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Biomedical subjects

B Sinervo

Publications and source records attributed to B Sinervo.

At least 19 recordsLinked to original sources
PubMed2007

Colour variation and alternative reproductive strategies in females of the common lizard Lacerta vivipara.

Within-sex colour variation is a widespread phenomenon in animals that often plays a role in social selection. In males, colour variation is typically associated with the existence of alternative reproductive strategies. Despite ecological conditions theoretically favourable to the emergence of such alternative strategies in females, the social significance of colour variation in females has less commonly been addressed, relative to the attention given to male strategies. In a population of the common lizard, females display three classes of ventral colouration: pale yellow, orange and mixed. These ventral colours are stable through individual's life and maternally heritable. Females of different ventral colourations displayed different responses of clutch size, clutch hatching success and clutch sex-ratio to several individual and environmental parameters. Such reaction patterns might reflect alternative reproductive strategies in females. Spatial heterogeneity and presence of density- and frequency-dependent feedbacks in the environment could allow for the emergence of such alternative strategies in this population and the maintenance of colour variation in females.

Animals↗
PubMed2003

Social competition, corticosterone and survival in female lizard morphs.

We examined the selective consequences of variation in behaviour and endocrine physiology in two female throat-colour morphs of the lizard, Uta stansburiana in the wild. Female morphs differed in home-range distribution patterns and corticosterone levels in relation to the density and frequency of their female neighbours. Levels of plasma corticosterone of yellow-throated females increased with increased density of both morphs. In contrast, orange-throated females had reduced levels of corticosterone in response to increased density of orange females. Additionally, females with lower corticosterone survived poorly, suggesting that social interactions and high local densities of orange females may be potentially costly for orange females. These results are consistent with decreased fitness effects and suppression of immune function previously reported for orange female morphs surrounded by more orange neighbours. These correlations, in conjunction with previous work in this system, indicate that corticosterone is likely to be an important physiological mechanism regulating female fitness in nature.

Animals↗
PubMed2003

Behavioral drive versus behavioral inertia in evolution: a null model approach.

Some biologists embrace the classical view that changes in behavior inevitably initiate or drive evolutionary changes in other traits, yet others note that behavior sometimes inhibits evolutionary changes. Here we develop a null model that quantifies the impact of regulatory behaviors (specifically, thermoregulatory behaviors) on body temperature and on performance of ectotherms. We apply the model to data on a lizard (Anolis cristatellus) and show that thermoregulatory behaviors likely inhibit selection for evolutionary shifts in thermal physiology with altitude. Because behavioral adjustments are commonly used by ectotherms to regulate physiological performance, regulatory behaviors should generally constrain rather than drive evolution, a phenomenon we call the "Bogert effect." We briefly review a few other examples that contradict the classical view of behavior as the inevitable driving force in evolution. Overall, our analysis and brief review challenge the classical view that behavior is invariably the driving force in evolution, and instead our work supports the alternative view that behavior has diverse--and sometimes conflicting--effects on the directions and rates at which other traits evolve.

Altitude↗
PubMed2002

Correlational selection and the evolution of genomic architecture.

We review and discuss the importance of correlational selection (selection for optimal character combinations) in natural populations. If two or more traits subject to multivariate selection are heritable, correlational selection builds favourable genetic correlations through the formation of linkage disequilibrium at underlying loci governing the traits. However, linkage disequilibria built up by correlational selection are expected to decay rapidly (ie, within a few generations), unless correlational selection is strong and chronic. We argue that frequency-dependent biotic interactions that have 'Red Queen dynamics' (eg, host-parasite interactions, predator-prey relationships or intraspecific arms races) often fuel chronic correlational selection, which is strong enough to maintain adaptive genetic correlations of the kind we describe. We illustrate these processes and phenomena using empirical examples from various plant and animal systems, including our own recent work on the evolutionary dynamics of a heritable throat colour polymorphism in the side-blotched lizard Uta stansburiana. In particular, male and female colour morphs of side-blotched lizards cycle on five- and two-generation (year) timescales under the force of strong frequency-dependent selection. Each morph refines the other morph in a Red Queen dynamic. Strong correlational selection gradients among life history, immunological and morphological traits shape the genetic correlations of the side-blotched lizard polymorphism. We discuss the broader evolutionary consequences of the buildup of co-adapted trait complexes within species, such as the implications for speciation processes.

