INBREEDING DEPRESSION IN TWO MIMULUS TAXA MEASURED BY MULTIGENERATIONAL CHANGES IN THE INBREEDING COEFFICIENT

In mixed-mating plant populations, one can estimate the relative fitness of selfed progeny w by measuring the inbreeding coefficient F and selfing rate s of adults of one generation, together with F of adults in the following generation (after selection). In the first application of this multigenerational method, we estimated F and s for adults over three consecutive generations in adjacent populations of two annual Mimulus taxa: the outbreeding M. guttatus and the inbreeding M. platycalyx. This gave estimates of w for the last two generations. Although average multilocus selfing rates were high in both taxa (0.63 in M. guttatus; 0.84 in M. platycalyx), the relative fitness of selfed progeny averaged only 0.19 in M. guttatus and 0.32 in M. platycalyx. An alternative estimator for w that incorporates biparental inbreeding gave even lower estimates of w. These values are significantly below the 0.5 threshold thought to favor selfing, and show that partially selfing populations can harbor substantial genetic load. In accordance with the purging hypothesis, the more highly selfing M. platycalyx showed marginally lower inbreeding depression than M. guttatus in both years (P = 0.08). Inbreeding depression and selfing rates also varied among years in concert among taxa. Several sources of bias are discussed, but computer simulations indicate it is unlikely that w is biased downwards by linkage of marker loci to load loci.     

ANNUAL DIFFERENCES IN FEMALE REPRODUCTIVE SUCCESS AFFECT SPATIAL AND COHORT-SPECIFIC GENOTYPIC HETEROGENEITY IN PAINTED TURTLES

Long-term ecological data were used to evaluate the relative importance of movements, breeding structure, and reproductive ecological factors to the degree of spatial and age-specific variation in genetic characteristics of painted turtles (Chrysemys picta) on the E. S. George Reserve in southeastern Michigan. Estimates of the degree of spatial genetic structuring were based on the proportion of total genotypic variance partitioned within and between subpopulations (inferred from hierarchical F-statistics based on variation at 18 protein loci), and in terms of gene correlations (co-ancestry among individuals derived from reproductive data on full-sib families of females nesting at specific nesting areas). Little variation in allele frequency was observed among turtles from different marshes (F_mt = 0.003), though significant variation was observed among turtles from different nesting areas associated with each marsh (F_nm = 0.046). Gene correlations among individuals within nesting areas varied greatly over years (0.032-0.171; mean = 0.069) and were negatively correlated to the proportion of females that successfully nested during each year. General concordance between independent estimates of genotypic correlations (i.e., F_nm derived from protein electrophoretic variation vs. mean co-ancestry) suggests that allozyme data, when collected over spatial scales consistent with species behavioral characteristics and reproductive ecology, may accurately reflect the apportionment of gene diversity within and among subpopulations. The magnitude and patterning of allelic variation among nesting areas and individuals appears to be primarily a function of gametic correlations among members of full-sib families, irrespective of the degree of gene flow or female nesting-site fidelity. Comparisons of genetic characteristics among 11 cohorts (1974-1984) revealed that heterozygosity (H) and inbreeding coefficients (F) varied greatly. Cohort estimates of H and F were correlated to female nesting success and to estimates of co-ancestry for the same years. Results clearly reflect the concomitant importance of ecological factors (principally the proportion of the female population that successfully produce offspring during each year) in determining the magnitude and patterning of gene correlations within and among groups, and to the genotypic composition of offspring born during each year.     

