Selection in Reference to Biological Groups. VI. Use of Extreme Forms of Nonrandom Groups to Increase Selection Efficiency

The strategy of using non-random groups to increase the efficiency of truncation selection is discussed. The present study, which considers extreme forms of non-rnadom groups, complements a previous study involving full-sib groups. It is shown that of the two kinds of non-randomness, i.e. that due to homozygosity or that due to homogeneity (as represented by cloning), the latter is the most effective. This suggests that with those plant crops in which intense competition among plants exists, use of clonal propagation to produce non-random groups should be investigated.     

Selection bias in quantitative trait loci mapping

A simulation study was performed to see whether selection affected quantitative trait loci (QTL) mapping. Populations under random selection, under selection among full-sib families, and under selection within a full-sib family were simulated each with heritability of 0.3, 0.5, and 0.7. They were analyzed with the marker spacing of 10 cM and 20 cM. The accuracy for QTL detection decreased for the populations under selection within full-sib family. Estimates of QTL effects and positions differed (P < .05) from their input values. The problems could be ignored when mapping a QTL for the populations under selection among full-sib families. A large heritability helped reduction of such problems. When the animals were selected within a full-sib family, the QTL was detected for the populations with heritability of 0.5 or larger using the marker spacing of 10 cM, and with heritability of 0.7 using the marker spacing of 20 cM. This study implied that when selection was introduced, the accuracy for QTL detection decreased and the estimates of QTL effects were biased. A caution was warranted on the decision of data (including selected animals to be genotyped) for QTL mapping.     

A theory of natural selection incorporating interaction among individuals. III. Use of random groups of inbred individuals

The present series of studies attempts to accommodate interaction among individuals in evolutionary theory. The interaction phenomenon is characterized by two dimensions (direct and associate) of gene activity. For optimal selection results, a balance between the two dimensions must occur. In the first paper of the series, it was shown that random interactions resulted in an unbalanced selection response in that the direct, but not associate, effects were included in the expression for gene frequency change. The next three papers of the series (II, III and IV) were designed to determine whether or not selection with life-history models that involved non-random interactions would be useful in ameliorating the problem of selection balance.In the present study, non-randomness is generated by restricting interactions to inbred individuals. It is demonstrated that this form of non-random gene association within interacting genotypes does not improve selection balance. Thus restricting interaction to groups of inbred individuals does not result in the introduction of associate effects into the expression for gene frequency change. It is shown that inbreeding in the base population merely accelerates the unbalanced response normally occurring when selection operates on random, non-inbred individuals.     

Individual and group selection with competition

Summary The response of a randomly mating population which is expected to follow selection of phenotypic units, comprising individuals or groups whose members have an arbitrary degree of relatedness, was formulated using a model which included additive and dominance competition effects. The derivation involved three steps. Twenty-two quadratic components were defined, six describing individual (direct) and neighbor (associate) effects, and 16 describing direct by associate interactions for different loci, for single loci with different alleles, and for identical alleles. Six covariances between pairs of individual phenotypes and three of individuals with their offspring were defined according to whether or not their direct or associate genotypes are common, and expressed in terms of the quadratic components. Finally, variances of selection units of different types and their covariance with their offspring were expressed as compounds of these individual covariances. Explicit formulations for mass, clonal and full-sib selection show that without constraints on the quadratic components, and hence on the magnitude and type of competition operative, no predictions as to the relative efficiencies of these three methods can be made.     

A theory of natural selection incorporating interaction among individuals. II. Use of related groups

The present series of studies attempts to accommodate interaction among individuals in evolutionary theory. The interaction phenomenon is characterized by two dimensions (direct and associate) of gene activity. For optimal selection results, a balance between the two dimensions must occur. In the first paper of the series, it was shown that random interactions resulted in an unbalanced selection response. The expression for gene frequency change involved direct, but not associate, effects. The next three papers of the series (II, III and IV) are designed to explore the possibility that restricting interactions to certain non-random patterns may ameliorate the problem of selection balance.In the present study the interactions are restricted to related individuals in a population that is in Hardy-Weinberg equilibrium. A preliminary analysis in which interactions are restricted to full-sibs is made. This analysis is extended to the more general case in which interactions occur among related individuals of any class whose coefficient of relationship is measured by ‘r’. The classical pairwise interaction results of Hamilton are verified and extended to include interactions among individuals in groups of arbitrary size, n.Restricting interactions to related individuals tends to improve the condition of selection balance. It does this by introducing associate effects into the expression for gene frequency change. The extent to which this is accomplished is a function of the coefficient of relationship (r), and the number of interacting genotypes.     

