ASYMMETRIC VIABILITY OF RECIPROCAL-CROSS HYBRIDS BETWEEN CRESTED AND MARBLED NEWTS (TRITURUS CRISTATUS AND T. MARMORATUS)

Hybridization between divergent lineages often results in reduced hybrid viability. Here we report findings from a series of independent molecular analyses over several seasons on four life stages of F₁ hybrids between the newts Triturus cristatus and T. marmoratus . These two species form a bimodal hybrid zone of broad overlap in France, with F₁ hybrids making up about 4% of the adult population. We demonstrate strong asymmetry in the direction of the cross, with one class ( cristatus -mothered) making up about 90% of F₁ hybrids. By analyzing embryos and hatchlings, we show that this asymmetry is not due to prezygotic effects, as both classes of hybrid embryos are present at similar frequencies, implicating differential selection on the two hybrid classes after hatching. Adult F₁ hybrids show a weak Haldane effect overall, with a 72% excess of females. The rarer marmoratus -mothered class, however, consists entirely of males. The absence of females from this class of adult F₁ hybrids is best explained by an incompatibility between the cristatus X chromosome and marmoratus cytoplasm. It is thus important to distinguish the two classes of reciprocal-cross hybrids before making general statements about whether Haldane's rule is observed.     

Fertility of roach × bream hybrids, Rutilus rutilus (L.) ×Abramis brama (L.), and their identification

Adult roach, bream and their presumed F₁ hybrid from an Anglian Water reservoir were identified on the basis of morphological and meristic characteristics. The hybrid was clearly intermediate. Four hybrid breeding crosses were induced to spawn by hypophysis. A bream × roach cross (female named first) failed to produce fertile eggs, whereas F₁ hybrid × roach, roach × F₁ hybrid and F₁ hybrid × F₁ hybrid all produced fry. Fertility (defined as survival of eggs to hatching) was high for the F₁ hybrid × roach back-cross (56%) but low for the others (<2%), in comparison to the pure species controls (roach 69%, bream 76%). Progeny from these crosses were reared until anal fin rays could be counted. These counts indicated intermediacy between the parents and back-crossed individuals, and similarity between F₁ hybrids and their F₂ progeny.     

The evolution of F1 postzygotic incompatibilities in birds

Abstract.— We analyzed the rate at which postzygotic incompatibilities accumulate in birds. Our purposes were to assess the role of intrinsic F₁ hybrid infertility and inviability in the speciation process, and to compare rates of loss of fertility and viability between the sexes. Among our sample more than half the crosses between species in the same genus produce fertile hybrids. Complete loss of F₁ hybrid fertility takes on the order of millions of years. Loss of F₁ hybrid viability occurs over longer timescales than fertility: some viable hybrids have been produced between taxa that appear to have been separated for more than 55 my. There is strong support for Haldane's rule, with very few examples where the male has lower fitness than the female. However, in contrast to Drosophila , fertility of the homogametic sex in the F₁ appears to be lost before viability of the heterogametic sex in the F₁. We conclude that the time span of loss of intrinsic hybrid fertility and viability is often, but not always, longer than the time to speciation. Premating isolation is an important mechanism maintaining reproductive isolation in birds. In addition, other factors causing postzygotic reproductive isolation such as ecological causes of hybrid unfitness, reduced mating success of hybrids, and genetic incompatibilities in the F₂s and backcrosses may often be involved in the speciation process.     

Mating patterns of Darwin's Finch hybrids determined by song and morphology

Three species of Darwin's Finches hybridize on the Galapagos island of Daphne Major. We examined mating patterns to determine if hybrids exhibit mate preferences. Geospiza fortis x G. scandens F₁ hybrids backcrossed to both of the parental species, whereas all backcrosses bred with the parental species to which they were most related, or with hybrids. Paternal song was shown to be the crucial factor determining the mating pattern of G. fortis x G. scandens F₁ hybrids and their offspring. Song is culturally inherited, transmitted faithfully from father to son (with few exceptions) as a result of an imprinting-like process. Size also contributes to the choice of mates. G. fortis x G. scandens F₁ hybrid females paired with large G. scandens -like G. fortis males. G. fortis x G. fuliginosa F₁ hybrids paired negatively assortatively with respect to the size of their G. fortis mates. Non-random mating of hybrids based on song, a non-generic trait, has interesting evolutionary consequences. Song characteristics and nuclear and mitochondrial genes flow from G. fuliginosa into the G. fortis population, whereas the direction of transfer of genetic and song information between G. fortis and G. scandens depends on which song was sung by the father of the hybrids.     

