Genetic differentiation and habitat connectivity across towhee hybrid zones in Mexico

Hybridization can either reinforce or erode species boundaries; therefore, hybrid zones offer a natural experimental setting in which to assess the dynamics of reproductive isolation. Secondary contact zones, in particular, present a partial separation of the original divergence mechanisms and the subsequent genomic architecture of reproductive isolation (or lack thereof). The spatial context of secondary contact and its consequent effect on dispersal play vital roles on the contact’s outcome. In a hybrid complex between two towhee species in Mexico, Pipilo maculatus and Pipilo ocai, two major hybrid gradients provide natural replicates for comparison. However, genetic analyses demonstrate significant divergence between geographically separate parental populations of each species and divergence of populations within each hybrid zone. The two hybrid transects (Teziutlán and Transvolcanic) are distinct and evidence suggests allelic introgression both across the species boundary and between the two transects. Habitat corridors for dispersal represent functional connectivity hotspots where the two transects meet. Both habitat connectivity and genetic differentiation between geographically disparate parental populations appear to influence the dynamics of gene flow across the hybrid gradients. In southern sympatric populations (Mt. Orizaba and Oaxaca) where morphological evidence for hybridization is scarce, opposing species’ alleles appear to traverse through the hybrid zones rather than arising from cryptic local hybridization. These results illustrate the importance of environmentally mediated gene flow in the context of secondary contact as an important force influencing evolutionary trajectory.     

Genetic mapping based on radiation hybrid data.

Radiation hybrid mapping is a recently developed technique for constructing high-resolution maps of mammalian chromosomes. The chromosome of interest is exposed to X rays, which cause it to break into numerous fragments. The fragments are randomly recovered into hamster cells. The resulting hybrids are then analyzed for the presence or absence of DNA markers of interest. A simple method for using this information to order the markers on the chromosome is developed here. For any particular ordering of the loci, the least number of breaks consistent with the data can be determined. The best order is taken to be that which minimizes this number of obligatory breaks. Some statistical properties of the method are examined, both theoretically and by use of a simulation study. Except for some extreme cases, the method gives good results, which compare well with results obtained by a full maximum likelihood analysis. The method is used to order a set of 14 loci on chromosome 21 with encouraging results.     

Genetic mapping of monoterpene composition in an interspecific eucalypt hybrid

Genetic control of foliar oil composition was investigated amongst half-sib progeny of an interspecific eucalypt hybrid. The oil was found to be largely composed of the monoterpenes, limonene, α−pinene, γ−terpinene, 1,8 cineole and p-cymene. Due to difficulties in the interpretation of the compositional data based on raw proportions, further analysis was conducted using log-ratio variables. A high degree of intercorrelation amongst log-ratios was thought to be a consequence of commonality in the biosynthetic origins of the monoterpenes. Quantitative trait locus (QTL) analysis of log-ratio variables indicated that a significant (68–81%) proportion of the variation in four out of the ten possible log-ratios were controlled by a single genomic region of the maternal Eucalyptus grandis parent. The impact of this genomic region upon oil composition was thought to be a consequence of a gene, or genes, controlling the production of limonene, as limonene was the predominant oil constituent in many hybrid individuals and was common to all log-ratios associated with the identified genomic region. Received: 20 November 1998 / Accepted: 16 June 1999     

Genotypic proportions in hybrid zones

We study a hybrid zone between two populations of a diploid organism. The populations differ at one locus. Homozygotes have equal fitnesses and the heterozygote fitness is reduced by + ( is the birth rate deviation and is the death rate deviation). The populations extend along a one dimensional continuous habitat, and migration occurs by diffusion of individuals. The model is formulated as a set of simple continuous time demographic models without age structure for the three genotypes, and the system is transformed into three new variables, the total population size N, the gene frequency p, and the deviation from Hardy-Weinberg proportions F. The gene frequency in a steady state cline always follows a hyperbolic tangent closely. Analysis of the asymptotic behavior of the cline far from the hybrid zone suggests a qualitative prediction of the shape of N, p and F over the zone. For weak selection the shape is determined by a central steepness of , as observed by Bazykin in 1969, where D is the diffusion coefficient. For strong selection the cline is less steep than the Bazykin cline, and the form is dominated by the migration process. The steepness at the center of the cline is close to where b is the birth rate of homozygotes.Supported in part by grant 11-9639-1 from the Danish Natural Science Research Council and grant 5.18.2023 from the Danish Biotechnological Research and Development Programme     