Animals↗
PubMed2001

Density-dependent competition and selection on immune function in genetic lizard morphs.

Density-dependent territorial interactions have been suggested to cause immunosuppression and thereby decrease fitness, but empirical support from natural populations is lacking. Data from a natural lizard population (Uta stansburiana) showed that breeding females surrounded by many territorial neighbors had suppressed immune function. Furthermore, variation in immunological condition had different effects on the fitness of the two heritable female throat-color morphs in this population. These interactive fitness effects caused correlational selection between female throat color and immune responsiveness. Population genetic theory predicts that this should have lead to the buildup and preservation of a genetic correlation between female morphotype and immunological condition. Accordingly, the throat color of a female was genetically correlated (r(A) = -1.36; SE = 0.55) with her daughter's immune responsiveness.

Journal Article↗
PubMed2001

Social causes of correlational selection and the resolution of a heritable throat color polymorphism in a lizard.

When selection acts on social or behavioral traits, the fitness of an individual depends on the phenotypes of its competitors. Here, we describe methods and statistical inference for measuring natural selection in small social groups. We measured selection on throat color alleles that arises from microgeographic variation in allele frequency at natal sites of side-blotched lizards (Uta stansburiana). Previous game-theoretic analysis indicates that two color morphs of female side-blotched lizards are engaged in an offspring quantity-quality game that promotes a density- and frequency-dependent cycle. Orange-throated females are r-strategists. They lay large clutches of small progeny, which have poor survival at high density, but good survival at low density. In contrast, yellow-throated females are K-strategists. They lay small clutches of large progeny, which have good survival at high density. We tested three predictions of the female game: (1) orange progeny should have a fitness advantage at low density; (2) correlational selection acts to couple color alleles and progeny size; and (3) this correlational selection arises from frequency-dependent selection in which large hatchling size confers an advantage, but only when yellow alleles are rare. We also confirmed the heritability of color, and therefore its genetic basis, by producing progeny from controlled matings. A parsimonious cause of the high heritability is that three alleles (o, b, y) segregate as one genetic factor. We review the physiology of color formation to explain the possible genetic architecture of the throat color trait. Heritability of color was nearly additive in our breeding study, allowing us to compute a genotypic value for each individual and thus predict the frequency of progeny alleles released on 116 plots. Rather than study the fitness of individual progeny, we studied how the fitness of their color alleles varied with allele frequency on plots. We confirmed prediction 1: When orange alleles are present in female progeny, they have higher fitness at low density when compared to other alleles. Even though the difference in egg size of the female morphs was small (0.02 g), it led to knife-edged survival effects for their progeny depending on local social context. Selection on hatchling survival was not only dependent on color alleles, but on a fitness interaction between color alleles and hatchling size, which confirmed prediction 2. Sire effects, which are not confounded by maternal phenotype, allowed us to resolve the frequency dependence of correlational selection on egg size and color alleles and thereby confirmed prediction 3. Selection favored large size when yellow sire alleles were rare, but small size when they were common. Correlational selection promotes the formation of a self-reinforcing genetic correlation between the morphs and life-history variation, which causes selection in the next density and frequency cycle to be exacerbated. We discuss general conditions for the evolution of self-reinforcing genetic correlations that arise from social selection associated with frequency-dependent sexual and natural selection.

Alleles↗
PubMed2001

Condition, genotype-by-environment interaction, and correlational selection in lizard life-history morphs.

We compared reproductive allocation and variation in condition and survivorship of two heritable female throat color morphs (orange and yellow) in a free-living population of side-blotched lizards (Uta stansburiana). Using path analysis and structural equation modeling, we investigated how variation in the social environment affected clutch size and egg mass and two condition traits (postlaying mass, immunological condition) and how these traits in turn affected female field survival. In the presence of many neighbors, both morphs increased their clutch sizes, although these effects were only significant in yellow females. In addition, yellow females increased their egg mass in the presence of many orange neighbors. Orange females surrounded by many orange neighbors showed sign of stress in the form of immunosuppression, whereas this effect was less pronounced in yellow females. The morphs also differed in the impact of variation in clutch size and egg mass on both condition traits. Finally, female morphotype and immune responsiveness affected fitness interactively, and hence these two traits showed signs of fitness epistasis: Selection gradients on this trait were opposite in sign in the two morphs. The correlational selection gradient (gamma throat x antibody response) between female throat color and antibody responsiveness was -0.365. Our data thus reveal important interactive effects such as genotype-by-environment interaction toward the social environment and morph-specific trade-offs as well as the occurrence of correlational selection. We discuss the use of naturally occurring and conspicuous genetic polymorphisms in field studies of selection and life-history allocation.

