Nonrandom mating in an open-pollinated maize population.

Randomness of fertilization was studied in an open-pollinated population of maize (Zea mays L.) through allozyme assays of seedlings from open-pollinated seeds produced on both tasseled and detasseled plants. Mixed-mating-model estimates of the amount of outcrossing (t) were not significantly different from t = 1.00 for four enzyme loci ( Adh1, Idh2, Got1 and Acp1), indicating that fertilizations were at random in the population. However, for loci Prx1 and Est4 , estimates of t were significantly smaller than unity-0.80 and 0.70 for tasseled plants and 0.81 and 0.80 for detasseled plants. The excesses of homogametic fertilizations detected on the detasseled plants could not have been due to selffertilization, s = 1 - t, because the detasseled plants shed no pollen. Analyses of allelic frequencies in the pollen that produced seed on the detasseled plants established that different maternal plants sampled genetically different populations of pollen from the outcross pollen pool. It was suggested that the causes of the differential sampling were temporal variation in the pollen pool, and/or gametophytic selection, correlated with marker-locus genotype. Two-, three- and four-locus interactions among the marker loci were often statistically significant, indicating that the factors responsible for the nonrandom gametic unions observed in the maize population studied were complexly interactive.

Estimation of life cycle components of selection in an experimental plant population.

Viability and fertility components of selection associated with linkage blocks marked by four electrophoretically detectable loci were estimated in an experimental population of barley [Composite Cross V (CCV)]. The intensity of selection affecting the distribution of pollen types in the outcross pool was also estimated and comparisons were made between the selective values of genes in the pools of uniting ovules and pollen. The estimates show that selection was intense at various stages of the life cycle and that viability and fertility components often opposed one another. Estimates of viability and fertility components of selection were also extended to the three-locus level. The multilocus estimates reveal large differences in viability and fertility among homozygous genotypes. It is concluded that strong selection operates at all life cycle stages in CCV, although often in differing directions.

Analysis of complex allozyme polymorphisms in a barley population.

Genotypes of 68,230 individuals taken from 10 generations (F(4)-F(6), F(14)-F(17), F(24)-F(26)) of an experimental population of barley were determined for four esterase loci. The results show that frequencies of gametic ditypes changed significantly over generations and that striking gametic phase disequilibrium developed within a few generations for each of the six pairwise combinations of loci which were monitored. The complex behavior of these four enzyme loci in the population is attributed to interactions between selection and restriction of recombination resulting from the effects of linkage and/or inbreeding.

The effect of selection on esterase allozymes in a barley population.

Changes in gene and genotypic frequencies at four esterase loci were monitored over 25 generations in Composite Cross V, an experimental population of barley, to obtain experimental evidence concerning the balance of forces responsible for: (1) the marked differences in allelic frequencies among barleys from different ecogeographical regions of the world; and (2) the extensive allelic variation found within local populations of barley. Analyses of the highly significant changes in allelic frequencies which occurred in CCV showed they were due to directional selection favoring particular alleles and not to mutation, migration or genetic drift. The results show that intense balancing selection, featuring consistent excesses of heterozygotes, also occurred in CCV. It is concluded that among the factors of neo-Darwinian evolution, natural selection plays the predominant role in determining the observed patterns of allelic variation in the barley species as a whole.

Further analysis of complex allozyme polymorphisms in a barley population.

New theory has recently been developed for two-locus models. In the light of this theory, an earlier analysis of esterase allozyme data from an experimental barley population has been modified to take proper account of initial gametic phase (linkage) disequilibria. The results show that the directions in which two-locus genotypic frequencies deviated from products of one-locus frequencies in this population followed those predicted by neutral descent theory. The observed departures were, however, much larger in size than predicted by the new descent measure theory, indicating that selection is operating in the population.

Mutation Rates for Enzyme and Morphological Loci in Barley ( HORDEUM VULGARE L.).

Spontaneous mutation rates were estimated by assaying 84,126 seedlings of a highly homozygous barley line (isogenic line 2025) for five enzyme loci. No mutants were observed in 841,260 allele replications. This result excludes, at probability level 0.95, a spontaneous mutation rate larger than 3.56 x 10(-6)/locus/gamete/generation for these enzyme loci. Isogenic line 2025 also was scored for mutants at four loci governing morphological variants. No mutants were observed in 3,386,850 allele replications which indicates that the upper bound for the mutation rate for these loci is 8.85 x 10(-7). It was concluded that, even though spontaneous mutation has been important in creating variability in the barley species at the loci scored, the rate is too low to have much affect on the short-term dynamics of barley populations.

RFLP genetic linkage maps from four F(2.3) populations and a joinmap of Gossypium hirsutum L.

