Showing papers in "Genetica in 1993"

Fluctuating asymmetry and sexual selection

Abstract: Fluctuating asymmetry occurs when an individual is unable to undergo identical development on both sides of a bilaterally symmetrical trait. Fluctuating asymmetry measures the sensitivity of development to a wide array of genetic and environmental stresses. We propose that fluctuating asymmetry is used in many signalling contexts for assessment of an individual’s ability to cope with its environment. We hypothesize that fluctuating asymmetry is used in sexual selection, both in fighting and mate choice, and in competition for access to resources. Evidence is reviewed showing that the patterns of fluctuating asymmetry in secondary sexual characters differ from those seen in other morphological traits. Secondary sexual characters show much higher levels of fluctuating asymmetry. Also, there is often a negative relationship between fluctuating asymmetry and the absolute size of ornaments, whereas the relationship is typically U-shaped in other morphological traits. The common negative relationship between fluctuating asymmetry and ornament size suggests that many ornaments reliably reflect individual quality.

Antisymmetry, directional asymmetry, and dynamic morphogenesis

Abstract: Fluctuating asymmetry is the most commonly used measure of developmental instability. Some authors have claimed that antisymmetry and directional asymmetry may have a significant genetic basis, thereby rendering these forms of asymmetry useless for studies of developmental instability. Using a modified Rashevsky-Turing reaction-diffusion model of morphogenesis, we show that both antisymmetry and directional asymmetry can arise from symmetry-breaking phase transitions. Concentrations of morphogen on right and left sides can be induced to undergo transitions from phase-locked periodicity, to phase-lagged periodicity, to chaos, by simply changing the levels of feedback and inhibition in the model. The chaotic attractor has two basins of attraction-right sidedominance and left side dominance. With minor disturbance, a developmental trajectory settles into one basin or the other. With increasing disturbance, the trajectory can jump from basin to basin. The changes that lead to phase transitions and chaos are thoseexpected to occur with either genetic change or stress. If we assume that the morphogen influences the behavior of cell populations, then a transition from phase-locked periodicity to chaos in the morphogen produces a corresponding transition from fluctuating asymmetry to antisymmetry in both morphogen concentrations and cell populations. Directional asymmetry is easily modeled by introducing a bias in the conditions of the simulation. We discuss the implications of this model for researchers using fluctuating asymmetry as an indicator of stress.

Integrons: Novel DNA elements which capture genes by site-specific recombination

Abstract: Integrons are unusual DNA elements which include a gene encoding a site-specific DNA recombinase, a DNA integrase, and an adjacent site at which a wide variety of antibiotic resistance and other genes are found as inserts One or more genes can be found in the insert region, but each gene is part of an independent gene cassette The inserted genes are expressed from a promoter in the conserved sequences located 5' to the genes, and integrons are thus natural expression vectors A model for gene insertion in which circular gene cassettes are inserted individually via a single site-specific recombination event has been proposed and verified experimentally The gene cassettes include a gene coding region and, at the 3' end of the gene an imperfect inverted repeat, a 59-base element The 59-base elements are a diverse family of elements which function as sites recognized by the DNA integrase Site-specific insertion of individual genes thus represents a further mechanism which contributes to the evolution of the genomes of Gram-negative bacteria and their plasmids and transposons Members of the most studied class of integrons, which include the sulI gene in the conserved sequences, are believed to be mobile DNA elements on the basis that they are found in many independent locations, and a discrete boundary is found at the outer end of the 5'-conserved segment However, the length of the 3'-conserved segment is variable in the integrons examined to date, and it is likely that this variability has arisen as the result of insertion and deletion events Though the true extent of the 3'-conserved segment remains to be determined, it seems likely that these integrons are mobile DNA elements The second known class of integrons comprises members of the Tn7 transposon family

Developmental stability in plants: symmetries, stress and epigenesis

Abstract: Plant developmental stability has received little attention in the past three or four decades. Here we review differences in plant and animal development, and discuss the advantages of using plants as experimental subjects in exploring developmental stability. We argue that any type of developmental invariant may be used to assess developmental stability and review the use of fluctuating asymmetry in studies of plant developmental stability. We also examine the use of deviations from translatory, radial, and self-symmetry as measures of developmental instability. The role of nonlinear dynamics and epigenesis in the production of the phenotype is also discussed.