Reproductive trait divergence and hybrid fertility patterns between chromosomal races of the house mouse in Tunisia: analysis of wild and laboratory-bred males and females

The divergence in reproductive features and hybrid fertility patterns between two chromosomal races (2 n  = 40, 40St, and 2 n  = 22, 22Rb) of the house mouse in Tunisia were re-assessed on a larger sample of wild and laboratory-bred individuals than studied hitherto. Results showed that litter sizes were significantly smaller in 40St than in 22Rb mice, contrary to previous analyses. This suggests that variation in litter size between the two chromosomal races is more likely related to selective and/or environmental factors acting locally than to interracial reproductive trait divergence. However, the significantly reduced litter size of F₁ hybrids compared with parental individuals was confirmed, and further highlighted a sex difference in hybrid infertility, as F₁ females produced fewer litters and of smaller size than males. Histological analyses of F₁ and backcrosses showed a breakdown of spermatogenesis in males and a significantly reduced primordial follicle pool in females. The degree of gametogenic dysfunction was not related to the level of chromosomal heterozygosity per se , but a significant effect of two Rb fusions on follicle number was observed in hybrid females. These results suggest that genetic incompatibilities contribute to primary gametogenic dysfunction in hybrids between the chromosomal races in Tunisia.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 407–416.     

Chloroplast ultrastructure in two Crassulacean species and an F1 hybrid with differing biomass δ13C values

Abstract. The chloroplasts of two species of the Crassulaceae and their F₁ hybrid were compared by electron microscopy. The two species had contrasting leaf tissue δ¹³C values of −25°/_∞ ( Sedum greggii ) and −13°/_∞ ( Cremnophila linguifolia ), and the F₁ hybrid had a value of − 18°/_∞. S. greggii had a mean of 8.9 thylakoids per granum in contrast to C. linguifolia which had a mean of only 3.8 thylakoids per granum. The F₁ hybrid had a mean of 6.4 thylakoids per granum. Crystaloids were observed in S. greggii and the hybrid but not in C. linguifolia     

Grey plumage colouration in the duck is genetically determined by the alleles on two different, interacting loci

Based on the observation of a grey phenotype in the F₁ generation from a cross between two white plumage duck varieties, the white Kaiya and the white Liancheng , we hypothesized a possible interaction between two autosomal loci that determine grey plumage. Using the parental and F₁ individuals, seven testing combinations including five different F₁ intercrosses (F₂) and two different backcrosses (BC₁ and BC₂) were designed to test our hypothesis. It was demonstrated by chi-squared analysis that six test matings produced offspring in the expected ratios between the grey and white, with P- values ranging from 0.50 to 0.99. Another mating, where all white offspring were expected, produced 33 white individuals. These results verified that the interaction between two loci produced the grey phenotype. The C locus, which carries the recessive allele ( c ), was previously thought to be the only gene responsible for white plumage in the duck. This is the first report that an allele ( t ), carried by the white Liancheng at a different autosomal locus, also determines white plumage in ducks. Furthermore, the dominant alleles at both loci can interact with each other to produce the grey phenotype, and a new dark phenotype, observed in some F₂ individuals, can be attributed to the dosage effect of the T allele.     

Artificial hybridization within Saxifraga pentadactylis (Saxifragaceae)

Saxifraga pentadactylis subsp. almanzorii , an endemic to the subalpine nucleus of Sierra de Gredos (central Spain), differs from its closest relative, subsp. willkommiana , by its less showy petals. An artificial crossing program was carried out in order to assess the degree of reproductive isolation between the subspecies. To facilitate interpretation of the results, the program was extended to 10 other interspecific hybrid combinations within sect. Saxifraga . All the data gathered are congruent with the occurrence of two evolutionary scenarios. Interspecific crossings, rendering moderate to high seed-set (in obtaining the F₁), and vigorous but relatively sterile F₁ offspring, reveal reproductive barriers at the level of the F₁ fertility, probably originated as a byproduct of divergent evolution. In contrast, intraspecific crossings within S. pentadactylis resulted in seed-set values lower than expected (in obtaining the F₁), in a majority of weak non-viable F₁ offspring but also in a few fertile F₁ hybrid specimens which were able to originate F₂ offspring. This second pattern reveals reproductive barriers at the level of the F₁ vitality, probably arisen in a quite abrupt fashion. The lower P/O for subsp. almanzorii as compared to subsp. willkommiana , together with the rest of the evidence suggest that the reproductive barriers between them might be the product of active selection against hybridization achieved by incrementing the levels of autogamy in the former.     