A theory of natural selection incorporating interaction among individuals. IV. Use of related groups of inbred individuals

The present series of papers attempts to accommodate interaction among individuals in evolutionary theory. The interaction phenomenon is genetically characterized by two dimensions (direct and associate) of gene activity. For optimal selection results, a balance between the two dimensions must occur. In the first paper of the series, it was shown that random interactions resulted in an unbalanced selection response in that the direct, but not associate, effects were included in the expression for gene frequency change. The next three papers of the series (II, III and IV) were designed to determine whether or not selection with life-history models that involved non-random interactions would be useful in ameliorating the problem of selection balance.In the present study, two kinds of non-random gene association are analyzed jointly by restricting interactions to related individuals that are derived from inbred base populations. The analyses are generalized to accommodate heterogeneous as well as homogeneous groups of interacting individuals. The joint contributions of inbreeding and consanguinity to selection response are analysed by use of the nine gene-identity parameters devised by Harris.It is demonstrated that consanguinity alone or in conjunction with inbreeding does improve selection balance. However, inbreeding alone does not. Also, the influence of inbreeding is not dependent on group size, whereas the influence of consanguinity is conditioned by the size of the group. Thus, by introducing associate effects into the selection process, the use of related groups can provide the genetical bases for the evolution of social behavior phenomena such as altruism.     

Choice of Female Groups by Male Mosquitofish (Gambusia holbrooki)

Even though females are usually more selective in choosing their mates, males are also capable of exercising mate choice. Despite the large body of evidence on the individual features preferred in sexual selection, little attention has been devoted to the first stage of male–female interaction. As poeciliid fish are known to be social, in the wild, initially mate choice may concern a preliminary selection among shoals. Only after this primary choice, males may subsequently direct their attention to a specific mate. We observed spontaneous preference of male mosquitofish (Gambusia holbrooki) when choosing between groups differing in size and sex ratio. In partial agreement with our predictions, males preferred to join a group of females rather than an isolated one (expt 1) and the larger group when two female groups were presented (expt 2). An all‐female group was preferred to a mixed‐sex group (expt 3), whereas no preference was observed when the two mixed‐sex groups differed in the number of males (expt 4) or in the size of the males (expt 5). These results suggest that male mosquitofish are capable of discriminating among different quantities of individuals within a group and use such information to select among groups in order to optimize the likelihood of successful matings.     

Comparison of selection methods for improvement of the population hybrid

Summary The objective of this study was to compare several selection procedures with respect to expected genetic gain in the population hybrid across a range of initial allelic frequencies, degrees of dominance, and environmental variances. The methods compared were intrapopulation recurrent selection using full-sib or S₁ families, full-sib and two half-sib reciprocal recurrent selection procedures, and convergent improvement applied to populations. Comparisons were made by calculating expected allelic frequency changes for each method. The optimal selection method for a given set of allelic frequencies and degree of dominance depended little on the environmental variance. Partly because of its short cycle, full-sib intrapopulation selection was the most effective method for the majority of allelic frequency combinations when the degree of dominance was small and an off-season nursery could be used to make recombinations. With larger values for the degree of dominance, S₁ and reciprocal full-sib methods became optimal, the former method especially when favorable alleles had a high frequency and the latter when populations were highly divergent. When off-season nursery use was restricted to making self-pollinations or was absent, S₁ selection was optimal for the majority of allelic frequency combinations. Convergent improvement was superior only for extremely divergent allelic frequencies and then only when the degree of dominance was less than 0.10. Half-sib reciprocal methods were never optimal, although the gain for the standard half-sib reciprocal procedure differed little from that of full-sib reciprocal selection when the degree of dominance was 0.75.     

A theory of natural selection incorporating interaction among individuals. V. Use of random synchronized groups

In the first paper of the current series, (I), a complex interaction model capable of describing any kind of interaction among individuals was developed. However, selection operating on random groups with regard to this model yielded short- and long-term results which were unbalanced. In subsequent kin-selection papers (II, III, and IV), a systematic analysis demonstrated that use of non-random groups could partially solve the balance problem.The present study is the first of several to employ a different approach to the problem of accommodating interaction. This approach involves changing the life-history model itself in such a way that the fitness components of individuals within groups are synchronized. Synchronization of fitness components produces total fitness values which are symmetrical. In the present study, selection operating on random groups for a model having symmetrical fitness values is evaluated for both balance and efficiency. It is demonstrated that the selection response is balanced and yields short- and long-term optimum results, but under a variety of conditions the efficiency can be low.     

