Gene flow across tribal barriers and its effect among the Amazonian Içana river Indians

Demographic information was obtained from 622 individuals of five communities of primarily Baniwa Amerindians living near the Içana river in Brazil. Four of these populations, plus another from the same area, were also studied genetically. The latter investigation included the blood and, in some cases, saliva of 531 subjects, variously tested in relation to 40 genetic systems. Demographically these groups are characterized by young age, high intertribal admixture, low non-Indian admixture, high exogamy but low marital distance and high inbreeding, high fertility but low variance in offspring number, and relatively low mortality. Their gene pool shows a peptidase B variant (PEPB^2BAN1) and “private” polymorphism of carbonic anhydrase₂ (CA₂^BAN1) until now observed only among them. Other distinctive characteristics are the low frequencies of L^NS (0.08), L^Ns (0.09), R^z (0.01), R^O or r (0.02), ACP^A (0.08), GALT^D (0.01), and the relatively high prevalences of Gm^1;11,13,16 (0.05) and Gc¹ (0.82). Tf^Dehi occurs with a low prevalence (0.01). Genetic distance analysis reveals that the one Baniwa sample by history comprised of minimally admixed individuals is quite similar genetically to the Wapishana, another Arawak-speaking tribe some 900 km to the east, and that the genetic distances between the Baniwa communities reflect the amount of historical admixture in a way that indicates which should be excluded from considerations of intertribal genetic distances. Finally, the genetic relation of the Baniwa to the nearby tribes is examined.     

The Genetic Structure of a Tribal Population, the Yanomama Indians. Xv. Patterns Inferred by Autocorrelation Analysis

Fifteen allele frequencies have previously been determined for 50 villages of the Yanomama, an Amerindian tribe from southern Venezuela and northern Brazil. These frequencies were subjected to spatial autocorrelation analysis to investigate their population structure. There are significant spatial patterns for most allele frequencies. Clinical patterns, investigated by one-dimensional and directional spatial correlograms, were relatively few in number and were moderate in strength. Overall, however, there is a marked decline in genetic similarity with geographic distance. The results are compatible with a hierarchic population structure superimposed on the geography, and generated by a stochastic fission-fusion model of village propagation, followed by localized gene flow. Strong temporal autocorrelations of allele frequencies based on linguistic-historical distances representing time since divergence were also found. There appears to be a stronger relation between geography and linguistic-historical hierarchic subdivisions than between either feature and genetic distances. These findings confirm by different approaches the results of earlier analyses concerning the important roles of both stochastic and social factors in determining village allele frequencies and the occurrence within this tribe of some allele frequency clines most likely due to the operation of chance historical processes.     

Maximum Likelihood Analysis of Population Differences in Allelic Frequencies

Statistical techniques are presented for the analysis of geographic variation in allelic frequencies. Likelihood ratio test criteria are derived from a multinominal sampling distribution, and are used to answer three questions. (1) Are there geographic differences in allelic frequencies? (2) Are population differences in allelic frequencies associated with environmental differences? (3) Is there any residual "lack of fit" variation among populations, after accounting for that variation associated with environmental differences? The two- and three-allele cases are explicitly treated, and the extension to more alleles is indicated.     

THE IMPACT OF ELECTROPHORETIC GENOTYPE ON LIFE HISTORY TRAITS IN PINUS TAEDA

Reports of positive associations between allozymic heterozygosity and measures of fitness are routine, but it has not been possible to distinguish between the two preeminent explanations of the phenomenon, dominance and overdominance. We tested several of the assumptions of these hypotheses in our study of the relationship between electrophoretic genotype and three life history traits in loblolly pines (Pinus taeda L.). Traits examined included the survival and growth of selfed and outcrossed progeny of 45 maternal trees, and maternal fecundity, measured as the number of surviving progeny per mother tree. Inbreeding depression was severe; the relative fitness of the selfed progeny was only 8% that of the outcrossed progeny. We found a heterozygote fecundity advantage, which should have resulted in an excess of rare alleles in the progeny. Instead, there was evidence of severe survival selection against rare alleles in both heterozygous and homozygous forms. The deficit of rare alleles averaged 69 and 50% in the selfed and outcrossed progeny, respectively. The one allele in the sample that we should have suspected of being maintained by overdominance (a PGI2 mid-frequency allele) appeared to be overdominant for outcrossed height growth and probably for fecundity as well. Multiple-locus genotype explained very little of the variation in growth, however, and rather than seeing evidence for overdominance as a force in maintaining most of the observed polymorphism, we were left to explain, in the face of the severe survival selection, why the rare alleles were present at all. Projection of the stand into the future through computer simulation showed how balancing selection acting on differential growth, fecundity, and mortality among genotypes could, over the life of the stand, account for the maintenance of the rare alleles in the population.     

THE FITNESS CONSEQUENCES OF MULTIPLE-LOCUS HETEROZYGOSITY: THE RELATIONSHIP BETWEEN HETEROZYGOSITY AND GROWTH RATE IN PITCH PINE (PINUS RIGIDA MILL.)