The genetic basis of developmental stability. I: Relationships between stability, heterozygosity and genomic coadaptation

Abstract: The genetic basis of developmental stability has been much debated over the last four decades. Two major hypotheses have been developed over this period, one that argues that the level of stability is a reflection of the underlying level of genomic heterozygosity and the other that stability reflects the general level of genomic coadaptation, or genic balance. In this paper I have attempted to critically review the evidence for these hypotheses from both practical and theoretical perspectives. On balance there is little convincing evidence to suggest that heterozygosity plays a significant role in the determination and maintenance of developmental stability, whereas there is considerable support for the genomic coadaptation hypothesis.

Enzyme heterozygosity, metabolism, and developmental stability

Abstract: Developmental homeostasis, measured as either fluctuating asymmetry or variance of morphological characters, increases with enzyme heterozygosity in many, but not all, natural populations. These results have been reported forDrosophila, monarch butterflies, honeybees, blue mussels, side-blotched lizards, killifish, salmonid fishes, guppies, Sonoran topminnows, herring, rufous-collared sparrows, house sparrows, brown hares, white-tailed deer, and humans. Because heterozygosity at a few loci can not predict heterozygosity of the entiry genome, these loci must be detecting localized zones that influence the developmental environment.

Morphological integration of fluctuating asymmetry in the mouse mandible

Abstract: Morphological integration of fluctuating asymmetry (FA) was assessed for nine mandibular characters in random-bred house mice to test the hypothesis that there should be a significant integration or concordance of FAs at the population level, but not at the individual level. FA estimates for each of the nine characters were made for each of 16 subpopulations (two replicates each for mice varying in sex and age) and their correlations indicated a moderate level of integration (index of integration = 0.42). A matrix permutation test of the differences of the correlations within two (‘incisor’ and ‘muscle’) developmentally different character groups versus correlations between groups was significant, indicating the presence of morphological integration.Kendall’s coefficient of concordance also indicated a significant population asymmetry parameter for FAs within the two character groups. Correlations among individual FA estimates were generally lower than those calculated from subpopulations, the index of integration being 0.21. The matrix permutation test failed to show significant morphological integration among individual FAs, but Kendall’s coefficient of concordance was significant, indicating the presence of an ’individual asymmetry parameter’ among the nine characters. Principal components analysis and canonical correlation analysis confirmed the overall higher level of integration of FAs among the subpopulations and within the two character groupings.

Developmental origins of variation in human hand preference

Abstract: Though right-handedness is a prominant characteristic within all human societies, a substantial and stable proportion of individuals are left-handed. Any comprehensive approach to the origin of variation in handedness must account for substantial evidence that left-handedness is associated with reduced fitness, neurodevelopmental disorders, and reduced neuroanatomical asymmetry. In this paper we investigate the hypothesis that developmental instability in early fetal development underlies variation in handedness. In two studies we note an increased incidence of minor physical anomalies (MPAs) and fluctuating asymmetries in both left-handers and extreme right-handers. Moreover, extreme right-handers were more apt to have left-handed parents than moderate right-handers. These data suggest that deviation from moderate right-handedness reflect imprecise expression of a near-universal design due to developmental instability. Preliminary attempts to elucidate the mechanisms underlying developmental instability suggest that both polygenic homozygosity and particular HLA alleles may be important factors. These observations are discussed with respect to current genetic theories of handedness and human evolution.