Effect of selection against deleterious mutations on the decline in heterozygosity at neutral loci in closely inbreeding populations.

Transition matrices for selfing and full-sib mating were derived to investigate the effect of selection against deleterious mutations on the process of inbreeding at a linked neutral locus. Selection was allowed to act within lines only (selection type I) or equally within and between lines (type II). For selfing lines under selection type I, inbreeding is always retarded, the retardation being determined by the recombination fraction between the neutral and selected loci and the inbreeding depression from the selected locus, irrespective of the selection coefficient (s) and dominance coefficient (h) of the mutant allele. For selfing under selection type II or full-sib mating under both selection types, inbreeding is delayed by weak selection (small s and sh), due to the associative overdominance created at the neutral locus, and accelerated by strong selection, due to the elevated differential contributions between alternative alleles at the neutral locus within individuals and between lines (for selection type II). For multiple fitness loci under selection, stochastic simulations were run for populations with selfing, full-sib mating, and random mating, using empirical estimates of mutation parameters and inbreeding load in Drosophila. The simulations results are in general compatible with empirical observations.     

A theory of natural selection incorporating interaction among individuals. VI. Use of non-random synchronized groups

The previous study, (V), in this series approached the problem of accommodating interaction among individuals in a population by considering a life-history model which resulted in the synchronization of fitness components (viability and fecundity) for members within groups of interacting individuals. It was shown that such synchronization resulted in symmetric fitness values, and that selection operating on symmetric fitness values produced optimum short- and long-term results. However, it was also shown that selection operating on the random groups of the model could be inefficient.The present paper demonstrates that use of non-random (related) groups can increase the efficiency of symmetric selection without destroying its short-term balance. The consequences of long-term selection are more complicated and depend on the complexity of the genetic model.     

Trait mean depression for second-generation inbred strawberry populations with and without parent selection

 Strawberry genotypes selected for superior fruit yield or chosen at random from first-generation self, full-sib, and half-sib populations were crossed to provide second-generation inbred progenies and composite cross-fertilized control populations. Mean yields for inbred offspring from crosses among selected parents exceeded those from the offspring of unselected parents by 87%, 23%, and 37% for self, full-sib, and half-sib populations, respectively; yields for offspring from unrelated crosses among selected parents were 54% larger than those for crosses among unselected parents. Selection for yield also resulted in significant correlated response for fruit number and plant diameter. Mean yields for second-generation half-sib and full-sib offspring from selected parents were greater than those for offspring from the unselected but non-inbred control population. This suggests that selection can be a powerful force in counteracting most of the inbreeding depression expected in cross-fertilized strawberry breeding programs. Selection treatment× inbreeding rate interactions were non-significant for all traits; thus, selection among partially inbred offspring did not have a large effect on the rate of genetic progress. Differential realized selection intensity among individuals with differing levels of homozygosity accumulated due to inbreeding is suggested as the most likely explanation for the absence of association between pedigree inbreeding coefficients and cross performance detected previously in strawberry. Received: 21 July 1996 / Accepted: 7 March 1997     

Phenotypic and Evolutionary Consequences of Social Behaviours: Interactions among Individuals Affect Direct Genetic Effects

Traditional quantitative genetics assumes that an individual''s phenotype is determined by both genetic and environmental factors. For many animals, part of the environment is social and provided by parents and other interacting partners. When expression of genes in social partners affects trait expression in a focal individual, indirect genetic effects occur. In this study, we explore the effects of indirect genetic effects on the magnitude and range of phenotypic values in a focal individual in a multi-member model analyzing three possible classes of interactions between individuals. We show that social interactions may not only cause indirect genetic effects but can also modify direct genetic effects. Furthermore, we demonstrate that both direct and indirect genetic effects substantially alter the range of phenotypic values, particularly when a focal trait can influence its own expression via interactions with traits in other individuals. We derive a function predicting the relative importance of direct versus indirect genetic effects. Our model reveals that both direct and indirect genetic effects can depend to a large extent on both group size and interaction strength, altering group mean phenotype and variance. This may lead to scenarios where between group variation is much higher than within group variation despite similar underlying genetic properties, potentially affecting the level of selection. Our analysis highlights key properties of indirect genetic effects with important consequences for trait evolution, the level of selection and potentially speciation.     

Response to Selection Under Non-Random Mating II. Prediction

A general expression for response to selection appropriate for both random and non-random mating situations is derived and illustrated with full-sibbing.     

