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Signatures of selection among sex-determining alleles of the honey bee.

Patterns of DNA polymorphisms are a primary tool for dissecting signatures of selection; however, the underlying selective forces are poorly understood for most genes. A classical example of diversifying selection is the complementary sex-determining locus that is found in the very large insect order Hymenoptera (bees, wasps, ants, and sawflies). The gene responsible for sex determination, the complementary sex determiner (csd), has been most recently identified in the honey bee. Females are heterozygous at this locus. Males result when there is only one functional allele present, as a result of either homozygosity (fertilized eggs) or, more commonly, hemizygosity (unfertilized eggs). The homozygotes, diploid males, do not reproduce and have zero fitness, which implies positive selection in favor of rare alleles. Large differences in csd cDNA sequences within and between four populations were found that fall into two major groups, types I and II. Type I consists of several allelic lineages that were maintained over an extended period, an indication of balancing selection. Diversifying selection has operated on several confined parts of the protein, as shown by an excess of nonsynonymous differences. Elevated sequence differences indicate another selected part near a repeat region. These findings have general implications about the understanding of both the function of the multiallelic mechanism and the adaptive processes on the level of nucleotide sequences. Moreover, the first csd sequence data are a notable basis for the avoidance of diploid males in bee selection programs by allele-assisted breeding.

Alleles↗

Direct benefits and the evolution of female-biased cooperative breeding in Seychelles warblers.

Inclusive fitness benefits have been suggested to be a major selective force behind the evolution of cooperative breeding. We investigated the fitness benefits selecting for cooperative breeding in the Seychelles warbler, Acrocephalus sechellensis. A microsatellite-based genotyping method was used to determine the relatedness of subordinates to group offspring in an isolated population of Seychelles warblers. The indirect and direct breeding benefits accruing to individual subordinates were then calculated for every successful breeding event over a three-year period. We show that female subordinates frequently gained parentage and that this, combined with high levels of extragroup paternity, resulted in low levels of relatedness between subordinates and nondescendent offspring within a territory. Direct breeding benefits were found to be significantly higher than indirect kin benefits for both female and male subordinates. As predicted, female subordinates gained significantly more direct breeding opportunities and therefore higher inclusive fitness benefits by being a subordinate within a group than did males. This may explain why most subordinates in the Seychelles warbler are female.

Animals↗

The spandrels of San Marco and the Panglossian paradigm: a critique of the adaptationist programme.

An adaptationist programme has dominated evolutionary thought in England and the United States during the past 40 years. It is based on faith in the power of natural selection as an optimizing agent. It proceeds by breaking an oragnism into unitary 'traits' and proposing an adaptive story for each considered separately. Trade-offs among competing selective demands exert the only brake upon perfection; non-optimality is thereby rendered as a result of adaptation as well. We criticize this approach and attempt to reassert a competing notion (long popular in continental Europe) that organisms must be analysed as integrated wholes, with Baupläne so constrained by phyletic heritage, pathways of development and general architecture that the constraints themselves become more interesting and more important in delimiting pathways of change than the selective force that may mediate change when it occurs. We fault the adaptationist programme for its failure to distinguish current utility from reasons for origin (male tyrannosaurs may have used their diminutive front legs to titillate female partners, but this will not explain why they got so small); for its unwillingness to consider alternatives to adaptive stories; for its reliance upon plausibility alone as a criterion for accepting speculative tales; and for its failure to consider adequately such competing themes as random fixation of alleles, production of non-adaptive structures by developmental correlation with selected features (allometry, pleiotropy, material compensation, mechanically forced correlation), the separability of adaptation and selection, multiple adaptive peaks, and current utility as an epiphenomenon of non-adaptive structures. We support Darwin's own pluralistic approach to identifying the agents of evolutionary change.

Adaptation, Biological↗

Physiology of peripheral neurons innervating otolith organs of the squirrel monkey. II. Directional selectivity and force-response relations.