Parasites increase fluctuating asymmetry of male Drosophila nigrospiracula: implications for sexual selection

Abstract: Fluctuating asymmetry (minor deviations from perfect bilateral symmetry) is manifested by individuals less able to buffer environmental stress during development. I utilized a system of two naturally-occurring parasites of Drosophila nigrospiracula to test whether parasitic infection during host development yields elevated degrees of fluctuating asymmetry in two morphological traits of males. This hypothesis has important implications for sexual selection, as it may explain why asymmetric males are often found to be sexually disadvantaged. In my system, nematodes infect larvae and therefore are more likely to disrupt development than mites which only parasitize adult flies. As predicted, nematode-infected male D. nigrospiracula had a higher degree of bristle asymmetry than did mite-infested and control (carrying neither parasite) males. There was also a significant relation between nematode number and degree of asymmetry. There was a significant negative relation between nematode load and size of adult males, implicating a causal link between nutritional stress during host development and fluctuating asymmetry. Patterns of wing length asymmetry were inconsistent with those of bristle asymmetry. Nematode-infected males did not differ in wing length asymmetry relative to mite-infested and control males, nor was there a significant relation between nematode number and wing asymmetry. This inconsistency in expression of asymmetry may reflect different intensities of selection operating on each morphological trait.

Evolutionary genetics of Drosophila esterases

Abstract: Over 30 carboxylester hydrolases have been identified in D. melanogaster. Most are classified as acetyl, carboxyl or cholinesterases. Sequence similarities among most of the carboxyl and all the cholinesterases so far characterised from D. melanogaster and other eukaryotes justify recognition of a carboxyl/cholinesterase multigene family. This family shows minimal sequence similarities with other esterases but crystallographic data for a few non-drosophilid enzymes show that the family shares a distinctive overall structure with some other carboxyl and aryl esterases, so they are all put in one superfamily of/beta hydrolases. Fifteen esterase genes have been mapped in D. melanogaster and twelve are clustered at two chromosomal sites. The constitution of each cluster varies across Drosophila species but two carboxyl esterases in one cluster are sufficiently conserved that their homologues can be identified among enzymes conferring insecticide resistance in other Diptera. Sequence differences between two other esterases, the EST6 carboxyl esterase and acetylcholinesterase, have been interpreted against the consensus super-secondary structure for the carboxyl/cholinesterase multigene family; their sequence differences are widely dispersed across the structure and include substantial divergence in substrate binding sites and the active site gorge. This also applies when EST6 is compared across species where differences in its expression indicate a difference in function. However, comparisons within and among species where EST6 expression is conserved show that many aspects of the predicted super-secondary structure are tightly conserved. Two notable exceptions are a pair of polymorphisms in the substrate binding site of the enzyme in D. melanogaster. These polymorphisms are associated with differences in substrate interactions in vitro and demographic data indicate that the alternative forms are not selectively equivalent in vivo.

The genetics of aging in the yeast Saccharomyces cerevisiae

Abstract: The yeast Saccharomyces cerevisiae possesses a finite life span similar in many attributes and implications to that of higher eukaryotes. Here, the measure of the life span is the number of generations or divisions the yeast cell has undergone. The yeast cell is the organism, simplifying many aspects of aging research. Most importantly, the genetics of yeast is highly-developed and readily applicable to the dissection of longevity. Two candidate longevity genes have already been identified and are being characterized. Others will follow through the utilization of both the primary phenotype and the secondary phenotypes associated with aging in yeast. An ontogenetic theory of longevity that follows from the evolutionary biology of aging is put forward in this article. This theory has at its foundation the asymmetric reproduction of cells and organisms, and it makes specific predictions regarding the genetics, molecular mechanisms, and phenotypic features of longevity and senescence, including these: GTP-binding proteins will frequently be involved in determining longevity, asymmetric cell division will be often encountered during embryogenesis while binary fission will be more characteristic of somatic cell division, tumor cells of somatic origin will not be totipotent, and organisms that reproduce symmetrically will not have intrinsic limits to their longevity.