Multilevel selection and neighbourhood effects from individual to metapopulation in a wild passerine

Multilevel selection has rarely been studied in the ecological context of animal populations, in which neighbourhood effects range from competition among territorial neighbours to source-sink effects among local populations. By studying a Dupont's lark Chersophilus duponti metapopulation, we analyze neighbourhood effects mediated by song repertoires on fitness components at the individual level (life-span) and population level (growth rate). As a sexual/aggressive signal with strong effects on fitness, birdsong creates an opportunity for group selection via neighbour interactions, but may also have population-wide effects by conveying information on habitat suitability to dispersing individuals. Within populations, we found a disruptive pattern of selection at the individual level and an opposite, stabilizing pattern at the group level. Males singing the most complex songs had the longest life-span, but individuals with the poorest repertoires lived longer than 'average' males, a finding that likely reflects two male strategies with respect to fitness and sexual trait expression. Individuals from groups with intermediate repertoires had the longest life-span, likely benefitting from conspecific signalling to attract females up to the detrimental spread of competitive interactions in groups with superior vocal skills. Within the metapopulation selection was directional but again followed opposite patterns at the two levels: Populations had the highest growth rate when inhabiting local patches with complex repertoires surrounded by patches with simple repertoires. Here the song may impact metapopulation dynamics by guiding prospecting individuals towards populations advertising habitat quality. Two fitness components linked to viability were therefore influenced by the properties of the group, and birdsong was the target of selection, contributing to linking social/sexual processes at the local scale with regional population dynamics.     

Effects of random and non-random errors on phenotypic selection in autotetraploids

Summary A theoretical investigation was made to ascertain the effects of random and non-random deviations, called errors, of phenotypic from genotypic values on population means and on the response to phenotypic recurrent selection. The study was motivated as a selection experiment for disease resistance where there was either variability in the inoculation or environment (the random errors) or where the inoculation was above or below the the optimum rate where genetic differences in resistance are maximized (the non-random errors). The study was limited to the genetics at a diallelic locus (alleles B and b) in an autotetraploid population in random mating equilibrium. The response to selection was measured as the covariance of selection and compared to the exact covariance which was the covariance of selection without errors in phenotype. The random errors were modeled by assuming that a given percentage () of the population was uniformly distributed among the five possible genotype classes independent of their true genotypes. This model was analyzed numerically for a theoretical population with the frequency of the B allele (p) ranging from 0.0 to 1.0 and assumed errors of=0.1 and 0.5 for the following six types of genic action of the B allele: additive, monoplex dominance, partial monoplex dominance, duplex dominance, partial duplex dominance, and recessive. The effect of random error was to consistently reduce the response to selection by a percentage independent of the type of genic action at the locus. The effect on the population mean was an upward bias when p was low and a downward bias when p approached unity. In the non-random error model below optimum inoculations altered the phenotypes by systematically including percentage of susceptible genotypes into one or more other genotype classes with more genetic resistance (a positive shift). With above optimum inoculations, some resistant genotypes are classed with the non-resistant genotypes (a negative shift). The effects on the covariance of selection were found by numerical analysis for the same types of genic action and's as investigated for random error. With a negative shift and a low p, the covariance of selection was always reduced, but for an increasing p the covariance approached and exceeded the exact covariance for all types of genic action except additive. With a positive shift and a low p, response to selection was greatly improved for three types of genic action: duplex dominance, partial duplex dominance, and recessive. The effect of a non-random error on population means was to greatly bias the means upwards for a low p and positive shift, but with increasing p the bias decreased. A relatively slight decrease in the mean occurred with a negative shift. This study indicated check varieties commonly used to monitor selection pressures in screening programs are very responsive to positive non-random shifts, but are relatively unresponsive to negative shifts. The interaction of selection pressure, types of genic action, and genotypes in the class shift models was suggested as a partial explanation for the lack of response to increasing selection pressures observed in some breeding programs.Cooperative investigations of the Alfalfa Production Research Unit, United States Department of Agriculture, Agricultural Research Service, and the Nevada Agricultural Experiment Station, Reno, Nevada. Paper No. 404 Scientific Journal Series. Nevada Agricultural Experiment Station     

Multivariate models for human genetic analysis: aggregation, coaggregation, and tracking of systolic blood pressure and weight.

