Genetic heterogeneity and accelerated responses to directional selection in Drosophila

Summary Strains set up from single inseminated founder females of D. melanogaster from the same population have been previously shown to differ genetically for the incidence of flies with more than four scutellar chaetae (additional chaetae). Therefore the base population is polymorphic for this trait. This was exploited by carrying out directional selection in lines derived from those strains initially having a high mean chaeta number. This led to far more rapid responses to selection than were obtained in lines derived from strains with lower means, such that in one line a continuous accelerated response was observed for the first 12 generations of selection. A correlated response was found for sternopleural chaeta number at generation 19 of selection, showing that at least some genes may affect both traits.     

Polygenes and polymorphism

Summary Single inseminated founder females in D. melanogaster derived from the same population have led to genetically discrete strains for three quantitative traits, namely scutellar chaeta number, mating speed and duration of copulation. The base population from which the females were derived can therefore be argued to be polymorphic for these traits. Although polymorphisms for genes (or polygenes) for quantitative traits are expected to be ubiquitous in outbreeding species on theoretical grounds, this has not previously been generally demonstrated.     

Genetic heterogeneity among the founders of laboratory populations ofDrosophila. I. Scutlllar chaetae

The incidence of flies with mor than four scutellar chaetae (additional chaetae) has been followed for 45 generations in three strains set up from single inseminated females ofDrosophila melanogaster collected in the wild from the same locality at the same time. Each strain differed in the incidence of additional chaetae over this time, and the differences between strains were found to be controlled largely by additive genes.Fifteen further such strains were followed for 9 generations and each maintained consistent incidences of additional chaetae.Thus there may be genetic differences between populations derived from single inseminated females present in a given wild population, so lending support to a role for genetic drift (as a founder effect) in these situations. These differences may be maintained for many generations.Some possibilities for obtaining rapid responses in directional selection experiments based on the initial selection of favourable strains derived from single individuals are discussed briefly.     

Correlation between characters as related to developmental pattern in Drosophila

In order to better understand the genetic basis of some body traits and their correlations in Drosophila, in relation to their developmental history, a biometrical study was performed on three lines selected for short wing (fourth vein) and a control strain.The correlated response to selection for short wing and four body traits (thorax length and width, scutellar length, head width) and of eight other dimensional wing traits was considered.The results show a strong correlated response to selection of all wing traits, low correlations for the thorax characters, while head width remains relatively constant. Two groups of wing characters, corresponding to compartments of development, show different levels of covariation with the selected trait, the covariation being greater when the characters included in the same compartment of the selected trait are considered.The results are discussed in terms of developmental genetics of Drosophila and suggest that quantitative studies may be suitable for studying the rôle of interactions between sets of genes controlling development.     

A pair of closely linked genes controlling high scutellar chaeta number in Drosophila

Summary 1. A line selected for high scutellar chaeta number reached a mean of about 16 chaetae in females and 13.5 in males at the 69th generation of selection following an accelerated response to selection which commenced at generation 65 and added five chaetae. 2. The accelerated response can probably be explained in terms of two recessive high chaeta number genes 1.05 cM apart, and which are located between po and vg on chromosome II. The gene closest to vg was found to be scabrous, sca, which causes rough eyes when homozygous and has a pleiotropic effect on scutellar chaeta number. The gene was found in one of the strains used in setting up the selection lines. 3. The results are discussed in relation to other theories of control of the scutellar chaeta system.     

Egg lengths inDrosophila melanogaster and correlated responses to selection

Egg lengths were studied inDrosophila using three lines, namely a high line, where selection for increased egg length was carried out, a low line where selection for decreased egg length was carried out, and a control line. Selection was more effective in the low than the high line. Adult fly weight assayed in the 18th and 19th generations of selection responded similarly to egg length. Sternopleural chaetae number, however, responded more in the high than in the low line. In the high line the response occurred earlier than for egg length. This rather surprising result can probably be explained by assuming that the genetic architectures controlling chaetae number and egg length differ.     

Selection for six scutellar chaetae in Drosophila melanogaster

Summary 1. From a base population showing some flies with more than the normal 4 scutellar chaeta phenotype directional selection was carried out and lead to a line with many flies having 6 chaetae. 2. Selection was then practised for 6 chaetae such that the extra 2 chaetae were in the anterior left and anterior right positions on the scutellum. This lead to a line with most flies having this chaeta number and pattern, therefore showing some canalization. 3. Additive genetic activity controlling the increased chaeta number was found on all the 3 major chromosomes.     