Positive correlations between measures of “fitness” and the number of electrophoretic loci for which an individual is heterozygous have been observed in many species. Two major hypotheses have been proposed to explain this phenomenon: inbreeding depression and overdominance. Until recently, there has been no way to distinguish between these hypotheses. The overdominance model devised by Smouse (1986) is used here in a reanalysis of Ledig et al.‘s (1983) study of heterozygosity and growth rate in eight populations of pitch pine and is contrasted with an inbreeding-depression analysis. Ledig et al. (1983) regressed mean growth rate per heterozygosity class on the number of heterozygous loci, a method of analysis which, although it points to general trends in the data, does not differentiate between hypotheses. The correlations they obtained in four populations were significant only because regressing on the means eliminates most of the sum of squares for error and does not weight the unequally sized heterozygosity classes. Reanalysis of Ledig et al.‘s data using individuals, not means, showed no significant correlations between heterozygosity and fitness. A major assumption of Smouse's overdominance model is that genetic polymorphism is in part a reflection of selection for heterozygotes at genetic equlibrium. The homozygote for the most frequent allele at a locus should be more fit than a homozygote for a less frequent allele, with the heterozygote superior to both homozygotes. Smouse's model predicts a negative, linear relationship between fitness and “adaptive distance,” a variable that for a heterozygote is zero and for homozygotes is equal to the inverse of the frequency of the corresponding allele. The adaptive-distance model accounted for between 15% and 50% of the variation in growth rate within eight P. rigida population samples by accounting for genotypic differences at eight polymorphic loci. This is over twice as much of the variation in growth rate accounted for by Ledig et al.'s (1983) analysis using individuals. Significant correlations were found between adaptive distance and growth rate in four of the eight populations, but in only two of the populations were more of the partial coefficients negative than positive, as would be predicted by the overdominance hypothesis. The remaining two populations in which correlations were significant did not lend themselves to such clear-cut interpretation, as the majority of the partial coefficients were positive. Positive partial coefficients indicate that the growth rate of the heterozygote is inferior to that of at least one of the homozygotes. The adaptive-distance analysis provides evidence that specific genotypes do play a role in determining growth rate in pitch pine. The correlation between growth rate and adaptive distance increased significantly with the age of the population, possibly reflecting competition subsequent to crown closure.     

THE FITNESS CONSEQUENCES OF MULTIPLE-LOCUS HETEROZYGOSITY UNDER THE MULTIPLICATIVE OVERDOMINANCE AND INBREEDING DEPRESSION MODELS

There is a growing body of literature suggesting that the fitness of an individual increases with the observed number of heterozygous loci. Broad theoretical considerations indicate that under various sorts of balancing selection, this is what one should generally expect in a population of multiple-locus genotypes. To date, however, it has not been possible to distinguish between two potential explanations of the phenomenon. The first explanation is that the loci examined are themselves responsible for the fitness differences observed (or, equivalents, are very closely linked to those that do). The genetic variation in question is thought to be maintained in polymorphic equilibrium by some form of balancing selection. The second explanation assumes that the observed loci are themselves selectively irrelevant but that their heterozygosity reflects that of the total genome. Genomic heterozygosity is thought to be predictive of fitness, being an obverse measure of generalized inbreeding depression. We provide a formal derivation of an explicit relationship between fitness and multiple-locus genotype for a simple form of the first explanation, the multiplicative overdominance model. The inbreeding depression model is a degenerate special case of this more general formulation. A formal estimation and testing framework is constructed that should facilitate evaluation of the two models with empiric data on heterozygosity and fitness.     

Parentage analysis in Chamaelirium luteum (L.) Gray (Liliaceae): why do some males have higher reproductive contributions?

Much of the contemporary study of adaptation in natural populations involves the regression of some component of fitness, usually survivorship or viability, on one or more characters of interest. It is difficult to apply this approach to measures of paternal reproduction, however, because paternity is typically estimated indirectly from genetic markers, rather than being measured directly from progeny counts. Here, we present maximum likelihood methods for modelling relative male reproductive success as a log-linear function of one or more potentially predictive features, as well as providing a framework for the assessment of pairwise (male:female) effects, as they affect male reproductive performance. We also provide nonparametric statistical tests for alternative models. Using this formulation, we examine the impact of inflorescence morphology on male reproductive success in Chamaelirium luteum L., and we also assess the importance of intermate distances between males and particular females. While male reproductive success and male inflorescence morphology are both quite variable, reproductive morphology does not appear to predict male reproductive success in this study. Intermate distance is an extremely effective predictor of pairwise success, however; but averaged over females, there is almost no net effect for different males.     

Paternity analysis in Ophiopogon xylorrhizus Wang et Tai (Liliaceae s.l.): selfing assures reproductive success

We assessed the utility of codominant allozyme markers for paternity analysis in natural populations of Ophiopogon xylorrhizus (Liliaceae s. l.) by means of likelihood‐based approach. In the three independent natural stands of 46 individual plants with 354 seeds of O. xylorrhizus (Liliaceae s. l), we assigned 319 seeds to a single individual with the highest LOD score. Among these, 179 seeds had a significant D‐value. An individual (acting either as female or male) mateds with an average of one other individual on average to reproduce, except when it self‐pollinated. If an individual that neither donated pollen to other individuals nor selfed, being strictly a pollen acceptor, the number of offspring produced was half that produced by as many offspring as individuals which both provided and accepted pollen (4.1 vs. 9.5). Significant positive relationships were found in all the three stands between the number of self‐pollen an individual produced and the number of seeds that individual set. Male outcrossing reproductive success was unevenly varied, both in amount and spatial location. The individuals located in the centre of the stand and the individuals closest to each other had higher male outcrossing reproductive success than those at the edge of the stand. Male selfing reproductive success showed a similar pattern, with an even distribution of female reproductive output. Most mating events were took place within 20 m. The distance between mates mirrored the active ranges of pollinators, and shaped the fine‐scale population spatial genetic spatial structure. The results indicate that selfing assures reproductive success in O. xylorrhizus.