A multivariate path model parameterizing the sources of familial aggregation and coaggregation of systolic blood pressure and weight, as well as their tracking across time, is applied to longitudinal data collected in Muscatine, Iowa. Genetic, common household, and individual environmental effects, pleiotropy, and a direct regression effect of blood pressure on weight are parameterized. The sample consisted of 998 individuals distributed in 261 families of whom 601 were measured on four successive occasions. The data were divided with times 1 and 2 forming group 1, and times 3 and 4, group 2. Model fitting and estimation was performed using group 1, followed by testing the model and estimates using the data in group 2. Heritability estimates for systolic blood pressure and weight were .15 and .54, respectively. The genetic correlation between these traits was nonsignificant, but there was a significant direct regression effect. The results indicate that 30% of the full-sib correlation for systolic blood pressure is attributable to the aggregation of weight. In terms of tracking, 59% and 60% of the predicted systolic blood pressure and weight correlations, respectively, were attributable to genetic effects. Testing the model from group 1 in group 2 indicates that the qualitative relationships between blood pressure and weight are stable with time.     

Kin groups and trait groups: population structure and epidemic disease selection

A Monte Carlo simulation based on the population structure of a small-scale human population, the Semai Senoi of Malaysia, has been developed to study the combined effects of group, kin, and individual selection. The population structure resembles D.S. Wilson's structured deme model in that local breeding populations (Semai settlements) are subdivided into trait groups (hamlets) that may be kin-structured and are not themselves demes. Additionally, settlement breeding populations are connected by two-dimensional stepping-stone migration approaching 30% per generation. Group and kin-structured group selection occur among hamlets the survivors of which then disperse to breed within the settlement population. Genetic drift is modeled by the process of hamlet formation; individual selection as a deterministic process, and stepping-stone migration as either random or kin-structured migrant groups. The mechanism for group selection is epidemics of infectious disease that can wipe out small hamlets particularly if most adults become sick and social life collapses. Genetic resistance to a disease is an individual attribute; however, hamlet groups with several resistant adults are less likely to disintegrate and experience high social mortality. A specific human gene, hemoglobin E, which confers resistance to malaria, is studied as an example of the process. The results of the simulations show that high genetic variance among hamlet groups may be generated by moderate degrees of kin-structuring. This strong microdifferentiation provides the potential for group selection. The effect of group selection in this case is rapid increase in gene frequencies among the total set of populations. In fact, group selection in concert with individual selection produced a faster rate of gene frequency increase among a set of 25 populations than the rate within a single unstructured population subject to deterministic individual selection. Such rapid evolution with plausible rates of extinction, individual selection, and migration and a population structure realistic in its general form, has implications for specific human polymorphisms such as hemoglobin variants and for the more general problem of the tempo of evolution as well.     

Phenotypic evolution under gene-culture transmission in structured populations.

I consider a simple model for the evolution of a quantitative character is structured populations when an offspring's phenotype is determined partly by his or her genetic constitution and partly by cultural transmission of the parental phenotype. Analysis of the model indicates that when individual and group selection are in the same direction, phenotypic evolution always proceeds faster under gene-culture vs. purely genetic transmission. When individual and group selection are countervailing, altruistic characters evolve faster under gene-culture transmission when individual selection is weak and migration among groups is limited, with increased individual selection and migration tending to decrease the advantage of gene-culture transmission over purely genetic transmission. Given the prevalence of cultural transmission in higher species, these results suggest that contrary to what is often assumed, group selection may indeed by a potent evolutionary force in the evolution of altruistic characters.     

Effective size of populations under partial inbreeding and selection on a set of linked additive genes

The prediction theory of effective population size (Ne) is extended to cover selection on a set of linked additive genes and partial inbreeding (partial selfing or partial full-sib mating). Ne under selection is generally expressed as a function of the cumulative change in frequency of a neutral gene due to the random association between the neutral and selected genes generated by finite sampling. In this study, the association under partial selfing was classified into two types, the association between the neutral and selected genes on the same gamete, and the association between the neutral and selected genes each on the different gametes in the same parent. For partial full-sib mating, an additional association, i.e., the association between the neutral and selected genes each in the different parents in the same family, was included in the model. According to this classification of the association, the coefficient accounting for the cumulative change in frequency of the neutral gene was partitioned into two or three components. A method for computing the partitioned coefficients was obtained from the transition matrix approach, in which the joint effect of linkage, selection and partial inbreeding was taken into account. To assess the joint effects of linkage, selection and partial inbreeding on Ne, numerical computations with the obtained expressions were carried out. The effect of linkage on Ne was generally small, except for an extremely small genome size, while the partial inbreeding resulted in a drastic reduction in Ne. For a given genome size, Ne was essentially independent of the length and number of chromosomes. Some of these results were verified by stochastic simulations.