Examination of DNA sequences undergoing chromatin conformation changes at a variegating breakpoint in Drosophila melanogaster

Position effect variegation in Drosophila melanogaster is associated with the inability of certain genes to be correctly expressed in a proportion of cells, giving a mosaic phenotype. The lack of expression is thought to be due to alterations in the gene's chromatin structure due to its proximity to a region of heterochromatin. Because of the difficulties involved, there is little biochemical data to support the intuitively appealing model of heterochromatin spreading used to explain this phenomenon.Differences in restriction fragment length were used to distinguish DNA regions from either normal (non-position affected) or rearranged (position affected) chromosomes so as to examine possible changes in gene copy number and the effects of endogenous nucleases. DNA sequences at the breakpoint of In (1)w ^m4, which variegates for the white gene, were assayed under conditions where the chromatin conformation was altered using second site modifier mutations (Su(var) or En(var)). No change in the DNA sequerice copy number was observed at either chromosome breakpoint, relative to wild type, when either suppressor or enhancer mutations were present. Therefore copy number change, through differential polyploidization or somatic gene loss, is not affected by Su(var) or En(var) induced changes in the chromatin conformation.Initial experiments showed a gross difference in the sensitivity of DNA to endogenous nucleases that appeared associated with Su(var) and En(var) mutations. En(var) mutation bearing samples appeared delayed in the digestion, relative to Su(var). This differential sensitivity seemed to be genome-wide as there was no detectable difference between either breakpoint of In(1)w ^m4 or the sequences on the homologous w ^- chromosome. However, after isogenizing the genetic background, the previously noted difference between the Su(var) and En(var) mutations was eliminated. In studies dealing with nuclease digestion of chromatin, the isogenization of genetic background is essential before meaningful comparisons can be made.     

Numerical analyses of RAPD data highlight the origin of cultivated tagasaste (Chamaecytisus proliferus ssp. palmensis) in the Canary Islands

Random Amplified Polymorphic DNA (RAPD) was used to generate molecular markers to trace the origin of the fodder legume tagasaste (Chamaecytisus proliferus (L. fil.) Link ssp. palmensis (H. Christ) Kunkel) in the Canary Islands. Results from multivariate analyses of data through Two Way Indicator Species Analysis (TWINSPAN) and Detrended Correspondence Analysis (DECORANA) showed that genotypes collected on the island of La Palma exhibited a wider range of variation than those from the other islands. This supports the existing hypothesis that tagasaste originated on La Palma and emphasizes the importance of conserving and evaluating germ plasm from this island.     

Acceptor stem and anticodon RNA hairpin helix interactions with glutamine tRNA synthetase

The class I glutamine (Gln) tRNA synthetase interacts with the anticodon and acceptor stem of glutamine tRNA. RNA hairpin helices were designed to probe acceptor stem and anticodon stem-loop contacts. A seven-base pair RNA microhelix derived from the acceptor stem of tRNA^Gln was aminoacylated by Gln tRNA synthetase. Variants of the glutamine acceptor stem microhelix implicated the discriminator base as a major identity element for glutaminylation of the RNA helix. A second RNA microhelix representing the anticodon stem-loop competitively inhibited tRNA^Gln charging. However, the anticodon stem-loop microhelix did not enhance aminoacylation of the acceptor stem microhelix. Thus, transduction of the anticodon identity signal may require covalent continuity of the tRNA chain to trigger efficient aminoacylation.     

DNA fingerprint bands applied to linkage analysis with quantitative trait loci in chickens

Summary
An efficient approach to detect association between quantitative traits and bands of DNA fingerprint patterns uses intra-family tail analysis, which compares fingerprints of DNA mixes from individuals at the two tails of a phenotypic distribution. In analysis of 67 paternal half-sibs of a meat-type chicken family, of 57 sire bands generated by two probes, one sire-specific band (S6–6) was associated with abdominal fat deposition. The band effect was estimated by a linear model analysis to be 0–88 standard deviations, or about 30% of the family mean. The association between band S6–6 and abdominal fat was further examined by testing progeny of paternal half-sibs of the chickens which were used in the tail analysis, establishing genetic linkage between the DNA marker and a genetic locus affecting abdominal fat deposition.     