An RFLP genetic linkage joinmap was constructed from four different mapping populations of cotton ( Gossypium hirsutum L.). Genetic maps from two of the four populations have been previously reported. The third genetic map was constructed from 199 bulk-sampled plots of an F(2.3) (HQ95-6x'MD51ne') population. The map comprises 83 loci mapped to 24 linkage groups with an average distance between markers of 10.0 centiMorgan (cM), covering 830.1 cM or approximately 18% of the genome. The fourth genetic map was developed from 155 bulk-sampled plots of an F(2.3) (119- 5 sub-okrax'MD51ne') population. This map comprises 56 loci mapped to 16 linkage groups with an average distance between markers of 9.3 cM, covering 520.4 cM or approximately 11% of the cotton genome. A core of 104 cDNA probes was shared between populations, yielding 111 RFLP loci. The constructed genetic linkage joinmap from the above four populations comprises 284 loci mapped to 47 linkage groups with the average distance between markers of 5.3 cM, covering 1,502.6 cM or approximately 31% of the total recombinational length of the cotton genome. The linkage groups contained from 2 to 54 loci each and ranged in distance from 1.0 to 142.6 cM. The joinmap provided further knowledge of competitive chromosome arrangement, parental relationships, gene order, and increased the potential to map genes for the improvement of the cotton crop. This is the first genetic linkage joinmap assembled in G. hirsutum with a core of RFLP markers assayed on different genetic backgrounds of cotton populations (Acala, Delta, and Texas plain). Research is ongoing for the identification of quantitative trait loci for agronomic, physiological and fiber quality traits on these maps, and the identification of RFLP loci lineage for G. hirsutum from its diploid progenitors (the A and D genomes).

Worldwide patterns of genetic variation among four esterase loci in Barley (Hordeum vulgare L.).

Electrophoretic assays of 1506 accessions of domestic (Hordeum vulgare L.) and wild (H. spontaneum Koch.) barley, maintained in the USDA World Barley Collection, led to the following conclusions: (1) worldwide the four esterase loci, Est 1, Est 2, Est 3, and Est 4, have a minimum of 7, 12, 6, and 7 alleles, respectively; (2) little or no genetic differentation has developed between H. vulgare and H. spontaneum at these four esterase loci; (3) substantial genetic polymorphism and heterozygosity occur within many of the accessions despite the heavy inbreeding which results from the mating system of predominant self fertilization and from genetic drift associated with maintenance in small populations; (4) patterns of geographical distribution of alleles at these four loci are not at random over both small and large geographical areas, including differences on a continental scale; (5) four among 16 four-locus combinations of alleles are found in excess and all other combinations occur in deficiency on a worldwide basis.

Associations between quantitative traits and enzyme loci in the F2 population of a maize hybrid.

Univariate and multivariate analyses were used to identify associations between eight enzyme marker loci and 11 quantitative traits of maize (Zea mays L.). The material analyzed included inbred lines Wf9 and Pa405, single-cross hybrid Wf9 X Pa405, and the F2 generation of the selfed single-cross hybrid. Each enzyme locus assayed was associated with at least one quantitative trait, and all quantitative traits were associated with genotypes at particular enzyme loci. Significant associations also were found between the level of heterozygosity per individual and nine of 11 quantitative traits. The total contribution to heterosis, for seed yield per plant, of genes linked with the eight enzyme loci, was 27% of the F2 mean and 18% of the difference in mean between the F1 hybrid and the inbred parents. Genes linked with Glu1 accounted for nearly one third of the total dominance effect detected by the eight enzyme loci. The chromosome segments marked by loci with significant effects on seed yield were markedly overdominant. The large heterotic effects of chromosome segments marked by particular loci suggest that enzyme loci could be used to help transfer genes responsible for heterosis to inbred lines. We conclude that analyses of additional inbred lines, F1 hybrids, and F2 populations in more environments will halp identify specific associations between enzyme loci, or chromosome segments which they mark, and important agronomic traits.

Evolutionary Changes in the Mating System of an Experimental Population of Barley (Hordeum vulgare L.).

An analysis is presented of the reproductive cycle in an experimental population of barley. The experimental design included growing three different generations of the population in the same year and environment, thus permitting an assessment of the mating system at three stages in the evolutionary history of the population, unconfounded by environmental differences. Gene frequencies sometimes differed significantly in adults and in the effective pollen pool. It was also found that out-crossing rate more than doubled during the twenty generations spanned by the study. These results provide evidence for selection during reproductive phases of the life cycle. They also demonstrate that evolution in this predominantly self-pollinating population was in the direction of increased recombinational potential.

A multilocus estimator of mating system parameters in plant populations.

A multilocus method of estimating mating system parameters in populations is presented that recovers information from classification over multiple loci that single-locus estimators do not detect. It is shown that the multilocus estimator provides a standard of reference (null hypothesis) that can be useful in analyzing the effects of factors such as population heterogeneity and post-mating zygotic selection on the transmission of genetic information at the population level.

Is the gene the unit of selection? Evidence from two experimental plant populations.

