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Evolution of delayed reproductive senescence in male fruit flies: sperm competition.

Populations of Drosophila melanogaster that had been subjected to long-term selection favoring either delayed or rapid senescence were compared with respect to age-specific components of male reproductive success involving sperm competition. These components of reproductive success were divided into those related to sperm 'defense' (protection of sperm from other males), and into those related to sperm 'offense' (ability to mate with previously mated females and to displace the sperm of other males). Males were tested at four ages ranging from 1-2 d to 5-6 wk after eclosion. Several aspects of sperm defense capability showed clear evidence of senescent decline. Furthermore, males from populations selected for delayed senescence were superior to males from control (rapid senescence) populations with regard to components of sperm defense. The superiority of males from populations with delayed senescence either increased as a function of male age, or was present at all ages tested. These results indicate that the rate of reproductive senescence in male D. melanogaster can be altered in predictable directions by artificial selection. There were no differences between selection regimes with regard to sperm offense, and most components of sperm offense did not show clear evidence of senescence. The improved late-age reproductive success of males from populations selected for delayed senescence did not appear to entail any cost or trade-off at early ages with respect to the reproductive traits examined in these experiments.

Aging↗

In silico protein recombination: enhancing template and sequence alignment selection for comparative protein modelling.

Comparative modelling of proteins is a predictive technique to build an atomic model for a given amino acid sequence, on the basis of the structures of other proteins (templates) that have been determined experimentally. Critical problems arise in this procedure: selecting the correct templates, aligning the query sequence with them and building the non-conserved surface loops. In this work, we apply a genetic algorithm, with crossover and mutation, as a new tool to overcome the first two. In silico protein recombination proves to be an effective way to exploit the variability of templates and sequence alignments to produce populations of optimized models by artificial selection. Despite some limitations, the procedure is shown to be robust to alignment errors, while simplifying the task of selecting templates, making it a good candidate for automatic building of reliable protein models.

Algorithms↗

Genetic diversity, population structure in a historical panel of Brazilian soybean cultivars.

Soybean [Glycine max (L.) Merrill] is one of the most widely grown legumes in the world, with Brazil being its largest producer and exporter. Breeding programs in Brazil have resulted from multiple cycles of selection and recombination starting from a small number of USA cultivar ancestors in the 1950s and 1960s years. This process has led to the successful adaptation of this crop to tropical conditions, a phenomenon known as tropicalization. Many studies describe a narrow genetic background in Brazilian soybean cultivars. Various factors can affect the genetic diversity in species, especially in cultivated crops, such as the reproduction type, artificial selection, and the number and sources of variability in the breeding programs. In turns, the genetic diversity can affect the linkage disequilibrium blocks (LD) patterns and, consequently, molecular breeding strategies for selection of target loci for agronomic traits. We used high-throughput genotyping with SoySNP50K Illumina SNP markers to assess a collection of 370 Brazilian soybean accessions covering more than 60 years of soybean breeding in Brazil. Our goal was to investigate population structure and genetic diversity in the Brazilian germplasm, detect patterns of LD blocks, and identify regions presenting signals of selective swaps linked with quantitative trait loci (QTLs) of agronomic interest. Population structure analysis revealed two major groups among all genotypes, primarily differentiated by the year of release, separating old and new cultivars (before and after 2000´s years), and by growth habit (stem termination type-SST). The group I comprises about 75% of the panel and includes cultivars release before 2000`s years, including the oldest cultivars released in Brazil, most of which exhibit a determinate growth habit and maturity groups VI and VII. Group II includes only 83 materials, but shows higher levels of diversity than group I, representing most recent introductions in Brazilian germplasm. Further analysis of substructure within Group I, identified seven subgroups with no clear trend for segregation based on maturity group, STT or year of release. Instead, these subgroups were based on the contribution of key donors of disease resistance and adaptability, as soybean cultivation expanded from the South to Central region of Brazil. This finding is consistent with the history of soybean expansion in Brazil. We identified 123 genomic regions under selection among the groups of Brazilian cultivars associated with 440 quantitative trait loci (QTLs), revealing regions fixed across the breeding process associated with yield, disease resistance, water efficiency use, and others.