Hybrid lethal systems in the Drosophila melanogaster species complex

Abstract: Lethal phases of the hybrids betweenDrosophila melanogaster and its sibling species,D. simulans are classified into three types: (1) embryonic lethality in hybrids carryingD. simulans cytoplasm andD. melanogaster X chromosome, (2) larval lethality in hybrids not carryingD. simulans X, and (3) temperature-sensitive pupal lethality in hybrids carryingD. simulans X. The same lethal phases are also observed when either of the two other sibling species,D. mauritiana orD. sechellia, is employed for hybridization withD. melanogaster. Here, we describe genetic analyses of each hybrid lethality, and demonstrate that these three types of lethality are independent phenomena. We then propose two models to interpret the mechanisms of each hybrid lethality. The first model is a modification of the conventional X/autosome imbalance hypothesis assuming a lethal gene and a suppressor gene are involved in the larval lethality, while the second model is for embryonic lethality assuming an interaction between a maternal-effect lethal gene and a suppressor gene.

Nonlinear growth dynamics and the origin of fluctuating asymmetry

Abstract: The nonlinear, complex nature of biosynthesis magnifies the impacts of small, random perturbations on organism growth, leading to distortions in adaptive allometries and, in particular, to fluctuating asymmetry. These distortions can be partly checked by cell-cell and inter-body part feedback during growth and development, though the latter mechanism also may lead to complex patterns in right-left asymmetry. Stress can be expected to increase the degree to which random growth perturbations are magnified and may also result in disruption of the check mechanisms, thus exaggerating fluctuating asymmetry.

Genes conferring late flowering inArabidopsis thaliana

Abstract: Three naturally occurring late flowering, vernalization responsive ecotypes ofArabidopsis thaliana, Pitztal, Innsbruck and Kiruna-2, were each crossed with the early flowering ecotypes of Landsbergerecta, Columbia and Niederzenz. Analysis of the subsequent generations suggested that late flowering in Kiruna-2 is recessive and mainly determined by a single, late flowering gene. This late flowering gene is not, however, the same as that in any of the late flowering mutants generated in the Landsbergerecta background. Both Pitztal and Innsbruck appear to contain the same dominant gene which confers late flowering to these ecotypes. The early flowering parents Niederzenz and Landsberg both contain genes which modify the phenotype of this dominant late flowering locus, causing F1 plants to flower either earlier (Landsberg) or later (Niederzenz) than the late parent. Mapping of the dominant late flowering locus from Pitztal demonstrated that late flowering co-segregated with an RFLP marker from one end of chromosome 4. This is a similar position to that ofFLA, the gene responsible for late flowering of theArabidopsis ecotypes Sf-2 and Le-O.

Evolutionary mechanisms of senescence

Abstract: This paper reviews theories of the evolution of senescence. The population genetic basis for the decline with age in sensitivity of fitness to changes in survival and fecundity is discussed. It is shown that this creates a presure of selection that disproportionately favors performance early in life. The extent of this bias is greater when there is a high level of extrinsic mortality; this accounts for much the diversity in life-history patterns among different taxa. The implications of quantitative genetic theory for experimental tests of alternative population genetic models of senescence are discussed. In particular, the negative genetic correlations between traits predicted by the antagonistic pleiotropy model may be obscured by positive correlations that are inevitable in a multivariate system, or by the effects of variation due to deleterious mutations. The status of the genetic evidence relevant to these theories is discussed.

Modification of developmental instability and fitness: Malathion-resistance in the Australian sheep blowfly,Lucilia cuprina

Abstract: The evolution of resistance to malathion byLucilia cuprina initially results in an increase in fluctuating asymmetry. Resistant flies are at a selective disadvantage, relative to susceptibles, in the absence of the insecticide. A fitness/asymmetry modifier of diazinon-resistant phenotypes ameliorates these effects resulting in malathion-resistant phenotypes of relative fitness and asymmetry similar to susceptibles. For the nine genotypic combinations of the modifier and malathion-resistance alleles, developmental time increases linearly with increasing asymmetry. Percentage egg hatch decreases linearly with increasing asymmetry. The initially disruptive effect of the malathion-resistant allele was partially dominant, the effect of the modifier dominant. The results are discussed in terms of developmental perturbation, asymmetry estimation and relative fitness to consider whether it is adequate to use changes in fluctuating asymmetry alone as measures of developmental instability. It is suggested that in some circumstances antisymmetry may indicate developmental instability and that the diazinon/malathion-resistance systems inL. cuprina may allow the relative importance of genetical and/or environmental developmental perturbations to be ascertained.