Admixture mapping of quantitative traits in Populus hybrid zones: power and limitations

Uncovering the genetic architecture of species differences is of central importance forunderstanding the origin and maintenance of biological diversity. Admixture mapping can beused to identify the number and effect sizes of genes that contribute to the divergence ofecologically important traits, even in taxa that are not amenable to laboratory crossesbecause of their long generation time or other limitations. Here, we apply admixturemapping to naturally occurring hybrids between two ecologically divergent Populusspecies. We map quantitative trait loci for eight leaf morphological traits using 77mapped microsatellite markers from all 19 chromosomes of Populus. We applymultivariate linear regression analysis allowing the modeling of additive and non-additivegene action and identify several candidate genomic regions associated with leaf morphologyusing an information-theoretic approach. We perform simulation studies to assess the powerand limitations of admixture mapping of quantitative traits in natural hybrid populationsfor a variety of genetic architectures and modes of gene action. Our results indicate that(1) admixture mapping has considerable power to identify the genetic architecture ofspecies differences if sample sizes and marker densities are sufficiently high, (2)modeling of non-additive gene action can help to elucidate the discrepancy betweengenotype and phenotype sometimes seen in interspecific hybrids, and (3) the geneticarchitecture of leaf morphological traits in the studied Populus species involvescomplementary and overdominant gene action, providing the basis for rapid adaptation ofthese ecologically important forest trees.     

Three-trophic-level interactions in cattail hybrid zones

Plant-herbivore and herbivore-parasitoid wasp interactions were examined in three hybrid zones of the cattails Typha latifolia and T. angustifolia in south-eastern Michigan over a 2-year period. Patterns of resource use by two lepidopteran species of seed-eating herbivores were studied and herbivore densities sustained by the hybrid cattail T. x glauca and its parental species were determined. Densities of the commoner seed-eating lepidopteran, Lymnaecia phragmitella, were found to be highest in seed heads of both parental species and lowest in hybrid seed heads in zones of hybridization, thus supporting the hybrid resistance hypothesis of Fritz et al. (1994). Densities of the second herbivore, Dicymolomia julianalis, on the hybrid were lower than on the parental T. latifolia, but did not differ from the mean of the combined herbivore densities of the two parental species. D. julianalis in cattails appears to fit the additive hypothesis (Fritz et al. 1994). Parasitism of L. phragmitella by four species of parasitoid wasp, Itoplectis conquisitor (Ichneumonidae), Scambus hispae (Ichneumonidae), Macrocentrus delicatus (Braconidae), and Temelucha gracilipes (Ichneumonidae), was studied to determine if rates of parasitization by these natural enemics on the different cattails were similar to the patterns of plant exploitation exhibited by their hosts. Parasitism rates of L. phragmitella were significantly higher in larvae reared on hybrid cattails. This study reveals two different patterns of hybrid utilization by two very similar herbivores. Opposite patterns of plant utilization were found for the parasitoids compared with their host (L. phragmitella) in cattail hybrid zones.     

Bimodal hybrid zones and speciation

Contact zones exemplify a series of stages in speciation. In unimodal hybrid zones intermediates predominate; in bimodal zones hybrids are rare and parental forms predominate; and finally, species might overlap, but never hybridize. Recent studies show bimodality to be associated strongly with assortative mating or fertilization, and only weakly with overall levels of genetic divergence or intrinsic genomic incompatibility. Ecological divergence across most bimodal hybrid zones suggests that ecology contributes more to speciation than genomic incompatibility. This continuum of stable contact zones provides empirical evidence for a route to speciation, which does not require allopatry.     

