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.     

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.     

Intergeneric Somatic Hybridization Between Brassica napus L. and Sinapis alba L.

Electrically induced protoplast fusion was used to produce somatic hybrids between Brassica napus L. and Sinapis alba L. Seven hybrids were obtained and verified by the simple sequence repeat and cleaved amplified polymorphic sequence analysis of the genefael, indicating that the characteristic bands from S. alba were present in the hybrids. The hybridity was also confirmed by chromosome number counting because the hybrids possessed 62 chromosomes, corresponding to the sum of fusion-parent chromosomes. Chromosome pairing at meiosis was predominantly normal, which led to high pollen fertility,ranging from 66% to 77%. All hybrids were grown to full maturity and could be fertilized and set seed after self-pollination or back-crosses with B. napus. The morphology of the hybrids resembled characteristics from both parental species. An analysis of the fatty acid composition in the seeds of F1 plants was conducted and the seeds were found to contain different amounts of erucic acid, ranging from 11.0% to 52.1%.     

Calcium taste preferences: genetic analysis and genome screen of C57BL/6J x PWK/PhJ hybrid mice

To characterize the genetic basis of voluntary calcium consumption, we tested C57BL/6J mice (B6; with low avidity for calcium), PWK/PhJ mice (PWK; with high avidity for calcium) and their F₁ and F₂ hybrids. All mice received a series of 96-h two-bottle preference tests with a choice between water and the following: 50 m m CaCl₂, 50 m m calcium lactate, 50 m m MgCl₂, 100 m m KCl, 100 m m NH₄Cl, 100 m m NaCl, 5 m m citric acid, 30 μ m quinine hydrochloride and 2 m m saccharin. Most frequency distributions of the parental and F₁ but not F₂ groups were normally distributed, and there were few sex differences. Reciprocal cross analysis showed that B6 × PWK F₁ mice had a non-specific elevation of fluid intake relative to PWK × B6 F₁ mice. In the F₂ mice, trait correlations were clustered among the divalent salts and the monovalent chlorides. A genome screen involving 116 markers showed 30 quantitative trait loci (QTLs), of which six involved consumption of calcium chloride or lactate. The results show pleiotropic controls of calcium and magnesium consumption that are distinct from those controlling consumption of monovalent chlorides or exemplars of the primary taste qualities.     

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.     

Some expression of heterosis in F1-hybrid barley

Small and large scale trials of F₁ barley hybrids were done in a glasshouse in the United Kingdom and in the field in the United Kingdom and Europe in 1970, 1971 and 1972. The results showed that there were few instances of significant positive heterosis in yield with either the small or field scale experiments. The hybrids studied were all derived from American varieties as male-sterile parents and therefore the hybrids were not adapted to European countries, thus the case for or against F₁-hybrids cannot be assessed solely on this material.     

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.     

Genetic and in vitro molecular hybridization of malate dehydrogenase isozymes in interspecific bass (Micropterus) hybrids

Supernatant malate dehydrogenase (MDH) isozymes (as visualized by starch gel electrophoresis) are encoded by two distinct gene loci in both the largemouth and smallmouth bass. When an interspecific F₁ hybrid is formed between these two fish, a unique MDH isozyme is generated. The results of freeze-thaw molecular hybridization (which is the first application of this technique to MDH) indicate that this unique isozyme in the F₁ hybrid is a heterodimer composed of one subunit of each parental type. The F₁ hybrids produced F₂ hybrids which in turn formed the F₃ hybrid population. The inheritance of alleles at the MDH-B locus is consistent with a single Mendelian autosomal locus. Furthermore, there is no evidence of linkage between the lactate dehydrogenase-E locus and the MDH-B locus.     