Genetics of food preference in Drosophila sechellia. I. Responses to food attractants

Abstract: To reveal the genetic mechanism of host selection in a monophagous fruit fly Drosophila sechellia, olfactory responses and oviposition preferences of this species were compared with those of closely related polyphagous species, D. simulans and D. melanogaster. Adult flies of D. sechellia were strongly attracted to the ripe fruit of Morinda citrifolia which is known to be the sole breeding site of this species. They were also attracted to the odor of n-caproic acid which is contained in the ripe fruit of M. citrifolia and is presumably responsible for the characteristic odor of the fruit. In contrast, D. simulans and D. melanogaster showed a strong repulsion to n-caproic acid. In parallel with the olfactory responses, D. sechellia females laid eggs preferentially on a medium containing n-caproic acid, to which the other two species showed an aversion. Genetic analyses using the hybrid progeny between D. sechellia and D. simulans suggested that the species differences in these behaviors are controlled by gene(s) located on the second chromosome.

Comparing mutants, selective breeding, and transgenics in the dissection of aging processes of Caenorhabditis elegans

Abstract: The genetic analysis of aging processes has matured in the last ten years with reports that long-lived strains of both fruit flies and nematodes have been developed. Several attempts to identify mutants in the fruit fly with increased longevity have failed and the reasons for these failures are analyzed. A major problem in obligate sexual species, such as the fruit fly, is the presence of inbreeding depression that makes the analysis of life-history traits in homozygotes very difficult. Nevertheless, several successful genetic analyses of aging in Drosophila suggest that with careful design, fruitful analysis of induced mutants affecting life span is possible. In the nematode Caenorhabditis elegans, mutations in the age-1 gene result in a life extension of some 70%; thus age-1 clearly specifies a process involved in organismic senescence. This gene maps to chromosome II, well separated from a locus (fer-15) which is responsible for a large fertility deficit in the original stocks. There is no trade-off between either rate of development or fertility versus life span associated with the age-1 mutation. Transgenic analyses confirm that the fertility deficit can be corrected by a wild-type fer-15 transformant (transgene); however, the life span of these transformed stocks is affected by the transgenic array in an unpredictable fashion. The molecular nature of the age-1 gene remains unknown and we continue in our efforts to clone the gene.

Population density effects on longevity.

Abstract: Population density, or the number of adults in an environment relative to the limiting resources, may have important long and short term consequences for the longevity of organisms. In this paper we summarize the way in which crowding may have an immediate impact on longevity, either through the phenomenon known as dietary restriction or through alterations in the quality of the environment brought on by the presence of large numbers of individuals. We also consider the possible long term consequences of population density on longevity by the process of natural selection. There has been much theoretical speculation about the possible impact of population density on the evolution of longevity but little experimental evidence has been gathered to test these ideas. We discuss some of the theory and empirical evidence that exists and show that population density is an important factor in determining both the immediate chances of survival and the course of natural selection.

The effect of superoxide dismutase alleles on aging in Drosophila

Abstract: The effects of superoxide dismutase on aging were tested using two differt experimental approaches. In the first, replicated populations with postponed aging were compared with their controls for frequencies of electrophoretic alleles at the SOD locus. Populations with postponed aging had consistently greater frequencies of the allele coding for more active SOD protein. This allele was not part of a segregating inversion polymorphism. The second experimental approach was the extraction of SOD alleles from different natural populations followed by the construction of different SOD genotypes on hybrid genetic backgrounds. This procedure did not uncover any statistical effect of SOD genotype on longevity or fecundity. There were large effects on longevity and fecundity due to the family from which a particular SOD genotype was derived. To detect the effects of SOD genotypes on longevity with high probability would require a ten-fold increase in the number of families used.