Mammalian hybrid zones: a review

相似文献   

Demographic history shapes genomic ancestry in hybrid zones

Demographic factors such as migration rate and population size can impede or facilitate speciation. In hybrid zones, reproductive boundaries between species are tested and demography mediates the opportunity for admixture between lineages that are partially isolated. Genomic ancestry is a powerful tool for revealing the history of admixed populations, but models and methods based on local ancestry are rarely applied to structured hybrid zones. To understand the effects of demography on ancestry in hybrids zones, we performed individual‐based simulations under a stepping‐stone model, treating migration rate, deme size, and hybrid zone age as parameters. We find that the number of ancestry junctions (the transition points between genomic regions with different ancestries) and heterogenicity (the genomic proportion heterozygous for ancestry) are often closely connected to demographic history. Reducing deme size reduces junction number and heterogenicity. Elevating migration rate increases heterogenicity, but migration affects junction number in more complex ways. We highlight the junction frequency spectrum as a novel and informative summary of ancestry that responds to demographic history. A substantial proportion of junctions are expected to fix when migration is limited or deme size is small, changing the shape of the spectrum. Our findings suggest that genomic patterns of ancestry could be used to infer demographic history in hybrid zones.     

Admixture in European Populus hybrid zones makes feasible the mapping of loci that contribute to reproductive isolation and trait differences

The use of admixed human populations to scan the genome for chromosomal segments affecting complex phenotypic traits has proved a powerful analytical tool. However, its potential in other organisms has not yet been evaluated. Here, we use DNA microsatellites to assess the feasibility of this approach in hybrid zones between two members of the 'model tree' genus Populus: Populus alba (white poplar) and Populus tremula (European aspen). We analyzed samples of both species and a Central European hybrid zone (N=544 chromosomes) for a genome-wide set of 19 polymorphic DNA microsatellites. Our results indicate that allele frequency differentials between the two species are substantial (mean delta=0.619+/-0.067). Background linkage disequilibrium (LD) in samples of the parental gene pools is moderate and should respond to sampling schemes that minimize drift and account for rare alleles. LD in hybrids decays with increasing number of backcross generations as expected from theory and approaches background levels of the parental gene pools in advanced generation backcrosses. Introgression from P. tremula into P. alba varies strongly across marker loci. For several markers, alleles from P. tremula are slightly over-represented relative to neutral expectations, whereas a single locus exhibits evidence of selection against P. tremula genotypes. We interpret our results in terms of the potential for admixture mapping in these two ecologically divergent Populus species, and we validate a modified approach of studying genotypic clines in 'mosaic' hybrid zones.     

Clustering of contact zones, hybrid zones, and phylogeographic breaks in North America

A recent test for the existence of suture zones in North America, based on hybrid zones studied since 1970, found support for only two of the 13 suture zones identified by Remington in 1968 (Swenson and Howard 2004). One limitation of that recent study was the relatively small number of hybrid zones available for mapping. In this study, we search for evidence of clustering of contact zones between closely related taxa using data not only from hybrid zones but from species range maps of trees, birds, and mammals and from the position of phylogeographic breaks within species. Digital geographic range maps and a geographic information system approach allowed for accurate and rapid mapping of distributional data. Areas of contact between closely related species and phylogeographic breaks within species clustered into areas characterized by common physiographic features or predicted by previously hypothesized glacial refugia. The results underscore the general importance of geographic barriers to dispersal (mountain chains) and climate change (periods of cooling alternating with periods of warming, which lead to the contraction and expansion of species ranges) in species evolution.     

Genetic and radiation hybrid mapping of the hyperekplexia region on chromosome 5q.