Animals↗
PubMed2001

Runaway social games, genetic cycles driven by alternative male and female strategies, and the origin of morphs.

Analysis of evolutionarily stable strategies (ESS) and decade-long field studies indicate that two color morphs of female side-blotched lizards exhibit density- and frequency-dependent strategies. Orange females are r-strategists: they lay large clutches of small progeny that are favored at low density. Conversely, yellow females are K-strategists: they lay small clutches of large progeny that are favored when carrying capacity is exceeded and the population crashes to low density. Interactions among three male morphs resembles a rock-paper-scissors (RPS) game. Fertilization success of males depends on frequency of neighboring morphs. Orange males usurp territory from blue neighbors and thereby mate with many females. However, orange males are vulnerable to cuckoldry by sneaky yellow males that mimic females. The yellow strategy is thwarted in turn by the mate-guarding strategy of blue. Sinervo and Lively (1996) developed a simple asexual model of the RPS game. Here, we model the dynamics of male and female morphs with one- and two-locus genetic models. Male and female games were considered in isolation and modeled as games that were genetically coupled by the same locus. Parameters for payoff matrices, which describe the force of frequency-dependent selection in ESS games, were estimated from free-ranging animals. Period of cycles in nature was 5 years for males and 2 years for females. Only the one locus model with three alleles (o, b, y) was capable of driving rapid cycles in male and female games. Furthermore, the o allele must be dominant to the y allele in females. Finally, the amplitude of male cycles was only reproduced in genetic models which allowed for irreversible plasticity of by genotypes, which is consistent with hormonally-induced changes that transform some males with yellow to dark blue. We also critique experimental designs that are necessary to detect density- and frequency-dependent selection in nature. Finally, runaway ESS games are discussed in the context of self-reinforcing genetic correlations that build and promote the formation of morphotypic variation.

Alleles↗
PubMed2000

Polygyny, mate-guarding, and posthumous fertilization as alternative male mating strategies.

Alternative male mating strategies within populations are thought to be evolutionarily stable because different behaviors allow each male type to successfully gain access to females. Although alternative male strategies are widespread among animals, quantitative evidence for the success of discrete male strategies is available for only a few systems. We use nuclear microsatellites to estimate the paternity rates of three male lizard strategies previously modeled as a rock-paper-scissors game. Each strategy has strengths that allow it to outcompete one morph, and weaknesses that leave it vulnerable to the strategy of another. Blue-throated males mate-guard their females and avoid cuckoldry by yellow-throated "sneaker" males, but mate-guarding is ineffective against aggressive orange-throated neighbors. The ultradominant orange-throated males are highly polygynous and maintain large territories; they overpower blue-throated neighbors and cosire offspring with their females, but are often cuckolded by yellow-throated males. Finally, yellow-throated sneaker males sire offspring via secretive copulations and often share paternity of offspring within a female's clutch. Sneaker males sire more offspring posthumously, indicating that sperm competition may be an important component of their strategy.

Animals↗
PubMed2000

Density cycles and an offspring quantity and quality game driven by natural selection.

A long-standing hypothesis posits that natural selection can favour two female strategies when density cycles. At low density, females producing many smaller progeny are favoured when the intrinsic rate of increase, r, governs population growth. At peak density, females producing fewer, high-quality, progeny are favoured when the carrying capacity, K, is exceeded and the population crashes. Here we report on the first example of a genetic r versus K selection game that promotes stable population cycles in lizards. Decade-long fitness studies and game theory demonstrated that two throat-colour morphs were refined by selection in which the strength of natural selection varied with density. Orange-throated females, r strategists, produced many eggs and were favoured at low density. Conversely, yellow-throated females, K strategists, produced large eggs and were favoured at high density. Progeny size should also be under negative frequency-dependent selection in that large progeny will have a survival advantage when rare, but the advantage disappears when they become common. We confirmed this prediction by seeding field plots with rare and common giant hatchlings. Thus, intrinsic causes of frequency- and density-dependent selection promotes an evolutionary game with two-generation oscillations.

Animals↗
PubMed2000

Testosterone, endurance, and Darwinian fitness: natural and sexual selection on the physiological bases of alternative male behaviors in side-blotched lizards.