Apparent directional selection by biased pleiotropic mutation

Pleiotropic effects of deleterious mutations are considered to be among the factors responsible for genetic constraints on evolution by long-term directional selection acting on a quantitative trait. If pleiotropic phenotypic effects are biased in a particular direction, mutations generate apparent directional selection, which refers to the covariance between fitness and the trait owing to a linear association between the number of mutations possessed by individuals and the genotypic values of the trait. The present analysis has shown how the equilibrium mean value of the trait is determined by a balance between directional selection and biased pleiotropic mutations. Assuming that genes act additively both on the trait and on fitness, the total variance-standardized directional selection gradient was decomposed into apparent and true components. Experimental data on mutation bias from the bristle traits of Drosophila and life history traits of Daphnia suggest that apparent selection explains a small but significant fraction of directional selection pressure that is observed in nature; the data suggest that changes induced in a trait by biased pleiotropic mutation (i.e., by apparent directional selection) are easily compensated for by (true) directional selection.     

黄瓜雄花性状的遗传分析

为了解黄瓜雄花花器的遗传特性,该研究以雄花器官较小的华南型黄瓜二早子为母本,花器较大的加工型黄瓜NC-76为父本,构建4世代遗传群体,并采用多世代联合分离分析方法,分析黄瓜雄花花器性状的遗传特性。结果表明:分离群体的雄花花梗和花冠长2个性状均表现为单峰分布,表明两性状为数量性状且有主基因控制;花梗长性状符合2对完全显性主基因+加性-显性多基因(E-5)模型,花冠长性状符合2对加性-显性-上位性主基因+加性-显性-上位性多基因(E-1)模型;控制花梗长性状的两对主基因的加性效应相等,为0.573,多基因的加性效应和显性效应值相差不大,且均为负向;控制花冠长度性状的2对主基因的加性效应均为0,显性效应分别为-0.226和-0.472,在上位性作用中以加性×加性和显性×显性互作为主,多基因以显性效应为主,正向显性效应值为0.613,大于负向的加性效应值。花梗和花冠长度两个性状在F2群体中主基因遗传率分别为61.04%和69.60%,多基因遗传率均为0。由此看出黄瓜雄花花器性状为数量遗传,遗传率相对较高。该研究结果显示在黄瓜杂交育种中对花器大小选择可以在较早世代选择。     

Cell interactions in the generation of chaetae pattern inDrosophila

Summary We have already shown that theachaetae-scute complex (AS-C) ofDrosophila is regulated by two genes,hairy andextramacrochaetae. Using mutants in these genes, we have analysed how different levels of expression of AS-C affect the pattern of chaetae. The results indicate that the spatial distribution of chaetae results from cell interactions, probably by a mechanism of lateral inhibition. The results are discussed in view of the different theories of pattern formation.     

Genetic heterogeneity among the founders of laboratory populations of Drosophila IV. Scutellar chaetae in different environments

The incidence of flies with more than four seutellar chaetae (additional chaetae) has been followed for up to 3 years in 4 lines at 15°C, 20°C, 25°C and 27.5°C derived from each of 3 strains, initially set up from single inseminated females of Drosophila melanogaster, collected in the wild single the same locality at the same time. Within each set of temperature lines, the incidence of additional chaetae was highest in one strain, intermediate in another strain, and lowest in the third. This shows that the differences between lines can be at least partly attributed to differences between the founder females of the three strains, although some divergence occurred between lines within strains in some cases. With respect to quantitative traits, the results indicate that in situations where new habitats are occupied by a small number of colonists, the nature of the founder female is possibly of greater importance than the different environments of the new habitats.Some of this work was supported by the Australian Research Grants Committee.     

Effects of assay conditions in life history experiments with Drosophila melanogaster

Selection experiments with Drosophila have revealed constraints on the simultaneous evolution of life history traits. However, the responses to selection reported by different research groups have not been consistent. Two possible reasons for these inconsistencies are (i) that different groups used different environments for their experiments and (ii) that the selection environments were not identical to the assay environments in which the life history traits were measured. We tested for the effect of the assay environment in life history experiments by measuring a set of Drosophila selection lines in laboratories working on life history evolution with Drosophila in Basel, Groningen, Irvine and London. The lines measured came from selection experiments from each of these laboratories. In each assay environment, we measured fecundity, longevity, development time and body size. The results show that fecundity measurements were particularly sensitive to the assay environment. Differences between assay and selection environment in the same laboratory or differences between assay environments between laboratories could have contributed to the differences in the published results. The other traits measured were less sensitive to the assay environment. However, for all traits there were cases where the measurements in one laboratory suggested that selection had an effect on the trait, whereas in other laboratories no such conclusion would have been drawn. Moreover, we provide good evidence for local adaptation in early fecundity for lines from two laboratories.     