1. The directional selectivity of peripheral otolith neurons was studied in the barbiturate-anesthetized squirrel monkey (Saimiri sciureus). Each unit's most sensitive axis was characterized by a functional polarization vector. The direction of a centrifugal force was varied with respect to the vector and to the macular plane. The neurons respond in an excitatory manner to shearing forces orthogonally disposed to the vector. The sensitivity to orthogonal shears was usually 10-15% of the sensitivity to parallel shearing forces. There was no significant response to orthogonal compressions, nor did compressions modify the response to shearing forces. 2. Force-response functions were obtained in the stimulus range of +/- 4.92 g. Forces were directed parallel to each unit's polarization vector. The functions are sigmoid shaped and possess both inhibitory and excitatory plateaus. The presumed physiological range of +/-1 g is represented in the lower (concave upward) portion of the function and has a dynamic range, expressed in terms of response magnitude, amounting to 20-40% of the potential dynamic range of the neuron. 3. There was considerable variation among units in their +/- 4.92 g force-response curves. The salient features of the functions are described by three factors, tentatively identified as a transduction gain, a receptor bias, and a mechanical gain. Both the resting discharge (do) and the +/-1 g sensitivity (so) vary in the same direction with changes in the factors related to transduction gain and receptor bias. It is shown that this covariation provides a quantitatively precise explanation for the positive relation between do and so.

Animals↗

Monitoring program of biological status of human populations related to environmental changes.

The paper aims at a synthesis of research carried out hitherto which might serve to evaluate the diagnostic value of human biological traits related to natural, economic and socio-cultural properties of the environment. The development of a monitoring program is the object of research of the Department of Human Ecology, the Polish Academy of Sciences, in the years 1986-1990 in Poland, and also of recommendations of the IUAES Human Ecology Commission for the last decade of the 20th century in various countries over the world. Generally, the stages of ontogenesis that are appropriate for monitoring are those which involve an intense proliferation of cells and a rapid growth. The appropriate subjects are the more ecosensitive individuals, that is the heterozygous ones as well as men. The more ecosensitive traits are those developed after birth in over 50 per cent of their adult value. Traits of the highest diagnostic (discriminant) value with respect to environmental conditions were selected. The results are based on the analysis of: 35 somatic traits in 1034 newborns from Białystok and Zambrów in relation to the traits of their families and parents; 21 somatic traits in 2461 of 11 year-old children from Lublin; 40 somatic, physiological and psychomotor traits in 1186 children, 6-19 years old from Poland; 26 morphological and physiological traits in 4771 subjects of 5-90 years of age in Poland; 9 somatic and 17 motor traits in 127,489 children, 7-19 years old, in relation to economic and demographic conditions in 98 regions of Poland; and 27 somatic, physiological and psychomotor traits in children and their parents from the ages of 3-80 years, from 3995 families from Poland. The age particularly important to the study includes: foetal growth effects recorded as newborn status, child development from about half a year (since weaning) to about 3 years of age, development at the early period of puberty, the period at adolescence, age at the peak value (maximal size, best results) of the main morphological and functional traits (usually between the age of 12 and 30), and finally, the rate of involution of these main traits in older age. For the "minimum program" (for public health and environmental conservation services) among somatic traits, 3 were selected for a minimum program: stature, Kaup index, and cephalic index. Among physiological traits, 2 were selected: forced respiratory volume per second and hemoglobin concentration. Among motor traits, 3 were selected for the minimum program: distant movement accuracy, grip strength, and standing long jump.(ABSTRACT TRUNCATED AT 400 WORDS)

Adolescent↗

The theory of neutral and weakly selected genes.

The controversy over whether a major fraction of evolutionary allele substitutions and protein polymorphisms in natural populations is due to random drift of neutral mutations remains unresolved; but the discussion has led to extensions of population genetic theory, particularly in stochastic models. The work of Maruyama in seeking quantities that are not strongly dependent on assumptions about population structure is particularly discussed. Under some circumstances, and perhaps quite generally, the probability of eventual fixation of a mutant gene, the total number of heterozygotes in which the gene is involved before its fixation or less, and the total number of heterozygotes during the time the gene has a specified frequency in the entire population have this desirable property. Whether there are many mutants that are so nearly neutral as to be unaffected by selective forces or not, it is clear that there are many loci at which selection is very weak and some of the evolutionary consequences of this are discussed briefly.