The mechanistic bases of natural and sexual selection on physiological and behavioral traits were examined in male morphs of three colors of the side-blotched lizard, Uta stansburiana. Orange-throated males are aggressive and defend large territories with many females. Blue-throated males defend smaller territories with fewer females; however, blue-throated males assiduously mate guard females on their territory. Yellow-throated males do not defend a territory, but patrol a large home range. They obtain secretive copulations from females on the territories of dominant males. Males with bright orange throats had higher levels of plasma testosterone (T), endurance, activity, and home range size and concomitantly gained greater control over female home ranges than blue- or yellow-throated males. Experimentally elevating plasma T in yellow- and blue-throated males increased their endurance, activity, home range size, and control over female territories to levels that were seen in unmanipulated orange-throated males that had naturally high plasma T. However, the enhanced performance of orange-throated males is not without costs. Orange-throated males had low survival compared to the other morphs. Finally, some yellow-throated males transformed to a partial blue morphology late in the season and the endurance of these transforming yellow-throated males increased from early to late in the season. In addition, yellow-throated males that transformed to blue also had significantly higher plasma T late in the season compared to the plasma T earlier in the season. T appears to play an important role in the physiological changes that all three color morphs undergo during the process of maturation. In some yellow males, T plays an additional role in plastic changes in behavior and physiology late in the reproductive season. We discuss natural and sexual selection on physiological and behavioral traits that leads to the evolution of steroid regulation in the context of alternative male strategies.

Aggression↗
PubMed2000

Reproductive burden, locomotor performance, and the cost of reproduction in free ranging lizards.

A reduction in the locomotor capacity of gravid females is considered to be a cost of reproduction if it leads to an increased risk of mortality. In this study, we measured the change in endurance between gravid and postgravid female side-blotched lizards (Uta stansburiana) as a test of the cost of reproduction. We also altered reproductive investment in some females by direct ovarian manipulation (yolkectomy), which decreased reproductive burden by 30%. Regardless of experimental treatment, all females had lower endurance when gravid. Endurance was 28% lower in gravid females from the yolkectomy treatment and 31% lower in the unmanipulated females relative to postoviposition females. The experimental reduction in clutch mass resulted in a 21% increase in endurance of gravid yolkectomy females relative to control females. Postovipositional endurance was significantly higher in the yolkectomized females than unmanipulated females, which suggests that the cost of reproduction carries over to postoviposition performance. Unmanipulated females exhibited a significant negative association between endurance and size-specific burden. Endurance was not correlated with clutch size or size-specific burden in the yolkectomy females. Survivorship to the second clutch was higher in the yolkectomy females. The results from a logistic regression showed the probability of survival to the second clutch was significantly and positively associated with endurance after controlling for the effects of treatment. Our analyses demonstrated that the decrement in performance associated with current reproductive investment represents a cost of reproduction expressed as diminished locomotor performance and lowered survivorship to the next clutch.

Analysis of Variance↗
PubMed2000

Experimental excursions on adaptive landscapes: density-dependent selection on egg size.

Theories of density-dependent natural selection suggest that intraspecific competition will favor juveniles of high competitive ability. Empirical evidence has been provided from laboratory selection experiments, but field studies are lacking due to the logistical difficulties of experimentally manipulating population densities in natural settings. Here, we present data from a decade-long experimental field study of side-blotched lizards, Uta stansburiana that overcomes these difficulties. We tested the hypothesis that density-dependent natural selection causes egg size to increase from early to late clutches in this and many other species. Using a novel combination of environmental manipulations of hatchling density and phenotypic manipulations of egg size, we demonstrate that the nature of selection on egg size changes dramatically in the absence of older competitors. The strength of selection on egg size among later-clutch hatchlings released in areas without competitors from early clutches became almost doubled in magnitude, compared to that among hatchlings released in the presence of older competitors. These experimental findings demonstrate density-dependent natural selection on egg size; however, they contradict the classical idea that egg size increases during the reproductive season because of competition between early and late hatchlings. The results indicate that competitive age or size asymmetries between early and late hatchlings can override within-cohort asymmetries due to egg size. We suggest that competition could be an important mediator of oscillating selection pressures in this and other systems. Finally, we discuss the utility of "double-level," simultaneous experimental manipulation of both phenotypic traits that are targets of selection (e.g., egg size) as well the environmental agents of selection (e.g., population density).