Crossability variation in four South American Hordeum species

Andersson, 2. 1995. Crossability variation in four South American Hordeum species— Nord. J. Bot. IS: 355–364. Copenhagen. ISSN 0107–055X.
Intra- and interpopulation crosses within the four South American Hordeum species, H. cordobense, H. chilense, H. muticum and H. stenostachys were performed. Seeds were obtained in all intrapopulation crosses within all four species. Hordeum cordobense showed little variation in crossability of interpopulation crosses and the F₁-families had rather high fertility. A wide range in crossability between populations was observed in the three species H. chilense, H. muticum and H. stenostachys . Some combinations failed to set seed while others had high seed set. The fertility of the F₁-_- families varied from almost sterile to fully fertile. All parental plants, intra and interpopulation hybrids in the four species had very high meiotic pairing frequency with 12–14 chiasmata per cell. Hordeum cordobense is a homogeneous species while the other three species are heterogeneous and genetically diverse.     

Relationship between species in the genus Rosa, section Pimpinellifoliae

The morphology of twelve species of Rosa is described and similarities between these species are assessed. Possible origins of the tetraploid species from diploid species are indicated on grounds of comparative morphology.
The wild origins of living and herbarium specimens are given in order to supplement published data on geographical distribution.
Meiosis in pollen mother cells, viability of pollen grains at anthesis and ability to set seed was studied in several F₁ hybrids: no indication of complete or even partial sterility was found. Reproductive isolation is therefore unlikely to be maintained by reduced fertility of interspecific hybrids.
Three species are reduced to synonymy with three other species, being retained as subspecific taxa. Two species are transferred from section Pimpinellifoliae to section Cinnamomeae.     

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.     

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.     

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.     

Genetic heterogeneity, reproductive isolation and host preferences in mealy aphids of the Hyalopterus pruni complex (Homoptera, Aphidoidea)

Three taxa were detected by allozyme markers within the mealy aphids of the Hyalopterus pruni complex, having different cultivated Prunus species as main primary hosts. The genetically closer H. pruni and H. amygdali A ( D _Nei= 0.10) never share primary hosts, whereas H. amygdali A and B ( D _Nei= 0.32) may occasionally share them, producing few F₁ hybrids. The three species proved reproductively isolated in the field, with no gene exchange. Their speciation seem to have occurred long before the agricultural revolution, crop colonization representing a host range expansion rather than a host shift, as in sympatric speciation.     

Revision of the genus Diplotaxis (Brassicaceae) in the Cape Verde Islands, W Africa

The genus Diplotaxis (Brassicaceae) in the Cape Verde Islands, W Africa, is revised. Nine taxa are accepted, of which five are described as new: D. antoniensis sp. nov., D. glauca, D. gorgadensis sp. nov., D. gorgadensis ssp. brochmannii ssp. nov., D. gracilis, D. hirta, D. sundingii sp. nov., D. varia sp. nov., and D. vogelii . All species are suffruticose, yellow-flowered perennials and belong to sect. Catocarpum . The species are endemic to the Cape Verde Islands, but show morphological and karyolog-ical affinity to the N African and Mediterranean D. harra s. lat. Analyses of morphological variation in 90 populations (400 plants) revealed a complex pattern, in particular in vegetative characters, most likely evolved by parallel ecogeographical differentiation in different islands. The chromosome number is 2n = 26 (n = 13) in the five taxa investigated. Self-pollination experiments indicate that the species are self-incompatible and outcrossing. Experimental F₁ hybrids with full seed set after open pollination were obtained in 30 interspecific combinations, and the taxa are considered fully interfertile. The taxa are thus isolated mainly by geographical and partly by ecological barriers and have evolved by vicariant evolution (most islands) and adaptive radiation (one island).     

Comparison of migratory tendency in European Quail Coturnix c. coturnix, domestic Japanese Quail Coturnix c. japonica and their hybrids