Assignment of five polymorphic ovine microsatellites to bovine syntenic groups

A panel of bovine somatic cell hybrids was used to map ovine microsatellites. Five of seven microsatellites were assigned to five bovine syntenic groups. These microsatellites were designated D5S10 (MAF23), D1S4 (MAF46), D13S1 (MAF18), D4S3 (MAF50), and DXS2 (MAF45), mapped to syntenic groups U3 (chromosome 5), U10 (chromosome 1), U11, U13, and the X chromosome, respectively. Two remaining sheep microsatellites amplified rodent DNA in the hybrid somatic cell panel, and were not assigned to bovine syntenic groups. Assignment of ovine-derived microsatellites to bovine syntenic groups provides additional evidence of the usefulness of microsatellites for mapping closely related species. The use of ovine and bovine microsatellites will aid in development of comparative genomic maps for these two species.     

Multiple origins of self-fertilization in tristylous Eichhornia paniculata (Pontederiaceae): Inferences from style morph and isozyme variation

Populations of Eichhornia paniculata (Pontederiaceae) exhibit a wide range of mating systems, from predominant outcrossing to high levels of self-fertilization. The origin of self-fertilization in this tristylous species is associated with the loss of style-length morphs from populations and the spread of self-pollinating, floral variants. We examined geographic variation in style morph and allozyme frequencies to determine whether the loss of style morphs and transition to selfing could have multiple origins in E. paniculata. Surveys of floral variation in 167 populations from six states in northeastern Brazil revealed that at least one style morph was absent from 29.3%. Non-trimorphic populations occurred in all states and ranged in frequency from 9% in Ceará to 68% in Alagoas. Selfing variants occurred in 8.5% and 55% of trimorphic and non-trimorphic populations, respectively, and were distributed among five of six states with primary concentrations in Alagoas and Pernambuco. A comparison of electrophoretic variation at 24 isozyme loci in 28 trimorphic, 13 dimorphic and 3 monomorphic populations indicated that non-trimorphic populations contained 84% of the allelic variation present in trimorphic populations and were markedly differentiated from one another. Analyses of genetic distance and the distribution of rare alleles indicated that non-trimorphic populations were often more similar to neighbouring trimorphic populations than to one another. Populations with selfing variants occurred at low frequency in three genetically distinct parts of the range. These results, in combination with genetic and morphological evidence suggest that style morphs are lost repeatedly from populations of E. paniculata and that selfing variants may have originated on at least three separate occasions in northeastern Brazil.     

Is drift ‘directional’? Unequal breeding sex ratio revisited

Unequal breeding sex ratio can significantly reduce effective population size, allowing a rare neutral allele to jump to a high frequency through genetic drift. However, this one-way alteration to allele frequency appears inconsistent with the concept that drift is non-directional. Based on binomial sampling distribution, this study developed a method to directly and exhaustively measure drift by calculating the mean deviation of change in allele frequency, then applied it to cases of unequal breeding sex ratio. The result shows that, under those cases, (1) the mean deviation can always be divided into two halves that are equal in size but opposite in direction; (2) each half consists of one or several categories represented by various allele proportions in the rare sex; (3) this proportion is another factor that determines the outcome of drift, in addition to effective population size and allele frequency; (4) drift is non-directional on a global scale, but whether an allele will drift up or down can be predicted based on the above factors. This method enables us to dissect every component of the expected change in allele frequency caused by drift and to find out the combined effect of population size, allele frequency and allele proportion in the rarer sex under neutrality but unequal breeding sex ratio.     

The maintenance of genetic diversity under host–parasite coevolution in finite,structured populations

As a corollary to the Red Queen hypothesis, host–parasite coevolution has been hypothesized to maintain genetic variation in both species. Recent theoretical work, however, suggests that reciprocal natural selection alone is insufficient to maintain variation at individual loci. As highlighted by our brief review of the theoretical literature, models of host–parasite coevolution often vary along multiple axes (e.g. inclusion of ecological feedbacks or abiotic selection mosaics), complicating a comprehensive understanding of the effects of interacting evolutionary processes on diversity. Here we develop a series of comparable models to explore the effect of interactions between spatial structures and antagonistic coevolution on genetic diversity. Using a matching alleles model in finite populations connected by migration, we find that, in contrast to panmictic populations, coevolution in a spatially structured environment can maintain genetic variation relative to neutral expectations with migration alone. These results demonstrate that geographic structure is essential for understanding the effect of coevolution on biological diversity.