Glycine max↗

Long day plants and the response to global warming: rapid evolutionary change in day length sensitivity is possible in wild beet.

Day length is a key factor in flowering induction in many plant species in a seasonal environment with flowering induction usually happening at shorter day lengths in lower latitudes. Now, the climate changes systematically at a considerable speed due to global warming. As a consequence, earlier flowering will be selected for in long day plants by favouring a lower threshold for day length sensitivity, on the condition of available genetic variability. Here, we show that there is considerable genetic variation for day length sensitivity in our study species, the seabeet Beta vulgaris subsp. maritima. In the northernmost natural populations without vernalization requirement, in southwest France, the necessary day length for flowering induction could be reduced by artificial selection in <10 generations from >13 h to <11 h, the latter value corresponding to populations in the Beta-species complex from Northern Africa and the eastern part of the Mediterranean tested under the same conditions. A quantitative genetic analysis provided evidence of a gradual change without detectable major genes. Additional experiments were carried out to separate the response to photoperiod from age and energy effects. A certain effect of energy availability has been found, whereas age effects could be excluded. These results indicate a considerable potential for evolutionary change in adjusting flowering time in a changing climate.

Adaptation, Physiological↗

Long-bone biomechanics in mice selected for body conformation.

Two lines of mice divergently selected from the control strain (CBi) against the positive phenotypic correlation between body weight (b.w.) and tail (skeletal) length were obtained (CBi/C: high weight, short tail; CBi/L: low weight, long tail). The selected animals showed a different relationship between body and skeletal masses. To compare the adequacy between biomass and load-bearing ability of the skeleton, and to describe the eventual role of bone mechanostat in the production of these changes, cross-sectional and bending properties of both femur diaphyses were determined in CBi, CBi/C, and CBi/L adult mice of both genders. Cortical bone material quality (elastic modulus) was reduced in the selected lines (p < 0.001), significantly less in CBi/C than in CBi/L. In contrast, cross-sectional design (b.w.-adjusted values of moment of inertia, CSMI) was largely improved (p < 0.001), significantly more in CBi/C than in CBi/L. These effects determined a greater stiffness and strength in CBi/C than in CBi/L or CBi weight-paired mice. The elevations of the negative regression lines between elastic modulus and CSMI ("distribution/quality" curves) decreased in the order CBi/C > CBi/L > CBi. Data show that selection improved diaphyseal stiffness and strength in CBi/C animals because of an architectural overcompensation for the reduced bone material quality. Therefore, an inadequate control of long-bone architectural design as a function of the mechanical quality of cortical bone and b.w. bearing could have been induced in that line. Assuming bone mechanostatic regulation to be genetically programmed, some of the corresponding biological determinants should be transmitted independently, because artificial selection separately affected material quality and architectural design. The possibility of transmission of an inadequate mechanostatic function (inability to adapt bone modeling to bone material quality as a function of the biomass to be supported) was also shown, as some genotypes could express architectural modifications that largely exceed bone material quality deterioration.

Animals↗

Preparation of gellan sulfate as an artificial ligand for removal of extra domain A containing fibronectin.

The extra domain A containing fibronectin (EDA(+)FN) concentration in plasma of rheumatoid arthritis (RA) is abnormally higher than the normal level. We synthesized various gellan-sulfate (GS) candidates as artificial ligands for removing EDA(+)FN from plasma. The interaction between these artificial ligands and EDA(+)FN was evaluated using affinity constants (KA), which were determined by surface plasmon resonance measurement. The KA (3.6 x 10(8) per M) of GS-25 [degree of substitution for sulfonation (DS) = 25%] with EDA(+)FN was higher than those of other molecules: GS-16 (DS=16%) at 8.3 x 10(7) per M, and GS-35 (DS = 35%) at 1.7 x 10(8) per M. Furthermore, GSs displayed selectivity of EDA(+)FN for binding with plasma FN (KAEDA(+)FN)/KA(plasma FN)>2). The removal ratio in plasma was measured by using GS-immobilized gel. Removals of 66, 11, 7.7, 6.2, 6.9, and 12% for EDA(+)FN, plasma FN, fibrinogen, albumin, immunoglobulin G (IgG) and antithrombin III from the patient-model plasma were, respectively, achieved with GS-25-immobilized gel. These results suggest that GS may be used as a selective artificial ligand for EDA(+)FN removal from plasma in RA treatment.

