Isonymy values, calculated from the different surname combinations made possible by the Ibero-American Surnames System, indicate a strong rejection of consanguineous marriages, particularly between patrilineal relatives, in agreement with the parental structure typical of Andean populations. The comparison between observed and expected repeated-pair values reveals a moderate level of subdivision within populations, which could be related to cultural and socioeconomic factors. Nonmetric multidimensional scaling was used to investigate temporal changes in the isonymous relationships among the communities. The results reveal a decrease in the interpopulational variability measured by surnames, in agreement with an increase in exogamy. Surnames and data contained in historical and demographic records yield reliable information, and they can be used to reconstruct the biological history of Amerindian populations over the last few centuries.
Porcine reproductive and respiratory syndrome (PRRS) virus was isolated at high frequency from the sera and lungs of pigs affected with Heko-Heko disease. In addition, a considerable amount of Mycoplasma hyorhinis (Mhr) was also isolated from the lungs. Inoculation of gnotobiote pigs with the first isolate of PRRS virus resulted in the reproduction of proliferative and interstitial pneumonia. The virus was recovered from the inoculated pigs for long periods. Superinfection with PRRS virus and Mhr appeared to produce more serious pneumonia than inoculation with PRRS virus alone. In this study, the presence of PRRS virus was confirmed in Japan, and the PRRS virus was considered as being the most important pathogen for Heko-Heko disease.
Patterns of reproductive isolation between two sympatric species of oaks, Quercus gambelii and Q. grisea, that exhibit strong ecological differentiation were examined. A full diallel cross using four trees of each species (i.e. all possible pollinations among eight trees) was performed. This design was repeated at two sites that represent different outcomes of sympatry: (1) a xeric mountain ridge where many hybrids are established (HZ); and (2) a mesic valley bottom where virtually no hybrids are established (MOCYN). By measuring fruit survival at several developmental stages, both the timing and strength of reproductive barriers within and between sites, species, cross types, and pollen dosage levels were examined. In three of four cases, heterospecific fruit set was significantly reduced compared to conspecific fruit set. This reduction occurred after the time of fertilization, but before the onset of embryo growth. Increasing the dose of pollen from an average of 9-194 grains/stigma did not affect this result. Thus, early postfertilization processes play a strong role in species fidelity in these oaks. Quercus gambelii experienced a five-fold decrease in conspecific fruit set at HZ relative to MOCYN. In contrast, heterospecific fruit set of Q. gambelii was the same at both sites. Poor Q. gambelii pollen performance is implicated as playing the major role in this result. One Q. gambelii individual at HZ was highly fecund, and had higher heterospecific than conspecific fruit set; slight introgression in this tree was detected uisng RAPD markers. The Environmental Emasculation Hypothesis that posits that environmental stress can increase the probability of hybrid formation by reducing the competitive ability of male gametes of one species is proposed.
A diverse array of sessile marine invertebrates mate by passive dispersal of sperm which fertilize the brooded eggs of neighbours. In two such species, a sea-mat (phylum Bryozoa) and an ascidian (phylum Chordata), vitellogenic egg growth is absent in reproductively isolated specimens, but is triggered by a water-borne factor released by conspecifics. In both of these colonial, hermaphroditic species, the active factor can be removed from water by filtration. The effect involves self-/non-self-recognition: water conditioned by a separate subcolony of the same genetic individual does not prompt oocyte growth. In each species, allosperm move from the surrounding water to the ovary and are then stored in close association with the growing oocytes. We concluded that sperm themselves are the water-borne factor that triggers the major phase of female reproductive investment. This mechanism is, to our knowledge, previously undescribed in animals, but has parallels with the initiation of maternal investment in flowering plants following the receipt of compatible pollen. The species studied may be representative of many other aquatic invertebrates which mate in a similar way. The stimulation of egg growth by allosperm could lead to intersexual conflict during oogenesis.