Adaptation, Physiological↗
PubMed1996

Role of Gene Interactions in Hybrid Speciation: Evidence from Ancient and Experimental Hybrids

The origin of a new diploid species by means of hybridization requires the successful merger of differentiated parental species' genomes. To study this process, the genomic composition of three experimentally synthesized hybrid lineages was compared with that of an ancient hybrid species. The genomic composition of the synthesized and ancient hybrids was concordant (rs = 0.68, P < 0.0001), indicating that selection to a large extent governs hybrid species formation. Further, nonrandom rates of introgression and significant associations among unlinked markers in each of the three synthesized hybrid lineages imply that interactions between coadapted parental species' genes constrain the genomic composition of hybrid species.

Journal Article↗
PubMed1994

Effects of corticosterone on activity and home-range size of free-ranging male lizards.

The purpose of our study was to examine whether corticosterone (B) affects the spacing behavior of free-ranging male side-blotched lizards (Uta stansburiana). Furthermore, we wanted to determine if the density, reflected in seasonal changes in population size, or behavior, as a result of hormonal manipulation, of "neighboring" males influences these effects. Field studies were conducted on four naturally isolated "neighborhoods" of lizards. Half of the males on three of the sites were randomly implanted with either saline or B, while on the fourth site all males were implanted with B. Pre- and postimplant home-ranges and activity levels were determined. Home-range size and activity level were significantly reduced by corticosterone if normally aggressive saline-implanted males were also present in the neighborhood. However, B had no effect on home-range if all males in the neighborhood were implanted with B. Space lost by B-implanted males was incorporated into the home-ranges of saline-implanted males so that the sum of all the male home-ranges in a neighborhood remained unchanged after implantation. These results suggest that elevated B levels put male lizards at a competitive disadvantage and, therefore, force these lizards to reduce their home-range.

Animals↗
PubMed1994

Effects of steroid hormone interaction on activity and home-range size of male lizards.

Steroid hormones play a major role in influencing the physiology and behavior of all animals, including reptiles. Oftentimes, it is an interaction between two or more hormones that is ultimately responsible for the observed response or behavior. We designed a pair of field studies on adjacent communities of side-blotched lizards (Uta stansburiana) to provide insight into the interrelationship of testosterone (T) and corticosterone (B) in regulating aggressive behavior. On one site, males were implanted with either T or saline (S), while on the other site males received either two S implants or both a T and a B implant (T + B). T increased both activity (by 31%) and home-range size (by 150%), whereas S-implanted cohorts exhibited a reduction in both of these parameters (by 24 and 50%, respectively). However, when B was given in combination with T, not only were the positive effects of T eliminated, but there was a reduction in activity (31%) and home-range size (72%) similar to that reported in lizards that received B implants alone. S-implanted cohorts in the T + B experiment increased their activity and home-range size by 15 and 43%, respectively. Although these latter changes in the S-implanted males are not statistically significant, they are indicative of a compensatory increase similar to that seen in the T and previously reported B outcrop experiments. Taken together, these results illustrate that regulation of aggressive behavior is complicated, with both hormonal and social interactions playing critical roles in determining an individual's home-range size and, hence, reproductive success.

Aggression↗
PubMed1989

Experimental phenocopy of a minute maternal-effect mutation alters blastoderm determination in embryos of Drosophila melanogaster.

Maternal haploinsufficiency for a third chromosome Minute, M(3)i55, lowers rates of protein synthesis by approximately 30% during the syncytial nuclear cycles of early embryogenesis. The maternal effect of Mi55 also produces segmentation defects (denticle belt fusions) in the posterior abdomen of larvae. Furthermore, embryos from Minute mothers show abnormal expression patterns of the segmentation gene fushi tarazu (ftz) at the cellular blastoderm stage of embryogenesis. We developed a computer-aided analysis to describe the deviations in ftz expression which demonstrates that abnormally narrow ftz stripes occur in segment primordia that become fused in the larva. Unexpectedly, an abnormally wide ftz stripe occurs in segment primordia which do not develop abnormally. In addition, Mi55 produces a general narrowing of all ftz- interstripes. We phenocopied the Minute mutation by injecting wild-type embryos with cycloheximide concentrations which decreased protein synthesis rates to levels comparable with those of Minute embryos. Thus, a general decrease in protein synthesis during early embryogenesis leads to abnormal determination of posterior abdominal segment primordia.

Abdomen↗