Adsorption↗

Response of Sod-2 enzyme activity to selection for high voluntary wheel running.

The objective of this study was to examine the correlated response of anti-oxidant enzyme activity to selective breeding for increased voluntary wheel running in house mice. Activity of liver superoxide dismutase-2 (Sod-2), a free radical scavenger, was measured in four groups of mice. 'Active' individuals were housed in cages with attached wheels for 8 weeks beginning at weaning; 'sedentary' individuals were housed in cages with attached wheels that were prevented from rotating. Both of these treatments were applied to male and female mice from generation 14 of a replicated artificial selection experiment, which is composed of four lines selected for high wheel running and four randomly bred lines that serve as controls. In females, Sod-2 activity was significantly lower in selected vs control animals, regardless of presence/absence of a free-turning wheel. This difference suggests a trade-off between early-age voluntary wheel-running activity and Sod-2 activity. In males, Sod-2 activity was significantly affected by an interaction between selection group and activity group, with males from selected lines having lower Sod-2 activity relative to control males only in the sedentary treatment. These negative correlated responses of Sod-2 activity to selection on wheel running are discussed in the context of antagonistic pleiotropy models of aging and with respect to potential effects on lifespan.

Analysis of Variance↗

The power of evo-devo to explore evolutionary constraints: experiments with butterfly eyespots.

Examples of adaptive radiation by Darwinian evolution will always enthral, and the theory of natural selection binds the whole of biology. But is selection all-powerful? Evolutionary developmental biology is beginning to provide the background to understanding how the internal organisation of organisms can influence the tempo and direction of evolutionary change. Do the mechanisms that generate the phenotype channel, bias or limit morphological evolution? The formation of eyespots on the wings of the butterfly Bicyclus anynana is a process which is becoming comparatively well understood from the genetical, developmental and ecological perspectives. We have begun to combine this knowledge with artificial selection experiments to examine the rates at which morphological changes of a particular eyespot pattern in different directions can be achieved. Our initial selection experiment performed over twenty-five generations has shown that in contrast to predictions based on shared genetical and developmental properties, two butterfly eyespots on the same wing surface can be freely uncoupled from each other with respect to their pattern of relative sizes. This flexibility in development and in response to selection may occur because our base population is part of a lineage with a long legacy of natural selection building up genetic variation that enables independent behaviour for eyespot size, and thus it no longer reflects a tightly modular organisation of eyespots as may well have existed close to their evolutionary origin in basal Lepidoptera. Comparisons can be made between such descriptions of the potential for short-term changes in morphology in B. anynana and observed patterns of divergence among all extant species in this species-rich genus. Being able to map phenotypes onto genotypes via developmental processes provides a powerful basis for exploring genetical and developmental constraints. This will eventually lead to examples of evolutionary constraints that represent more than 'just-so' stories, and thus to a more balanced view of Darwinian evolution.

Journal Article↗

Metabolic models of selection response.

Consequences of directional selection on metabolic flux are explored in models for which variation in flux among individuals is generated by segregation of allelic variants at enzyme activity loci. The pattern of selection response is strongly affected by the presence of genetic dominance and epistasis, which are automatically generated in metabolic systems. The expected magnitudes of dominance and epistasis effects on flux are evaluated. Small differences in enzyme activity generate little dominance, but a null allele will tend to be recessive for the pathway in which it occurs and for metabolically distant pathways. Epistasis is found to be greatest in short pathways in which large differences in enzyme activity occur. Under divergent artificial selection asymmetrical responses can occur due to the presence of directional dominance and epistasis, and lead to departures from the classic infinitesimal model of quantitative genetic variation. The effects of epistasis and dominance are in opposite directions, however, and partially cancel each other out in a diploid population.

Alleles↗

Heritability of the timing of autumn migration in a natural bird population.