Hyperekplexia, or startle disease (STHE), is an autosomal dominant neurologic disorder characterized by muscular rigidity of central nervous system origin, particularly in the neonatal period, and by an exaggerated startle response to sudden, unexpected acoustic or tactile stimuli. STHE responds dramatically to the benzodiazepine drug clonazepam, which acts at gamma-aminobutyric acid type A (GABA-A) receptors. The STHE locus (STHE) was recently assigned to chromosome 5q, on the basis of tight linkage to the colony-stimulating factor 1-receptor (CSF1-R) locus in a single large family. We performed linkage analysis in the original and three additional STHE pedigrees with eight chromosome 5q microsatellite markers and placed several of the most closely linked markers on an existing radiation hybrid (RH) map of the region. The results provide strong evidence for genetic locus homogeneity and assign STHE to a 5.9-cM interval defined by CSF1-R and D5S379, which are separated by an RH map distance of 74 centirays (roughly 2.2-3.7 Mb). Two polymorphic markers (D5S119 and D5S209) lie within this region, but they could not be ordered with respect to STHE. RH mapping eliminated the candidate genes GABRA1 and GABRG2, which encode GABA-A receptor components, by showing that they are telomeric to the target region.     

Genetic mapping in maize with hybrid progeny across testers and generations: plant height and flowering

DNA markers were used to identify quantitative trait loci (QTLs) for plant height, ear height, and three flowering traits in hybrid progeny of two generations (F_2:3, F_6:8) of lines from a Mo17×H99 maize population. For both generations, testcross (TC) progeny were developed by crossing the lines to three inbred testers (B91, A632, B73). The hybrid progeny from the two generations were evaluated at the same locations but in different years as per an early generation testing program. QTLs were identified within each TC population and for mean testcross (MTC) performance. Overall, more QTLs were detected in the F_6:8 than the F_2:3 generation. Totalled over all five traits, 41 (B91) to 69% (B73) of the QTLs for tester effects and 67% of the QTLs for MTC detected in the F_2:3 generation were verified in the F_6:8 generation. Although differences in relative rank of the QTL effects across generations were observed, especially for the flowering traits, parental contributions were nearly always consistent. Several (8–11) QTLs were identified with effects for all three tester populations and for all traits except the anthesis-silk interval, which had only two such regions. Over all five traits, previous evaluations in this population identified 26 QTLs with consistent effects for two (F_2:3, F_6:8) inbred-progeny evaluations, and 20 (77%) were also associated with MTC in at least one of the generations evaluated herein. In all instances of common inbred and TC QTLs, parental contributions were the same. Received: 26 November 1999 / Accepted: 18 April 2000     

Levels of herbivory and parasitism in host hybrid zones

Hybrid zones often are intermediate in their habitat, and have individuals that are intermediate genetically to those of either parental species. Recently, a number of studies has addressed the distribution of herbivores and parasites in hybrid zones of their hosts. Mechanisms that underlie these distributions include behavioral, environmental and genotypic differences between hybrids and their parental species.     

Introgression at differentially aged hybrid zones in red-tailed chipmunks

Hybrid zones allow us to investigate the maintenance and the break down of reproductive isolation; they are a window into the speciation process. Tamias ruficaudus (red-tailed chipmunk) has a roughly ring-like distribution in the Inland Northwest and includes two morphologically well-differentiated subspecies, T. r. ruficaudus (in the eastern portion of its range) and T. r. simulans (in the western portion). These taxa meet at two contact zones: the Lochsa River in Idaho and 200 km to the north, near Whitefish, Montana. The Lochsa Zone is encompassed within the Clearwater River Drainage, which has been proposed as a glacial refugium for many taxa throughout the Pleistocene, whereas the Whitefish Zone was under the Cordilleran ice sheet during the most recent glacial maxima approximately 10,000 years ago. Mitochondrial DNA introgression has been documented at both contact zones, yet the subspecies remain significantly distinct with respect to bacular morphology and no intermediate morphologies have ever been reported. Here, we elucidate differentiation and introgression of the nuclear genome using ten microsatellite loci and compare findings to previously described mitochondrial DNA haplotype distribution and introgression. We found significant substructure in the nuclear data; each subspecies is divided into at least two genetically distinct demes. At the Lochsa contact zone, individuals restricted to the mtDNA zone of introgression form a distinct deme at microsatellite loci whereas in the younger, Whitefish contact zone, there is no hybrid-zone specific group. The genetic distances of the demes within these two subspecies indicate recent northward expansion.