Bilateral variation and the evolutionary origin of macroscopic asymmetries

Abstract: Given that characters exhibiting macroscopic asymmetry have evolved in a wide variety of taxa, heritable variation for bilateral asymmetry must have arisen at some point in their history. The recognition that heritable variation may underlie some statistical asymmetries not only raises concerns about the incautious use of statistical estimates of FA in studies of developmental stability, but it suggests some intriguing questions about the possible evolutionary origins of macroscopic asymmetries. First, we developed an additive model of bilateral variation based on some simple assumptions about the developmental control of bilateral variation. Second, using a new approach for studying statistical asymmetries, we conducted an analysis of bilateral variation in eight metrical traits of a harpacticoid copepod (Tigriopus californicus) to search for novel forms of statistical asymmetries. The model we developed revealed three independent statistical asymmetries of potential evolutionary significance: a) a previously unrecognized form of asymmetry (referred to here as normal covariant asymmetry), b) antisymmetry, and c) directional asymmetry. Because each pattern of variation would seem to require different amounts and kinds of developmentalgenetic information [a- only negative feedback between sides (bilateral inhibition), b- both bilateral inhibition and average departure from symmetry (bilateral offset), c- bilateral inhibition, bilateral offset, and a consistent overdevelopment of one side or the other (side-bias control)], those requiring less information would seem more likely to represent earlier stages in the evolution of macroscopic asymmetries. Our analysis of bilateral variation in Tigriopus revealed no evidence for any form of statistical asymmetry other than fluctuating asymmetry. However, a significant positive covariation between sides, even after correction for body size variation, suggested that factors influencing relative limb length (whether genetic or environmental) affected both sides equally rather than one side at the expense of the other. Finally, we note that certain statistical asymmetries (directional asymmetry, any form of covariant asymmetry) may render characters unreliable for estimating developmental stability because, unlike pure fluctuating asymmetry, they may signal a genetic component to asymmetry variation.

Genetic engineering of grain and pasture legumes for improved nutritive value

Abstract: This review describes work aimed at the improvement of the nutritive value of grain and forage legumes using gene transfer techniques. Two traits which are amenable to manipulation by genetic engineering have been identified. These are plant protein quality and lignin content. In order to increase the quality of protein provided by the legume grains peas and lupins, we are attempting to introduce into these species chimeric genes encoding a sunflower seed protein rich in the sulphur-containing amino acids methionine and cysteine. These genes are designed to be expressed only in developing seeds of transgenic host plants. Chimeric genes incorporating a similar protein-coding region, but different transcriptional controls, are being introduced into the forage legumes lucerne and subterranean clover. In this case the genes are highly expressed in the leaves of transformed plants, and modifications have been made to the sunflower seed protein-coding sequences in order to increase the stability of the resultant protein in leaf tissue. Another approach to increasing plant nutritive value is represented by attempts to reduce the content of indigestible lignin in lucerne.

Inbreeding depression and male-mating behavior in Drosophila melanogaster.

Abstract: There have been relatively few studies designed to investigate the effects of inbreeding on behavioral traits. To study this phenomenon, five experimental lines ofDrosophila melanogaster made isogenic for chromosome 2 were evaluated for their male-mating ability and, subsequently, male courtship behavior. All lines showed significant reductions in overall mating ability, and males from all of these lines displayed impaired mating behavior, with two lines displaying particularly aberrant courtship patterns. Line 16 displayed an inability to successfully initiate copulation following successful courtship, while line 17 displayed significant reduction in locomotor activity, resulting in virtually no successful courtship or copulatory activity. The implications of these findings for competitive mating ability in wildDrosophila populations are presented. Further, the importance of mating success as a fitness component in the management of potentially highly inbred populations of endangered species is discussed.

Shaping intraspecifíc variation: development, ecology and the evolution of morphology and life history variation in tiger salamanders