In recent decades, global temperature has increased at an unprecedented rate. This has been causing rapid environmental shifts that have altered the selective regimes determining the annual organization of birds. In order to assess the potential for adaptive evolution in the timing of autumn migration, we estimated heritabilities of the onset of migratory activity in a southern German blackcap (Sylvia atricapilla) population. Heritabilities (h2=0.34-0.45) and coefficients of additive genetic variation (CV(A)=4.7-5.7) were significant and consistent when estimated by different methods, irrespective of whether they were derived from birds hatched in the wild or bred in captivity. In an artificial selection experiment, we selected for later onset of migratory activity, simulating expected natural selection on this trait. We obtained a significant delay in the mean onset of migratory activity by more than one week after two generations of selection. Realized heritability (h2=0.55) was in agreement with expected heritability in the cohort that the selection line was derived from. Our results suggest that evolutionary changes in the timing of autumn migration may take place over a very short time period and will most probably be unconstrained by the lack of additive genetic variation.

Acclimatization↗

Complete maternal inheritance of bifenazate resistance in Tetranychus urticae Koch (Acari: Tetranychidae) and its implications in mode of action considerations.

Bifenazate is a selective hydrazine carbazate acaricide launched in 1999 and reported to be neurotoxic, since preliminary studies on the mode of action suggested that bifenazate may act on GABA-gated chloride channels. However, this information has not yet been supported by mechanistic studies. Therefore bifenazate is still considered as a neuronal inhibitor, but with unknown mode of action. Here we report an alternative hypothesis on the mode of action of bifenazate, i.e. its possible interference with a non-neuronal target site. An acaricide susceptible strain of Tetranychus urticae Koch (Acari: Tetranychidae), LS-VL, was artificially selected for bifenazate resistance, and after 36 generations an extremely high resistance ratio (RR) of >164,000 was obtained. This bifenazate-resistant strain (BR-VL) lacks cross-resistance to many different chemical classes and modes of action of other acaricides. In order to check for metabolic resistance mechanisms, synergists known to inhibit well-known detoxification routes were used together with in vitro enzymatic assays. No synergism or highly increased detoxification activity was observed in the resistant strain. However, the organophosphorous esterase inhibitor S,S,S-tributylphosphorotrithioate (DEF) applied to the susceptible strain could completely antagonise the acaricidal efficacy of bifenazate, suggesting that bifenazate is a pro-acaricide, not active by itself, that needs in vivo activation by esterases. Reciprocal crosses of diploid females and haploid males of strains LS-VL (susceptible) and BR-VL (bifenazate resistant) revealed that bifenazate resistance was inherited completely maternally, i.e. resistance is fully dominant when susceptible males were crossed with resistant females, and fully recessive when resistant males were crossed with susceptible females. Such an inheritance pattern has to our knowledge never been observed before in the case of insecticide/acaricide resistance. This observation may suggest a target-site for bifenazate encoded by the mitochondrial genome. Further evidence supporting such a hypothesis was obtained when measuring the ATP-level in spider mites treated with bifenazate. The ATP content in bifenazate treated mites declined progressively between 0 and 4h after treatment, similarly to mites treated with the complex I inhibitor fenpyroximate, an acaricide known to interfere with mitochondrial function. The obtained results suggest a target-site other than GABA-gated chloride channels, most likely encoded by and located in the mitochondria.

Adenosine Triphosphate↗

Genomic imprinting of the insulin-like growth factor 2 gene in sheep.

A number of genes in the human and mouse genomes are subject to genomic imprinting, with selective inactivation of one allele of a gene in a parent-of-origin specific manner. One of the first imprinted genes identified was the Insulin-like Growth Factor 2 gene (IGF2), which promotes growth of the fetus and is expressed from only the paternal allele in most tissues in both the mouse and human. The aim of this study was to establish the imprinting status of IGF2 in sheep (Ovis aries). Sheep provide an interesting model to study imprinting, owing to differences in their placental development and the fact that they have been subject to strong artificial selection for various production traits. We report the identification of a length polymorphism in the transcribed 3'-untranslated region of the ovine IGF2 gene. This polymorphism was used to map IGF2 to sheep Chromosome (Chr) 21 and demonstrate that IGF2 is indeed imprinted in sheep, being expressed from the paternal allele. We also report that the developmental switch from imprinted IGF2 expression in the fetal liver to biallelic IGF2 expression in the adult liver, which occurs in the human but not mouse, also occurs in sheep. Differences in male- and female-specific recombination values reported around the IGF2 locus in the human were also observed around the ovine IGF2 locus. The techniques developed in this study will enable the imprinting status of IGF2 to be assessed in a variety of tissues and stages of development in normal sheep.