Environmental influences on epistatic interactions: viabilities of cytochrome c genotypes in interpopulation crosses

Abstract .The genetic incompatibilities that underlie F₂ hybrid breakdown and reproductive isolation between al-lopatric populations may be susceptible to environmental interactions. Here we show that epistatic interactions between cytochrome c ( CYC ) alleles and mitochondrial DNA (mtDNA) variation are dramatically influenced by environmental temperature in interpopulation hybrids of the copepod Tigriopus californicus . CYC is a nuclear-encoded gene that functionally interacts with electron transport system (ETS) complexes composed in part of mtDNA-encoded proteins. Previous studies have provided evidence for functional coadaptation between CYC and ETS complex IV (cytochrome c oxidase) and for cytoplasmic effects on the fitness of CYC genotype in copepod hybrids. In this study, selection on CYC genotype is shown to continue into advanced generation hybrids (F₂-F₈) increasing the likelihood that CYC itself is involved in the interaction (and not a linked factor). Relative viabilities varied markedly between copepods raised in two different temperature/light regimes. These results suggest that both intrinsic coadaptation and extrinsic selection will influence the outcome of natural hybridizations between populations. Furthermore, the results indicate that the fitness of particular hybrid genotypes depends on additional non-mtDNA encoded genes that interact with CYC.     

Can Extensive Reticulation and Concerted Evolution Result in a Cladistically Structured Molecular Data Set?

Hierarchy is the main criterion for informativeness in a data set, even if no explicit reference to evolution as a causal process is provided. Sequence data (nuclear ribosomal DNA ITS) from Armeria (Plumbaginaceae) contains a certain amount of hierarchical structure as suggested by data decisiveness and distribution of tree lengths. However, ancillary evidence suggests that extensive gene flow and biased concerted evolution in these multicopy regions have significantly shaped the ITS data set. This argument is discussed using parsimony analysis of four data sets, constructed by combining wild sequences with those from different generations of artificial hybrids (wild + F₁, F₂, and backcrosses; wild + backcrosses; wild + F₁; wild + F₂). Compared to the F₁ hybrids, F₂ show a certain degree of homogenization in polymorphic sites. This effect reduces topological disruption caused by F₁ and is considered to be illustrative of how extensive gene flow and biased concerted evolution may have modeled the wild ITS data. The possibility that hierarchy has arisen as a result of—or despite a significant contribution from—those two such potentially perturbing forces raises the question of what kind of signal are we recovering from this molecular data set.     

A test of ecological selection against young-of-the-year hybrids of sympatric sticklebacks

Ecological selection against hybrids, the reduction in hybrid fitness attributed solely to environmental factors, was tested by introducing young-of-the-year benthic, limnetic and F₁ hybrid sticklebacks Gasterosteus aculeatus to divided experimental ponds and lake enclosures. The frequency of hybrids in samples taken at the end was significantly lower than their frequency at introduction. Hybrid survival was significantly lower in pond-sides in which they were initially the most common cross type than in pond-sides in which they were initially rare, suggesting that hybrid survival may be frequency-dependent. Growth rate of F₁ hybrids was marginally lower than benthic growth rates, being significantly lower than in ponds and not different in lake enclosures. The diet of hybrids overlapped with both parent species in ponds and with benthic diets in lake enclosures. The results suggest that ecological selection is acting against young-of-the-year hybrid sticklebacks.     