Amino Acid Sequence↗

Does a parasite have a better chance of survival if an Inuit or a Mayan spits on it?

An earlier puzzling observation [Shields and Mann, J Craiofac Genet Dev Biol, 16:126-136, 1996] that the prevalence of a polymorphic male predominate, major salivary gland-associated, static lesion of the mandible increased exponentially from the Arctic to the Tropics was explained by both positive and negative selection (balancing) on major salivary gland endocrine and exocrine factors. Additional prevalence rates presented here identified three high prevalence high-Temperate zone cultures that were unusually exposed to alimentary parasites. A correlation between macroparasite exposure and the mandibular lesion helped refine the potential selective forces that fashioned major salivary gland size variation. The data suggests that positive selection occurred for androgen-induced enlargement of the suite of major salivary glands and consequently increased quantities of factors. Increased quantities of salivary gland biologically active factors enhance innate protection against infestation of macroparasites per se, especially gut parasites. The data further suggests that negative selection against enlarged salivary glands occurred as protection against electrolyte imbalances in electrolyte stressed environments and in females.

Climate↗

Recent developments in U.S. Air Force pilot candidate selection and classification.

Recent U.S. Air Force policy decisions regarding Specialized Undergraduate Pilot Training (SUPT) have eliminated the need to classify pilot candidates into training specialties prior to entering primary jet training. Under the new system, specialized training assignments will occur at the completion of primary jet training and will be based on flying and academic performance, student preferences, and aircraft availability. Another significant change to the SUPT program was the development of a refined pilot candidate selection model that reduced potential threats to test compromise and gaming strategies.

Aptitude Tests↗

Measles: extinction or evolution?

The human body is the biological environment to which measles virus must adapt. In this paper the immunity conferred by vaccination is viewed as a selective force on the evolution of this virus. The possible consequences of this type of selection pressure include an increased Ro, antigenic variation, and immune depression.

Antigens, Viral↗

Some population genetic models combining artificial and natural selection pressures.

The evolutionary behavior of a diploid population characterized by a trait determined at one or two major loci subject to the combined effects of artificial and natural selection pressures is investigated. A number of different genotypic phenotypic associations are set forth, including additive allelic effects and additive loci effects with a variety of culling programs. Threshold selection schemes as well as culling favoring intermediate phenotypic values are considered. For these formulations results are reported concerning the dynamic progress of the population, a delineation of the numbers and properties of the stable equilibria outcomes, and a discussion of their qualitative and quantitative dependence on the two kinds of selection forces. The level of culling can be used as a control through which natural selection parameters can be estimated.

Alleles↗

Geographical distribution of selected and putatively neutral SNPs in Southeast Asian malaria parasites.

Loci targeted by directional selection are expected to show elevated geographical population structure relative to neutral loci, and a flurry of recent papers have used this rationale to search for genome regions involved in adaptation. Studies of functional mutations that are known to be under selection are particularly useful for assessing the utility of this approach. Antimalarial drug treatment regimes vary considerably between countries in Southeast Asia selecting for local adaptation at parasite loci underlying resistance. We compared the population structure revealed by 10 nonsynonymous mutations (nonsynonymous single-nucleotide polymorphisms [nsSNPs]) in four loci that are known to be involved in antimalarial drug resistance, with patterns revealed by 10 synonymous mutations (synonymous single-nucleotide polymorphisms [sSNPs]) in housekeeping genes or genes of unknown function in 755 Plasmodium falciparum infections collected from 13 populations in six Southeast Asian countries. Allele frequencies at known nsSNPs underlying resistance varied markedly between locations (F(ST) = 0.18-0.66), with the highest frequencies on the Thailand-Burma border and the lowest frequencies in neighboring Lao PDR. In contrast, we found weak but significant geographic structure (F(ST) = 0-0.14) for 8 of 10 sSNPs. Importantly, all 10 nsSNPs showed significantly higher F(ST) (P < 8 x 10(-5)) than simulated neutral expectations based on observed F(ST) values in the putatively neutral sSNPs. This result was unaffected by the methods used to estimate allele frequencies or the number of populations used in the simulations. Given that dense single-nucleotide polymorphism (SNP) maps and rapid SNP assay methods are now available for P. falciparum, comparing genetic differentiation across the genome may provide a valuable aid to identifying parasite loci underlying local adaptation to drug treatment regimes or other selective forces. However, the high proportion of polymorphic sites that appear to be under balancing selection (or linked to selected sites) in the P. falciparum genome violates the central assumption that selected sites are rare, which complicates identification of outlier loci, and suggests that caution is needed when using this approach.