3' Untranslated Regions↗

Evolutionary ecology of egg size and number in a seed beetle: genetic trade-off differs between environments.

In many organisms, large offspring have improved fitness over small offspring, and thus their size is under strong selection. However, due to a trade-off between offspring size and number, females producing larger offspring necessarily must produce fewer unless the total amount of reproductive effort is unlimited. Because differential gene expression among environments may affect genetic covariances among traits, it is important to consider environmental effects on the genetic relationships among traits. We compared the genetic relationships among egg size, lifetime fecundity, and female adult body mass (a trait linked to reproductive effort) in the seed beetle, Stator limbatus, between two environments (host-plant species Acacia greggii and Cercidium floridum). Genetic correlations among these traits were estimated through half-sib analysis, followed with artificial selection on egg size to observe the correlated responses of lifetime fecundity and female body mass. We found that the magnitude of the genetic trade-off between egg size and lifetime fecundity differed between environments--a strong trade-off was estimated when females laid eggs on C. floridum seeds, yet this trade-off was weak when females laid eggs on A. greggii seeds. Also differing between environments was the genetic correlation between egg size and female body mass-these traits were positively genetically correlated for egg size on A. greggii seeds, yet uncorrelated on C. floridum seeds. On A. greggii seeds, the evolution of egg size and traits linked to reproductive effort (such as female body mass) are not independent from each other as commonly assumed in life-history theory.

Analysis of Variance↗

The effect of testosterone in mice divergently selected on fat content or body weight.

Lines of mice have been divergently selected on one of two traits: (i) estimated fat content at 14 weeks of age, which has resulted in a 5-fold divergence, and (ii) body weight at 10 weeks of age, which has resulted in a 3-fold divergence. Individuals from each line were castrated or sham operated at 10 days of age and subsequently given either exogenous testosterone or the appropriate control from 14 days of age. Castration increased fat content and decreased lean weight in all lines, an effect which was not reversed by administration of testosterone. Body weight was reduced by around 10% as a result of castration and this effect was at least partially reversed by exogenous testosterone. Analysis of body weight, fat content and lean mass at 10 weeks of age failed to detect any interaction between these treatments and genetic background. It is therefore concluded that testosterone metabolism has not contributed disproportionately to the response to artificial selection in spite of its known effects on growth and body composition.

Adipose Tissue↗

Allomorphic relationships from hatching to 56 days in parental lines and F1 crosses of chickens selected 27 generations for high or low body weight.

Allomorphic relationships in chickens selected for high or low juvenile body weight and their reciprocal crosses were examined from hatch to 56 days of age (doa). Those organs whose relationships to whole body weight were most divergent in the parental populations were weights of the gizzard, small intestine, breast, legs, feathers, abdominal fat and lengths of the shank, esophagus and small intestine. Relative weights of feathers, abdominal fat and small intestine were influenced mainly by additive genetic variation, whereas relative weights of breast, legs, and gizzard were influenced by additive genetic variation at younger ages (0 to 24 doa) and by both additive and non-additive genetic variation thereafter. In utilization of resources for high growth velocity and growth rate factor, crosses were 20 to 30% more efficient than their parental populations, demonstrating hybrid vigor. Relationships of rate of development of specific organs to total body weight varied with age and were noted as correlated responses to artificial selection for body weight. Selection for high and for low body weight at 56 days of age resulted in compromises in resource allocations. Specifically, selection for high juvenile body weight favored relatively heavier breasts, legs, abdominal fat depots and small intestines while selection for low juvenile body weight favored relatively heavier feathers and gizzard.

Animals↗

Further evidence for mirror-reversed laterality in lines of fish selected for leftward or rightward turning when facing a predator model.