Abstract: The tiger salamander,Ambystoma tigrinum, is a geographically widespread, morphologically variable, polytipic species. It is among the most variable species of salamanders in morphology and life history with two larval morphs (typical and cannibal) and three adult morphs (metamorphosed, typical branchiate, cannibal branchiate) that vary in frequency between subspecies and between populations within subspecies. We report morphometric evidence suggesting that branchiate cannibals arose through intraspecific change in the onset or timing of development resulting in the wider head and hypertrophied tooth-bearing skull bones characteristic of this phenotype. We also quantified bilateral symmetry of gill raker counts and abnormalities, then evaluated fluctuating asymmetry as a measure of the developmental stability of each morph. There was a significant interaction between fluctuating asymmetry of developmental abnormalities in cannibals and typicals and the locality where they were collected, suggesting that relative stability of each phenotype could vary among populations. While altered timing of developmental events appears to have a role in the evolution and maintenance of morphs, novel phenotypes persist only under favorable ecological conditions. Predictability of the aquatic habitat, genetic variation, kinship, body size, intraspecific competition and predation all affect expression and survival of the morphs inA. tigrinum. This taxon provides an excellent model for understanding the diversity and complexity of developmental and ecological variables controlling the evolution and maintenance of novel phenotypes.

Evolution of aging: Testing the theory using Drosophila

Abstract: Evolutionary explanations of aging (or senescence) fall into two classes. First, organisms might have evolved the optimal life history, in which survival and fertility late in life are sacrificed for the sake of early reproduction or high pre-adult survival. Second, the life history might be depressed below this optimal compromise by the influx of deleterious mutations; since selection against late-acting mutations is weaker, deleterious mutations will impose a greater load on late life. We discuss ways in which these theories might be investigated and distinguished, with reference to experimental work with Drosophila. While genetic correlations between life history traits determine the immediate response to selection, they are hard to measure, and may not reflect the fundamental constraints on life history. Long term selection experiments are more likely to be informative. The third approach of using experimental manipulations suffers from some of the same problems as measures of genetic correlations; however, these two approaches may be fruitful when used together. The experimental results so far suggest that aging in Drosophila has evolved in part as a consequence of selection for an optimal life history, and in part as a result of accumulation of predominantly late-acting deleterious mutations. Quantification of these effects presents a major challenge for the future.

Antagonistic pleiotropy, mutation accumulation, and human genetic disease

Abstract: The antagonistic pleiotropy theory of senescence is the most convincing theoretical explanation of the existence of aging. As yet, no locus or allele has been identified in a wild population with the features predicted by the pleiotropic theory. Human genetic diseases offer the opportunity to identify potentially pleiotropic alleles/loci. Four human genetic diseases—Huntington's disease, idiopathic hemochromatosis, myotonic dystrophy, and Alzheimer's disease—may exhibit pleiotropic effects and further study of these diseases might result in the identification of pleiotropic genes causing aging. Inability to find an early life selective benefit associated with these disease-causing alleles would favor the major alternative genetic explanation for aging, the mutation accumulation theory.

Persistence of subtle departures from symmetry over multiple molts in individual brachyuran crabs: relevance to developmental stability

Abstract: Fluctuating asymmetry (FA) is generally viewed as a population-level character. It is described by some measure of the variance of the difference between the right and left sides for a collection of individuals. Very little is known of the developmental origins of FA, despite the fact that FA is widely used to estimatedevelopmental stability. We present a novel technique for examining the growth trajectory of the asymmetries that give rise to FA, and we explore two sample data sets for the brachyuran crabHemigrapsus nudus. We have traced the fate of these small, random deviations from perfect symmetry through three successive molts of individual crabs. Invertebrates that molt, and hence grow in discrete steps, provide an easily preserved record of their growth. Model II regressions of measurements from one molt versus the previous molt can help describe the stability of subtle departures from symmetry over time. Although any number of different patterns may occur, we identify four general cases: a) asymmetries vary at random throughout growth (random determination), b) asymmetries remain unchanged in sign and magnitude (constant asymmetries), c) asymmetries increase in proportion to character size and hence increase with growth (size-dependent asymmetries), and d) asymmetries persist, but are reduced in magnitude (damped asymmetries). Data from tenHemigrapsus nudus, measured for between 21 and 28 metrical, limb-segment characters over three successive molts, yielded associations most similar to our pattern ‘b’, although some subtle departures in the direction of pattern ‘c’ were also observed. Persistent asymmetries accounted for 26% and 20% of the variance among asymmetries between molts 1 and 2, and molts 2 and 3 respectively. Thus, in spite of large and rapid increments in the external size of the crab, these subtle asymmetries tended to persist in both direction and magnitude, from molt to molt. This result suggests either i) that individual crabs have a genetic predisposition towards asymmetry in a particular direction but contribute to a continuous and normal distribution ofR-L differences at the population level, or ii) that these subtle asymmetries arose at some earlier ontogenetic stage and were preserved through growth. Either interpretation has important ramifications for the study of FA. The first suggests that under some circumstances FA may not provide a valid measure of developmental instability, because subtle departures from symmetry in an individual may have a genetic basis. The second implies that subtle departures from bilateral symmetry are not ‘corrected’ as an individual grows.