Animals↗

Effects of natural and sexual selection on adaptive population divergence and premating isolation in a damselfly.

The relative strength of different types of directional selection has seldom been compared directly in natural populations. A recent meta-analysis of phenotypic selection studies in natural populations suggested that directional sexual selection may be stronger in magnitude than directional natural selection, although this pattern may have partly been confounded by the different time scales over which selection was estimated. Knowledge about the strength of different types of selection is of general interest for understanding how selective forces affect adaptive population divergence and how they may influence speciation. We studied divergent selection on morphology in parapatric, natural damselfly (Calopteryx splendens) populations. Sexual selection was stronger than natural selection measured on the same traits, irrespective of the time scale over which sexual selection was measured. Visualization of the fitness surfaces indicated that population divergence in overall morphology is more strongly influenced by divergent sexual selection rather than natural selection. Courtship success of experimental immigrant males was lower than that of resident males, indicating incipient sexual isolation between these populations. We conclude that current and strong sexual selection promotes adaptive population divergence in this species and that premating sexual isolation may have arisen as a correlated response to divergent sexual selection. Our results highlight the importance of sexual selection, rather than natural selection in the adaptive radiation of odonates, and supports previous suggestions that divergent sexual selection promotes speciation in this group.

Adaptation, Physiological↗

Horizontal gene transfer from transgenic plants to terrestrial bacteria--a rare event?

Today, 12 years after the first field release of a genetically modified plant (GMP), over 15,000 field trials at different locations have been performed. As new and unique characteristics are frequently introduced into GMPs, risk assessment has to be performed to assess their ecological impact. The possibilities of horizontal gene transfer (HGT; no parent-to-offspring transfer of genes) from plants to microorganisms are frequently evaluated in such risk assessments of GMPs before release into the field. In this review we indicate why putative HGT from plants to terrestrial (soil and plant associated) bacteria has raised concern in biosafety evaluations. Further, we discuss possible pathways of HGT from plants to bacteria, outline the barriers to HGT in bacteria, describe the strategies used to investigate HGT from plants to bacteria and summarize the results obtained. Only a few cases of HGT from eukaryotes such as plants to bacteria have been reported to date. These cases have been ascertained after comparison of DNA sequences between plants and bacteria. Although experimental approaches in both field and laboratory studies have not been able to confirm the occurrence of such HGT to naturally occurring bacteria, recently two studies have shown transfer of marker genes from plants to bacteria based on homologous recombination. The few examples of HGT indicated by DNA sequence comparisons suggest that the frequencies of evolutionarily successful HGT from plants to bacteria may be extremely low. However, this inference is based on a small number of experimental studies and indications found in the literature. Transfer frequencies should not be confounded with the likelihood of environmental implications, since the frequency of HGT is probably only marginally important compared with the selective force acting on the outcome. Attention should therefore be focused on enhancing the understanding of selection processes in natural environments. Only an accurate understanding of these selective events will allow the prediction of possible consequences of novel genes following their introduction into open environments.

Bacteria↗

A computer simulation of evolutionary forces controlling the size of a multigene family.

A Monte Carlo-type simulation of the evolution of a multigene family was performed. The model was designed to study the selective forces which may control the size of a multigene family. As expected, we find that direct selection on the size of the multigene family can control its size. More important, we find that selection acting upon the family as a single functional unit, in conjunction with homologous but unequal crossing over, can also control the size of a multigene family.