In the teleost fish Girardinus falcatus eye preferences for inspecting a potential predator is highly heritable and this consented to select lines with opposed laterality. In previous studies individuals from a RD line (rightward turning when facing a dummy predator) and those from a LD line (leftward turning) were subjected to several other laterality tests (most of which, possibly all, were visually based). Since they obtained opposite scores in all tests, it was suggested that LD and RD fish have complete mirror-reversed organizations of the brain. Here, we studied fish from selected lines in a new set of laterality tests including some non-visual tests. They included measures of (1) rotational preference in the home tank (2) direction of spontaneous swimming in the dark (3) escape trajectories after delivery of an auditory stimulus (4) escape turning direction to a fast approaching visual stimulus. The results of the first three tests are congruent with the finding of previous studies in that fishes of the two lines showed opposite direction of lateralisation. When tested for laterality in the escape response to a fast approaching stimulus, fish of the LD and RD lines showed no differences and both were biased toward leftward escape. Overall these results suggest the existence in G. falcatus of a single mechanism controlling a co-ordinate placement of the great majority of lateralised functions. Yet the results of the fourth test suggest that a few lateralized functions are controlled by different mechanisms that were unaffected by artificial selection.

Acoustic Stimulation↗

Avian leukosis virus and selection on oviposition interval in Australorp lines.

1. Shedding of group-specific antigen of avian leukosis virus (ALV) into egg albumen was determined in three Australorp lines: AS line that had been selected primarily for short oviposition interval, ASS line that had been derived from the AS line and developed as a commercial dam line for egg laying, and AC line that had been kept as a randombred control. 2. The proportion of shedders was 13.1 to 16.7% in the AS line in 1984-88, 16.3% in the ASS line in 1984 (before culling of the shedders), and 6.1% and 6.6% in the AC line in 1984 and 1988, respectively. 3. In the AS line, shedders were 1.8% lower in rate of lay to 300 d of age, 1.3 g lower in average egg weight at 34 weeks of age, 5.6% lower in hatchability of fertile eggs and 0.24 h shorter in oviposition interval than non-shedders. In the ASS line (1984 only), the differences between shedders and non-shedders were in the same direction, but in magnitude greater for rate of lay and smaller for oviposition interval. 4. The shedders were favoured by the artificial selection because of their shorter oviposition interval and this appeared to be responsible for the higher levels of ALV shedding in the selection lines.

Animals↗

Selection, evolution of behavior and animal models in behavioral neuroscience.

We investigated whether genetic differences in various forms of intraspecific aggression and anxiety in four different genetic lines of mice (i.e. wild, outbred Swiss-CD1, inbred DBA/2 and inbred C57/BL6N) may reflect modifications in behavioral strategy. Experiments 1 and 2 used ethologically based paradigms to analyze aggressive and anxiety responses both in social (i.e. aggression) and non-social (i.e. novel environment exploration) contexts. In Experiment 3, an anxiolytic drug (chlordiazepoxide (CDP)) was used to examine possible differences in proximal mechanisms underlying anxiety-related behaviors. The data show that intrasexual aggression, infanticide and maternal aggressions are related and covarying. Genetic lines with the highest levels of intermale attack (i.e. Wild and Swiss-CD1) also have highest levels of infanticide, interfemale attack and maternal aggression but, interestingly, the lowest levels of anxiety. In fact, exploratory behavior is lower and risk assessment behavior markedly higher in DBA/2 and C57/BL6N mice (i.e. the less aggressive strains) compared to Swiss and Wild genetic lines. Although reproductive status influences anxiety levels in female mice, our findings show that (contrary to previous studies) lactating mice are more anxious than virgin females in terms of risk assessment activities. These data demonstrate the importance of studying behavior in a more ecologically-relevant context which emphasizes the function of behavior in a specific situation. Moreover, differential strain sensitivity to the behavioral effects of CDP suggests that genetic lines of mice may differ in the underlying mechanisms mediating behavior. It is therefore possible that artificial selection of different genotypes has resulted in differences in proximate mechanisms modulating the levels of aggression and anxiety, thereby leading to modification of social behavior. Overall, the results presented here suggest that subtle genetic alterations in specific underlying neural mechanisms are likely to cause profound effects on behavioral responses and their adaptive significance. Implications for behavioral neuroscience research that seeks to understand both the proximal and ultimate mechanisms of behavior are discussed.

Animals↗