The dynamics of gametic frequency change have been analyzed in two experimental plant populations. Individual plants were scored for their genotype at four enzyme loci, and four-locus joint gametic frequencies were estimated from the genotypic data over generations. Striking correlations developed in allelic state over loci, including correlations between nonlinked loci, as these populations evolved. Furthermore, the same pair of four-locus complementary gametic types came into marked excess in the late generations of both populations. The results demonstrate that natural selection acted to structure the genetic resources of these populations into sets of highly interacting, coadapted gene complexes. They also provide evidence, at the level of the gene, that selection operating on correlated multilocus units is an important determinant of population structure.

Allozyme polymorphisms within and among open-pollinated and adapted exotic populations of maize.

Twelve U.S. Corn Belt open-pollinated and five adapted exotic populations of maize (Zea mays L.) were assayed for allozyme (allele) variation at 13 enzyme marker loci. Extensive allozyme variability was observed in all populations studied. No locus was monomorphic over all populations. Each of the lociIdh2, Got1, Mdh2, Pgd1, andPgd2 expressed two allozymes over all populations,Adh1, Acp1, Prx1, andEst1 each had three allozymes present,Est4, Glu1, andEnp1 had five allozymes, andAcp4 had six allozymes present. Significant deviations of genotypic frequencies were detected from Hardy-Weinberg equilibrium frequencies and 94% of average Fixation Index values indicated heterozygote deficiencies, which suggested that nonrandom mating and/or natural selection favoring homozygotes were possible factors affecting the maintenance or loss of genetic variability marked by these enzyme loci. Genetic distance and cluster analyses indicated that the observed genetic variability at the 13 enzyme loci was closely related to 'Dent' and 'Flint' types of maize.

Associations between isozyme phenotypes and environment in the slender wild oat (Avena barbata) in Israel.

Collections from 31 populations of A. barbata from diverse habitats in Israel were assayed electrophoretically for seven enzyme systems. Phenotype frequencies were scored in nine enzyme zones, probably representing 27 loci, to determine isozyme variability within and among populations. Many different isozyme phenotypes were found in all of the populations; also the array of isozyme phenotypes found in each population differed distinctly from that found in each other population. Overlays of phenotypic frequencies on map locations showed that isozyme variability is distributed in mosaic patterns not related to geographical distance. Principal-component and multiple-regression analyses revealed that temperature and moisture-related variables are significantly correlated with particular isozyme phenotypes. Further, the mosaic patterns of isozyme variation were found to correspond closely to mosaic patterns of the habitat. This structuring of the genetic variability into multilocus combinations was attributed to the combined effects of directional and diversifying selection. Comparisons of patterns and extent of genetic variation in Israel and California led to the conclusion that the evolution of 'ecotypes,' each adapted to a specific habitat and marked by a particular set of enzyme alleles, has proceeded further in Israel, where A. barbata is endemic, than in California, where it is a recent introduction.

Evidence for coadaptation in Avena barbata.

Two populations of Avena barbata have been sampled intensively and allelic frequencies have been determined on a fine grid for six enzyme loci. The distribution of alleles among microniches within the location occupied by one of these populations (CSA-1) was correlated with environment following patterns of macro- and microdifferentiation previously described in California. Multilocus analyses revealed striking gameticphase disequilibrium for both linked and unlinked loci, and also that two balanced five-locus gametic types, one associated with mesic conditions and the other with more xeric conditions, are in substantial excess at the CSA-1 location. Evidence is presented that these two gametic types are maintained by selection acting on two coadapted combinations of alleles, held together by restriction of recombination resulting from linkage and/or inbreeding. Gametic-phase disequilibrium featuring different allelic complexes were found in each of three microniches in the second population (Geyserville), which occupies a site that is environmentally atypical for A. barbata.

Establishment of molecular markers and linkage groups in two F2 populations of Upland cotton.

Two F2 populations of cotton (Gossypium hirsutum L.) from the crosses of HS46 x MARCABUCAG8US-1-88 (MAR) and HS46 x Pee Dee 5363 (PD5363) were characterized for restriction fragment length polymorphisms (RFLPs) using DNA probes. Seventy-three probe/enzyme combinations were used in the HS46 x MAR population analysis, which resulted in 42 informative polymorphic fragments. These 42 moleclar markers represented 26 polymorphic loci, which consisted of 15 codominant and 11 dominant (+/-) genotypes. Chi-square analyses of these loci fit expected genotypic ratios of 1∶2∶1 and 3∶1, respectively An analysis of these loci with the MAPMAKER program resulted in the establishment of four linkage groups A, B, C, and D with 4,2,2, and 2 loci, respectively, as well as 16 unlinked loci. Six probe-enzyme combinations were assayed on the HS46 x PD5363 population, which resulted in 11 informative polymorphic fragments. These 11 fragments represented 6 polymorphic loci, 1 dominant (+/-) and 5 codominant genotypes. The MAPMAKER analysis of these loci yielded 2 linked loci. Thus, a total of 53 polymorphic fragments and 32 polymorphic loci, representing five linkage groups, were identified among the two families.