Since the 1970s, the decline of the occidental populations of European Quail Coturnix c. coturnix , a partial migrant Galliform species, has driven the release of domestic Japanese Quail Coturnix c. japonica into the wild as game birds. Recent experiments suggest that reproductive isolating mechanisms have not developed to prevent hybridization between European and Japanese Quail. To assess the potential impact of hybridization between the subspecies in the wild, this study aims to compare the migratory characteristics of captive European Quail, domestic Japanese Quail, F₁ hybrids and a backcross. In a laboratory experiment, birds hatched in summer and reared under a natural photoperiod were transferred during winter to an artificial photoperiod (light–dark) of 14 : 10 h. Behavioural (migratory restlessness, crowing activity in males) and physiological (sexual development, fat accumulation, body mass) measures were recorded. Domestic Japanese Quail showed no migratory tendency in response to the increased day length. F₁ hybrids showed rapid sexual development and few of them exhibited the typical profile of a migrant bird. Backcrosses showed great variation in their response, comparable with those of the European Quail. Therefore, hybridization could lead to an increased proportion of quail showing sedentary rather than migratory behaviour. Furthermore, hybrids that do show migratory behaviour have the potential to facilitate Japanese gene flow into areas where the release of domestic Quail as game birds is not yet practised.     

Analysis of the oviductal glycoprotein 1 polymorphisms and their effects on components of litter size in rabbits

The objective of this work was to study the effect of the oviductal glycoprotein 1 ( OVGP1 ) genotype and mRNA expression on litter size and other fertility measures, as OVGP1 has positive effects on fertilization and early embryo development. We have analysed an F₂ cross of two lines of rabbits divergently selected for uterine capacity. The OVGP1 mRNA expression was analysed in both lines, but no differences were observed between them. The promoter region and mRNA were sequenced in the F₀ generation, and 17 polymorphic sites were found to co-segregate in three haplotypes (A, B and C). An association study was performed between several reproductive traits and a triallelic microsatellite identified in the promoter region as well as a non-synonymous SNP located in exon 11 [g.12944C>G (p.Arg468Gly)]. The alleles g.12944G and g.325(GT)₁₄T(G)₅ of the B haplotype have a positive effect on the total number of kits born, number born alive, number of implanted embryos and foetal and prenatal embryo survival.     

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.     

Costs of transgenic herbicide resistance introgressed from Brassica napus into weedy B. rapa

Wild relatives of genetically engineered crops can acquire transgenic traits such as herbicide resistance via spontaneous crop–wild hybridization. In agricultural weeds, resistance to herbicides is often a beneficial trait, but little is known about possible costs that could affect the persistence of this trait when herbicides are not used. We tested for costs associated with transgenic resistance to glufosinate when introgressed into weedy Brassica rapa . Crosses were made between transgenic B. napus and wild B. rapa from Denmark. F₁ progeny were backcrossed to B. rapa and BC₁ plants were selected for chromosome numbers similar to B. rapa . Further backcrossing resulted in a BC₂ generation that was hemizygous for herbicide resistance. We quantified the reproductive success of 457 BC₃ progeny representing six full-sib families raised in growth rooms (plants were pollinated by captive bumblebees). Pollen fertility and seed production of BC₃ plants were as great as those of B. rapa raised in the same growth rooms. Segregation for herbicide resistance in BC₃ plants was 1:1 overall, but the frequency of resistant progeny was lower than expected in one family and higher than expected in another. There were no significant differences between transgenic and nontransgenic plants in survival or the number of seeds per plant, indicating that costs associated with the transgene are probably negligible. Results from this growth-chamber study suggest that transgenic resistance to glufosinate is capable of introgressing into populations of B. rapa and persisting, even in the absence of selection due to herbicide application.     

Genetic evidence for asymmetric hybridization between menhadens (Brevoortia spp.) from peninsular Florida

Hybridization between the North American menhadens Brevoortia smithi , Brevoortia tyrannus and Brevoortia patronus has been reported by numerous authors, based primarily on the morphological intermediacy of hybrids relative to each parental type. In this study, genetic characterization of individuals in areas previously designated as hybrid zones validates the presence of F₁ hybrids. Evidence from microsatellite genotypes indicates that B. patronus and B. smithi are parental species of hybrids on both sides of the Florida peninsula, and mitochondrial DNA (mtDNA) haplotypes suggest a maternal contribution by B. smithi in each case. Hybrids between B. tyrannus and B. smithi were not identified in this study. The presence of B. patronus on the Atlantic coast of Florida, coupled with admixture between this species and B. tyrannus , has likely influenced the mixed lineage assortment of mtDNA haplotypes demonstrated in this study and in previous studies.