Ecology and genetics of insecticide resistance in Helicoverpa armigera: interactions between selection and gene flow

Abstract: Pyrethroid resistance inHelicoverpa armigera provides a model system in which to study evolution in natural populations. Resistance is seen to evolve as a consequence of selection pressure that varies within and between life-stages and gene flow. Although three different mechanisms are involved, present day fluctuations in phenotype frequency can be explained by variation in only one of these, metabolic resistance, that is inherited as a single, incompletely dominant gene,mfo. Selective mortality of phenotypes occurs in both larvae and adults in the presence of the pyrethroid insecticides. Although most individuals of all three genotypes are killed in young larvae, selection in this age-class contributes significantly to evolution of resistance. While there is some evidence of reduced fitness of resistant pupae during winter diapause, most of the decline in resistance frequencies each spring occurs as a result of immigration of susceptible individuals into insecticide-treated populations.

Evolutionary genetics of the Drosophila alcohol dehydrogenase gene-enzyme system

Abstract: Evolutionary genetics embodies a broad research area that ranges from the DNA level to studies of genetic aspects in populations. In all cases the purpose is to determine the impact of genetic variation on evolutionary change. The broad range of evolutionary genetics requires the involvement of a diverse group of researchers: molecular biologists, (population) geneticists, biochemists, physiologists, ecologists, ethologists and theorists, each of which has its own insights and interests. For example, biochemists are often not concerned with the physiological function of a protein (with respect to pH, substrates, temperature, etc.), while ecologists, in turn, are often not interested in the biochemical-physiological aspects underlying the traits they study. This review deals with several evolutionary aspects of the Drosophila alcohol dehydrogenase gene-enzyme system, and includes my own personal viewpoints. I have tried to condense and integrate the current knowledge in this field as it has developed since the comprehensive review by van Delden (1982). Details on specific issues may be gained from Sofer and Martin (1987), Sullivan, Atkinson and Starmer (1990); Chambers (1988, 1991); Geer, Miller and Heinstra (1991); and Winberg and McKinley-McKee (1992).

The origin and evolution of parthenogenesis in the Heteronotia binoei complex: synthesis

Abstract: The Heteronotia binoei complex includes several cryptic species of sexually reproducing lizards and parthenogenetic lineages derived from them. This paper synthesizes analyses of distribution and variation of allozymes, chromosomes, mitochondrial DNA and ribosomal DNA genes in order to make inferences about the origins of the parthenogenetic lineages, the extent and source of their genetic diversity, their current and historical biogeography and their ecological properties. The parthenogens appear to have arisen recently (relative to geographic differentiation within the sexual taxa) via episodes of repetitive hybridization between two of the sexual taxa. These events probably occurred within one or two small geographic areas of western Australia, after which some of the parthenogenetic lineages rapidly expanded their ranges to central Australia. The parthenogenetic form has extraordinarily high genetic diversity, mostly derived from the repetitive origins, but with some contribution from mutation and biased gene conversion/recombination being apparent. The rapid and extensive range expansion of the parthenogenetic lineages from western to central Australia attests to the short-term success of this reproductive strategy, in this case perhaps reinforced by the parthenogenetic females having higher fecundity than their smaller sexual relatives. However, the parthenogens are orders of magnitude more susceptible to infection by ectoparasitic mites, suggesting that they could be at a long-term disadvantage. The detailed characterization of this system provides a basis for critical evaluation of hypotheses about the evolutionary advantage of sexual reproduction derived from broad comparative studies.