Spontaneous hybridizations between oilseed rape and wild radish

The occurence of spontaneous hybridization between Brassica napus (oilseed rape) and Raphanus raphanistrum (wild radish) was investigated under different density conditions in cages and open-field experiments. Hybrids with wild radish as the seed parent were identified by screening for herbicide resistance belonging to rape. Small seed size and intermediate morphology were used to screen for hybrids with rape as the seed parent. Leaf isozyme patterns and flow cytometry provided confirmation of hybrids. Wild radish in an oilseed rape field produced as many as three interspecific hybrids per 100 plants. This is the first report of such a spontaneous event. The frequency of hybrids is expected to range from 0.006 to 0.2% of the total seed produced, at P = 0.05. Male-sterile oilseed rape plants surrounded by wild radish can produce up to 37 hybrids per plant. Seed production of the F₁ hybrids and their F₂ descendants was up to 0.4% and 2%, respectively, of that of wild radish. Gene escape from transgenic oilseed rape to wild related species is discussed.     

Mating Types and Mating Type Inheritance in Euplotes minuta Yocom (Ciliata, Hypotrichida)

SYNOPSIS. Thirty-one stocks of a marine ciliate, Euplotes minuta Yocom, collected from different localities, can be grouped in seven mating types. True pairs are formed only in mixtures of stocks belonging to different mating types. No selfing pairs or intraclonal conjugation have ever been observed. Synclonal inheritance of mating types is the rule, although about 10% of pairs show clonal inheritance. The latter can be explained by assuming syncaryon formation through cytogamy or through caryogamy of pronuclei derived from different products of meiosis. Mating type determination is due to 7 alleles at the single locus mt . There is complete dominance among the 7 alleles which can be orderly seriated, as shown in Table 3, according to their dominance relationship. The 5 stocks, and only these 5, of mating type VII have the autogamy trait. The mortality rate in F₁ and F₂ is very low–a maximum of 10%; however, the F₂'S obtained by autogamy from F₁ progenies in which mating type VII is involved have a very high mortality rate. The two facts (high mortality rate in F₂ and strict association of autogamy trait with mating type VII in natural populations) have been considered as evidences of a probable isolation mechanism existing between mating type VII and the other 6 mating types. Thus, the 7 mating types have been assigned to the same syngen only tentatively.     

INHERITANCE OF RESISTANCE TO CYHALOTHRIN IN THE HOUSEFLY (DIPTERA: MUSCIDAE)

Abstract The genetic inheritance of resistance to cyhalothrin in housefly, Musca domstica (L) was investigated.
Reciprocal crosses between susceptible (S) and resistant (R) strains were used to determine the characteristics of resistance. Analysis of probit line from the F₁ generation and F₂ generation obtained by inbreeding the F₁ hybrids indicated that cyhalothrin resistance was controlled by more than one factors and degree of resistance dominance to cyhalothrin was -0.10, indicating cyhalothrin resistance is conferred by incompletely recessive gene(s). The realized heritability of resistance to cyhalothrin cyhalothrin calculated from data collected routinely from laboratory selection was 0.12.     

Electrophoretic analysis of protein systems of Ctenopharyngodon idella (Val.), Hypophthalmichthys nobilis (Rich.) and their F1 triploid hybrid

Electrophoretic analysis was performed on eight protein systems (lactate dehydrogenase, liver esterases, superoxide dismutase, haemoglobin, isocitrate dehydrogenase, glucose-6-phosphate dehydrogenase, 6-phosphogIuconate dehydrogenase, transferrin) in Ctenopharyngodon idella, Hypophthalmichthys nobilis , and their F₁ triploid hybrid. The data demonstrated codominant Mendelian inheritance at individual loci for each protein system. Transferrin and superoxide dismutase were the only proteins found to be polymorphic in either parental species. Both parental alleles were present and functional at each enzyme locus examined in the F₁ triploid hybrid.     

Rare and asymmetrical hybridization of the endemic Barbus carpathicus with its widespread congener Barbus barbus