A method was developed for gene fingerprinting, combining the principles of restriction fragment length polymorphism and single-strand conformational polymorphism (PCR-RFLP-SSCP). Taking advantage of this method, we analysed the genotypes of 20 isolates from five species of Trichinella (Trichinella spiralis, Trichinella britovi, Trichinella nativa, Trichinella murrelli and Trichinella pseudospiralis) and two uncertain genotypes (Trichinella T6 and Trichinella T8). Target genes for the analysis included three kinds of random amplified polymorphic DNA (RAPD), the gene encoding 43 kDa excretory-secretory protein, the gene encoding mitochondrial cytochrome c -oxidase subunit I and the gene of 18 S rRNA. The genotype revealed by this method was in good concordance with the taxonomy of Trichinella (six species and two uncertain genotypes which is currently accepted based on morphology, isozyme pattern and reproductive isolation). More important, this method revealed intraspecies polymorphism among geographical isolates of T. spiralis and T. britovi.
The endemic New Zealand Hector's dolphin is considered the rarest species of marine dolphin with a total abundance of less than 4000. The species is listed as vulnerable because of fisheries-related mortality due to entanglement in set nets. The vulnerability of this species is further increased by its fidelity to local natal ranges and the genetic isolation of regional populations. Here we present evidence, based on 108 contemporary samples and 55 historical samples dating back to 1870, of a significant loss of mitochondrial DNA (mtDNA) diversity in two regional populations of Hector's dolphin. The haplotype diversity (h) was calculated from sequences of a 206 bp fragment in the mtDNA control region, designed to identify 13 out of the 14 known maternal lineages. Over the last 20 years, the North Island population has been reduced from at least three lineages (h = 0.41) to a single lineage (h = 0; p < 0.05). Given its small size, reproductive isolation and reduced genetic diversity, this population is likely to become extinct. The diversity of the East Coast South Island population has declined significantly from h = 0.65 to h = 0.35 (p < 0.05). Based on trend analysis of the mtDNA diversity, we predict that the East Coast population will lose all mtDNA diversity within the next 20 years. This time-series of reduction in genetic variation provides independent evidence of the severity of population decline and habitat contraction resulting from fisheries and perhaps other human activities.
Recent studies indicate that postcopulatory sexual selection may represent an important component of the speciation process by initiating reproductive isolation via the evolutionary divergence of fertilization systems. Using two geographically isolated populations of the polyandrous beetle Callosobruchus maculatus, we investigated divergence in fertilization systems by determining the extent of postcopulatory functional incompatibility. Through reciprocal, cross-population matings we were able to separately estimate the effects of male and female population origin and their interaction on the extent of last-male sperm precedence, female receptivity to further copulation and female oviposition. Our results indicate partial incompatibility between the fertilization systems of the two populations at all three functional levels. Males derived from the same population as females outcompete rival, allopatric males with respect to sperm preemption, sperm protection, and ability to stimulate female oviposition. This pattern is reciprocated in both populations indicating that postcopulatory, prezygotic events represent important mechanisms by which between-population gene flow is reduced. We suggest the partial gametic isolation observed is a by-product of the coevolution of male and female fertilization systems by a process of cryptic female choice. Our results are consistent with a mechanism akin to conventional mate choice models although they do not allow us to reject antagonistic sexual coevolution as the mechanism of cryptic female choice.
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)
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.
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.
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.
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.
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.
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.
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.
Crosses between Drosophila melanogaster and D. simulans normally result in progeny that are either inviable or sterile. Recent discovery of strains that rescue these inviability and sterility phenotypes has made it possible to study the developmental basis of reproductive isolation between these two species in greater detail. By producing both rescued and unrescued hybrids and examining the protein product staining patterns of genes known to be involved in early germline development and gametogenesis, we have found that in crosses between D. simulans and D. melanogaster, hybrid female sterility results from the improper control of primordial germline proliferation, germline stem cell maintenance, and cystoblast formation and differentiation during early oogenesis. Rescued hybrid females are fertile, yet they generally have lower amounts of adult germline from the outset and show a premature degeneration of adult germline cells with age. In addition, older rescued hybrid females also exhibit mutant egg phenotypes associated with defects in dorso-ventral patterning which may result from the improper partitioning of cytoplasmic factors during early oogenesis that could stem from the early defect. Although a variety of germline and oogenic defects are described for the hybrid females, all of them can potentially result from the same underlying primary defect. Hybrid males from these same crosses, on the other hand, have no detectable germline in adult reproductive tissues, even when hybrid sterility rescue strains are used, indicating that male sterility and female sterility stem from distinctly different developmental defects.