Explaining stasis: microevolutionary studies in natural populations

Microevolution, defined as a change in the genetic constitution of a population over time, is considered to be of commonplace occurrence in nature. Its ubiquity can be inferred from the observation that quantitative genetic divergence among populations usually exceeds that to be expected due to genetic drift alone, and from numerous observations and experiments consistent with local adaptation. Experimental manipulations in natural populations have provided evidence that rapid evolutionary responses may occur in the wild. However, there are remarkably few cases where direct observations of natural populations have revealed microevolutionary changes occurring, despite the frequent demonstration of additive genetic variation and strong directional selection for particular traits. Those few cases where responses congruent with expectation have been demonstrated are restricted to changes over one generation. In this article we focus on possible explanations as to why heritable traits under apparently strong directional selection often fail to show the expected evolutionary response. To date, few of these explanations for apparent stasis have been amenable to empirical testing. We describe new methods, derived from procedures developed by animal breeding scientists, which can be used to address these explanations, and illustrate the approach with examples from long-term studies of collared flycatchers (Ficedula albicollis) and red deer (Cervus elaphus). Understanding why most intensively studied natural populations do not appear to be evolving is an important challenge for evolutionary biology.     

Complex patterns of local adaptation in heat tolerance in Drosophila simulans from eastern Australia

Latitudinal clines are considered a powerful means of investigating evolutionary responses to climatic selection in nature. However, most clinal studies of climatic adaptation in Drosophila have involved species that contain cosmopolitan inversion polymorphisms that show clinal patterns themselves, making it difficult to determine whether the traits or inversions are under selection. Further, although climatic selection is unlikely to act on only one life stage in metamorphic organisms, a few studies have examined clinal patterns across life stages. Finally, clinal patterns of heat tolerance may also depend on the assay used. To unravel these potentially confounding effects on clinal patterns of thermal tolerance, we examined adult and larval heat tolerance traits in populations of Drosophila simulans from eastern Australia using static and dynamic (ramping 0.06 °C min^?1) assays. We also used microsatellites markers to clarify whether demographic factors or selection are responsible for population differentiation along clines. Significant cubic clinal patterns were observed for adult static basal, hardened and dynamic heat knockdown time and static basal heat survival in larvae. In contrast, static, hardened larval heat survival increased linearly with latitude whereas no clinal association was found for larval ramping survival. Significant associations between adult and larval traits and climatic variables, and low population differentiation at microsatellite loci, suggest a role for climatic selection, rather than demographic processes, in generating these clinal patterns. Our results suggest that adaptation to thermal stress may be species and life‐stage specific, complicating our efforts to understand the evolutionary responses to selection for increasing thermotolerance.     

Power of the joint segregation analysis method for testing mixed major-gene and polygene inheritance models of quantitative traits

Understanding the genetic architecture of quantitative traits can greatly assist the design of strategies for their manipulation in plant-breeding programs. For a number of traits, genetic variation can be the result of segregation of a few major genes and many polygenes (minor genes). The joint segregation analysis (JSA) is a maximum-likelihood approach for fitting segregation models through the simultaneous use of phenotypic information from multiple generations. Our objective in this paper was to use computer simulation to quantify the power of the JSA method for testing the mixed-inheritance model for quantitative traits when it was applied to the six basic generations: both parents (P₁ and P₂), F₁, F₂, and both backcross generations (B₁ and B₂) derived from crossing the F₁ to each parent. A total of 1968 genetic model-experiment scenarios were considered in the simulation study to quantify the power of the method. Factors that interacted to influence the power of the JSA method to correctly detect genetic models were: (1) whether there were one or two major genes in combination with polygenes, (2) the heritability of the major genes and polygenes, (3) the level of dispersion of the major genes and polygenes between the two parents, and (4) the number of individuals examined in each generation (population size). The greatest levels of power were observed for the genetic models defined with simple inheritance; e.g., the power was greater than 90% for the one major gene model, regardless of the population size and major-gene heritability. Lower levels of power were observed for the genetic models with complex inheritance (major genes and polygenes), low heritability, small population sizes and a large dispersion of favourable genes among the two parents; e.g., the power was less than 5% for the two major-gene model with a heritability value of 0.3 and population sizes of 100 individuals. The JSA methodology was then applied to a previously studied sorghum data-set to investigate the genetic control of the putative drought resistance-trait osmotic adjustment in three crosses. The previous study concluded that there were two major genes segregating for osmotic adjustment in the three crosses. Application of the JSA method resulted in a change in the proposed genetic model. The presence of the two major genes was confirmed with the addition of an unspecified number of polygenes. Received: 18 August 2000 / Accepted: 9 March 2001     