In endemic species that co-occur with widespread congeners, hybridization can lead to an influx of novel and beneficial genetic variation, but high rates of introgression may cause genetic swamping of the endemic species and have detrimental effects on its survival potential. This study examines hybridization between sympatric populations of the Carpathian barbel Barbus carpathicus , a recently discovered cryptic species with a restricted range, and the widespread common barbel Barbus barbus . Based on six diagnostic allozyme loci, a microsatellite locus and mtDNA, hybrids were found to be present at multiple localities within the Vistula River drainage (Baltic Sea) as well as in the Tisza River system of the Danube River drainage (Black Sea). However, the numbers of hybrids were very low; four individuals of 230 fish sampled from the Vistula drainage. Bayesian assessment of their nuclear genotypes suggested that two hybrids in the Vistula drainage and nine in the Tisza system were F₁ generation, and one in the Vistula drainage and one in the Tisza system were backcrosses (BC) to B. barbus , while no F₂ or BC to B. carpathicus were detected. No hybrid carried B. carpathicus mtDNA and cytonuclear linkage disequilibria showed significant positive associations between hybrid genotypes and B. barbus mtDNA, suggesting unidirectionality in the interspecific mating with a disproportionate contribution of B. barbus mothers. Despite geographically broad occurrence of hybrids, these data provide evidence of strong constraints on hybridization in the native breeding habitats and the lack of introgression towards B. carpathicus .     

Breeding experiments to distinguish two sibling species of Halicarcinus (Crustacea, Brachyura)

Hybridization experiments and various salinities were used in investigating the status of two Halicarcinus lacustris-like , forms, one of which has no free larval stages (direct-development) and the other of which has normal larval stages (indirect-development).
Hybrids were produced, at least from indirect-development female x direct-development male crosses, but introgression of genetic material was limited to F₂ hybrids from the female parental form. There were no intergrades between the two modes of development in F₁ and F₂ hybrids or as a response to salinity.
These forms are established as sibling species. The name lacustris applies to the direct-development form and the other is a new species, H. paralacustris , which is described. Hybridization in the field is unlikely due to geographical and reproductive isolating mechanisms. Contrary to a previous view, it seems that H. lacustris had a recent origin in Australia and transport by water birds is suggested as the mode of dispersal through south-east Australasia.     

Population differentiation in an annual legume: genetic architecture

Abstract. The presence or absence of epistasis, or gene interaction, is explicitly assumed in many evolutionary models. Although many empirical studies have documented a role of epistasis in population divergence under laboratory conditions, there have been very few attempts at quantifying epistasis in the native environment where natural selection is expected to act. In addition, we have little understanding of the frequency with which epistasis contributes to the evolution of natural populations. In this study we used a quantitative genetic design to quantify the contribution of epistasis to population divergence for fitness components of a native annual legume, Chamaecrista fasciculata . The design incorporated the contrast of performance of F₂ and F₃ segregating progeny of 18 interpopulation crosses with the F₁ and their parents. Crosses were conducted between populations from 100 m to 2000 km apart. All generations were grown for two seasons in the natural environment of one of the parents. The F₁ often outperformed the parents. This F₁ heterosis reveals population structure and suggests that drift is a major contributor to population differentiation. The F₂ generation demonstrated that combining genes from different populations can sometimes have unexpected positive effects. However, the F₃ performance indicated that combining genes from different populations decreased vigor beyond that due to the expected loss of heterozygosity. Combined with previous data, our results suggest that both selection and drift contribute to population differentiation that is based on epistatic genetic divergence. Because only the F₃ consistently expressed hybrid breakdown, we conclude that the epistasis documented in our study reflects interactions among linked loci.     

Postzygotic isolation between the two European subspecies of the house mouse: estimates from fertility patterns in wild and laboratory-bred hybrids

We assessed the fertility (reproductive success, litter size, testis weight, spermatocyte-to-spermatid ratio) of F₁s and backcrosses between different wild-derived outbred and inbred strains of two mouse subspecies, Mus musculus domesticus and M. m. musculus . A significant proportion of the F₁ females between the outbred crosses did not reproduce, suggesting that female infertility was present. As the spermatocyte-to-spermatid ratio was correlated with testis weight, the latter was used to attribute a sterile vs. fertile phenotype to all males. Segregation proportions in the backcrosses of F₁ females yielded 11 (inbred) to 17% (outbred) sterile males, suggesting the contribution of two to three major genetic factors to hybrid male sterility. Only one direction of cross between the inbred strains produced sterile F₁ males, indicating that one factor was borne by the musculus X-chromosome. No such differences were observed between reciprocal crosses in the outbred strains. The involvement of the X chromosome in male sterility thus could not be assessed, but its contribution appears likely given the limited introgression of X-linked markers through the hybrid zone between the subspecies. However, we observed no sterile phenotypes in wild males from the hybrid zone, although testis weight tended to decrease in the centre of the transect.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 379–393.