Animals↗

The influence of GAP promoter variants on hirudin production, average plasmid copy number and cell growth in Saccharomyces cerevisiae.

The yeast Saccharomyces cerevisiae has been engineered to synthesize and secrete desulfato-hirudin (hirudin), a thrombin inhibitor from the leech Hirudo medicinalis. The synthetic gene coding for hirudin was expressed constitutively under the control of four size-variants of the yeast glyceraldehyde-3-phosphate dehydrogenase promoter (GAP) and cloned into a 2 mu based multicopy yeast vector. The constitutive action of the four promoter variants was confirmed by demonstrating that the expression and secretion of hirudin is growth-related. The different efficiencies of the promoter variants not only affected hirudin expression but also led to changes in several cellular parameters, such as cell growth, average plasmid copy number and plasmid stability. The observed changes show that yeast cells establish a specific equilibrium for each promoter variant. We conclude, that the adjustment of cellular parameters in response to the expression levels of a heterologous protein is regulated by two counteracting selective forces: (1) the need for complementation of the auxotrophic host marker by the plasmid-encoded selection gene which, in the case of dLEU2, requires several plasmid copies; and (2) a selective advantage of cells with a lower copy number enabling them to escape the burden of heterologous protein production.

Base Sequence↗

Evidence of positive selection on the Atlantic salmon CD3gammadelta gene.

Atlantic salmon are typically anadromous, spending the majority of their lifetime in oceans and returning to fresh water to breed. This diversity of environments likely results in strong selective forces shaping their genome. In this paper, we present the first genomics approach to detect positive selection operating on the Salmo salar (salmon) lineage, an important aquaculture species. We identify a panel of candidate genes that may have been subject to adaptive evolution in this species. In particular, we identify a robust signature of positive selection operating on the salmon CD3gammadelta gene, which encodes one of the protein chains essential for formation of the T-cell receptor complex and for T-cell activation. Furthermore, we identified the particular codon sites that have been subject to positive selection in fish and highlight two sites flanking an important N-glycosylation site in this molecule.

Amino Acid Sequence↗

Near-neutrality in evolution of genes and gene regulation.

The nearly neutral theory contends that the interaction of drift and selection is important and occurs at various levels, including synonymous and nonsynonymous substitutions in protein coding regions and sequence turnover of regulatory elements. Recent progress of the theory is reviewed, and the interaction between drift and selection is suggested to differ at these different levels. Weak selective force on synonymous changes is stable, whereas its consequence on nonsynonymous changes depends on environmental factors. Selection on differentiation of regulatory elements is even more dependent on environmental factors than on amino acid changes. Of particular significance is the role of drift in the evolution of gene regulation that directly participates in morphological evolution. The range of near neutrality depends on the effective size of the population that is influenced by selected linked loci. In addition to the effective population size, molecular chaperones such as heat shock protein 90 have significant effects on the range of near neutrality.

Amino Acid Substitution↗

The potential for floral mimicry in rewardless orchids: an experimental study.

More than one-third of orchid species do not provide their pollinators with either pollen or nectar rewards. Floral mimicry could explain the maintenance of these rewardless orchid species, but most rewardless orchids do not appear to have a rewarding plant that they mimic specifically. We tested the hypothesis that floral mimicry can occur through similarity based on corolla colour alone, using naive bumble-bees foraging on arrays of plants with one rewarding model species, and one rewardless putative mimic species (Dactylorhiza sambucina) which had two colour morphs. We found that when bees were inexperienced, they visited both rewardless morphs randomly. However, after bees had gained experience with the rewarding model, and it was removed from the experiment, bees resampled preferentially the rewardless morph most similar to it in corolla colour. This is the first clear evidence, to our knowledge, that pollinators could select for floral mimicry. We suggest that floral mimicry can be a selective force acting on rewardless orchids, but only under some ecological conditions. In particular, we argue that selection on early-flowering rewardless orchids that receive visits from a large pool of naive pollinators will be weakly influenced by mimicry.

Adaptation, Biological↗