The Isolation of Polygenic Factors Controlling Bristle Score in Drosophila Melanogaster. II. Distribution of Third Chromosome Bristle Effects within Chromosome Sections

In the present study an attempt has been made to characterize the genetic ``factors' controlling quantitative characters, bristle numbers, in Drosophila melanogaster. A low sternopleural bristle multiple recessive marker third chromosome was used to analyze a high sternopleural third chromosome, in a high sternopleural bristle background. An attempt was made to estimate the minimum number of ``effective factors' involved in the difference in bristle score between the tested and marker chromosomes. Apart from sternopleural, scutellar and ocellar bristles, a new character, subprimal bristles, was also scored. The unselected characters were used to help in the factor locations, and an attempt made to detect epistasis. Concentrations of bristle effects were found, as were a few `factors' of large effect. At least 17 sternopleural bristle factors are required to account for the difference in bristle score between the high tested third chromosome and the low tester third chromosome. There was an ascertainment problem for polygenes with effects of less than about 0.6 phenotypic standard deviation. Only an estimate of the minimum number of factors and approximate locations can be given with any degree of certainty. The results are compatible with the hypothesis (among others) that quantitative characters are under the control of a few major genes supported by numerous genes with smaller effect.     

Use of recombinant inbred strains to map genes of aging

Recombinant inbred strains have been used in a number of organisms for segregation and linkage analysis of quantitative traits. One major advantage of the recombinant inbred (RI) methodology is that the genetic identity of individuals within a strain permits replicate measures of the same recombinant genotype. Such replicability is important for traits such as aging inDrosophila, where phenotypic expression is highly influenced by different environmental conditions. RI strain methodology has an added advantage for DNA marker-based linkage analysis of traits measured over the lifespan of the organism. The DNA can be extracted from individuals of the same genotype as those measured in a longevity study. In this paper an argument is presented for the use of a set of recombinant inbred strains to map the quantitative trait loci involved in the aging process inDrosophila. A unique use of a set of stable, transposable moleular markers to trace the quantitative trait loci involved is suggested.     

Genetic control of the opaque-2 gene and background polygenes over some kernel traits in maize (Zea mays L.)

Some kernel traits of agronomical importance in maize are affected by the opaque-2 (o2) gene and background polygenes, which express in different genetic systems such as embryo, endosperm, cytoplasm and maternal plant. A genetic model for seed quantitative traits with the o2 gene effects and polygenic effects as well as their GE interactions was used for protein content, lysine content, oil content and kernel density in maize. The results suggested that the o2 gene was involved in the traits investigated but the effects of the o2 gene were distinctive on various traits. The effects of the o2 gene were large on lysine content and protein content while minor on oil content. There was a substantially wide quantitative variation from polygenes expressing in different genetic systems for the traits evaluated. Significant GE interactions of the o2 gene and background polygenes declared that not only the main effects but also specific expressions depending on environments were responsible for variation of the traits studied. There seemed to have strong maternal heterosis and slight embryo heterosis for kernel density.     

P element transposon-induced quantitative genetic variation for inebriation time in Drosophila melanogaster

Summary Bi-directional selection was carried out in coisogenic stocks with and without mobilised P element transposons to determine whether P elements induce quantitative genetic variation for inebriation time in Drosophila. There was significant response to 11 generations of selection in both pairs of replicates of bi-directional selection from an isogenic base stock in which P elements had been mobilised. Conversely, there was no significant response to 11 generations of identical selection in the control lines derived from a relatively inbred line lacking P elements. Thus, P elements have induced quantitative genetic variation for inebriation time.     

MORPHOLOGICAL EVOLUTION OF THE DROSOPHILA VIRILIS SPECIES GROUP AS ASSESSED BY RATE TESTS FOR NATURAL SELECTION ON QUANTITATIVE CHARACTERS

Two rate tests for assessing natural selection on quantitative traits are discussed for their usefulness in macroevolutionary and adaptational studies. The underlying assumptions and parameter estimation for the constant-heritability (CH) and mutation-drift-equilibrium (MDE) models, which are the bases for these tests, are discussed. The purpose of these rate tests is to determine whether morphological change has occurred either too fast to be explained by neutral drift, which suggests directional selection, or too slow, which suggests stabilizing selection. Previous formulations of these rate tests have ignored the phylogenetic component. Several models of evolution are considered to help account for phylogeny in the context of rate tests. The MDE rate test for stabilizing selection was performed on nine morphological characters among several species of the Drosophila virilis species group. These tests can be interpreted to suggest that stabilizing selection has probably been a major factor in producing the observed similarity among the Drosophila species examined.