Summary1. Natural selection can generate correlated suites of phenotypic traits by acting independently on physiological and behavioural characters or on mechanisms that exert pleiotropic actions. 2. Current theory, supported by artificial selection studies, suggests that physiological and behavioural stress responses are at least partially under genetic control and covary in a predictable manner. Indeed, physiological mechanisms such as hormonal stress responsiveness may underlie variation in behaviour, including consistent behaviours described as temperament or personality, with bolder, more exploratory and active individuals being less hormonally responsive to stressors. 3. This relationship, however, has yet to be demonstrated in natural populations. We investigated the relationship between hormonal and behavioural stress responsiveness in multiple natural populations of a tropical freshwater poeciliid fish, Brachyrhaphis episcopi, that experience different levels of predation pressure and hence encounter different rates of stressful events. Predation can impose a strong selection pressure, and living with a high risk of predation is known to select for specific behavioural traits. 4. We quantified variation in stress responsiveness via cortisol release rates (exp. 1) and behaviour in an open field test followed by cortisol release rates (exp. 2). Populations exposed to high levels of predation were consistently more exploratory and active and had lower release rates of cortisol in response to a stressor than conspecifics sampled at sites with few predators. 5. However, this difference in stress responsiveness was only apparent after fish had experienced the mild stress of behaviour testing (in exp. 2), which resulted in elevation of cortisol levels. The relationship between hormone release and behaviour was also not apparent within populations once independent factors were controlled for, highlighting the importance of factors such as size and sex on individual variability. 6. This study demonstrates that the relationship between hormonal and behavioural stress responsiveness can result from natural selection pressures, such as that imposed by predation.
Imbalances in phosphorus (P) intake relative to demand negatively affect animal growth, but their consequences are less understood for vertebrates, in which bone represents a significant and potentially flexible pool of P. Flexibility in body-P content could buffer vertebrates from the effects of imbalances between P intake and demand, reducing the likelihood of a sharp stoichiometric ''knife-edge'' in the relationship between growth rate and diet-P level. We conducted a meta-analysis of published aquaculture experiments that tested effects of diet %P on fish growth rate (49 studies, 28 species) and body-P content (27 of the studies in the main data set, 20 species). Our meta-analysis revealed significant P limitation of growth, as well as significant negative effects of excess P on growth rate. Diet-P thresholds for these effects occurred at ecologically relevant levels (optimal diet-P of 1.2% 6 0.45%, mean 6 SD, under experimental conditions of high ration). Finally, the analysis also suggested a pattern of relatively shallow relationships between growth rate and diet-P level, coupled with surprisingly flexible body-P content in fishes. This result is consistent with fish using flexible body-P content (presumably mediated through bone P) to buffer imbalances between P intake and demand. Together, our results provide evidence for a relatively ''dull'' stoichiometric ''knife-edge'' in fishes, driven in part by flexible body-P content.
Summary1. The evolution of sexual dimorphism presents a challenge because males and females must express two phenotypes from the same underlying genome. In vertebrates, one solution to this challenge is to link the expression of shared traits to sex steroids. However, even 'male-biased' steroids such as testosterone (T) circulate at biologically significant levels in females, raising the question of whether sexual dimorphism evolves not only through the coupling of trait expression to T in males, but also through the decoupling of trait expression from T in females. 2. We tested these alternatives by asking whether male and female brown anoles (Anolis sagrei) respond to exogenous T in similar fashion with respect to a suite of sexually dimorphic traits: growth, skeletal morphology, resting metabolism, fat storage, dewlap size and dewlap colour. 3. First, we established the ontogeny of sexual dimorphism in a colony raised in a laboratory common garden. Next, we treated juveniles of each sex with either T implants or empty implants at 5-8 months of age, when sexual dimorphism first began to develop for most traits. 4. T stimulated growth in both sexes and largely abolished natural sex differences in growth. This effect was associated with the stimulation of resting metabolism and the diversion of energy from fat and liver stores in both sexes. T also enlarged the dewlap in both sexes, though females never developed dewlaps equal in size to those of males. Finally, T altered the brightness and saturation of the dewlap in both sexes, inducing coloration similar to that of adult males. 5. Female brown anoles retain many of the same tissue-specific responses to T that occur in males, suggesting that the evolution of androgen-mediated sexual dimorphism has been achieved largely through the coupling of trait expression to sex differences in circulating T, without an associated decoupling of trait expression from T in females.
The mangrove rivulus (Kryptolebias marmoratus) is a small fish native to mangrove ecosystems in Florida, the Caribbean, Central America, and South America. This species is one of only two self-fertilizing, hermaphroditic vertebrates capable of producing offspring that are genetically identical to both the parent and all siblings. Long bouts of selfing result in individuals with completely homozygous genotypes, effectively allowing for the production of "clones." Rivulus is also extremely sensitive to environmental change, both during development and adulthood. Life-history traits, behavior, physiology, morphology, and even sexual phenotype are shaped to a large extent by the interaction of genes with the environment, and many of these traits appear to co-vary. True reaction norms can be generated for this species in much the same way as has been done for clonally reproducing invertebrates and plants that have contributed immensely to our understanding of the evolution of phenotypic plasticity. That is, rivulus provides the opportunity to place individuals with identical genotypes in many different environments at any point during ontogeny or adulthood. In addition, rivulus populations are characterized by high genotypic diversity, a luxury not afforded by many clonal vertebrates, which allows us to evaluate variation among genotypes in the shape of reaction norms and in patterns of covariance among traits. We provide background information on phenotypic plasticity and phenotypic integration, coupled with a description of characteristics that we feel qualify rivulus as a potentially powerful model in which to study the evolution of reaction norms and covariance among traits.
Using the Xiphophorus fish melanoma model we show a strong male bias for cutaneous malignant melanoma, consistent with that seen in the human population. To examine underlying factors, we exposed adult X. couchianus fish to a single, sub-lethal dose of UVB and measured circulating sex steroid hormones and expression of associated hormone receptor genes over a 24 hour period. We found that a single exposure had profound effects on circulating levels of steroid hormones with significant decreases for all free sex steroids at 6 and 24 h and increases in conjugated 2-estradiol and 11-ketotestosterone at 6 and 24 h, respectively. Whereas ARα expression increased in male and female skin, neither ARβ nor either of the ER’s showed significant responses to UVB in either sex. The rapid response of male androgens and their receptors in the skin after UVB irradiation implicates hormones in the male-bias of skin cancer and suggests that the photoendocrine response immediately after UV exposure may be relevant to melanomagenesis.
Usually no distinction is made between female and male salivary glands although cyclic changes of and ⁄ or differences in serum and salivary sex steroid concentrations characterize women and men. Moreover, sexual dimorphism is well recognized in salivary glands of rodents. Salivary glands contain estrogen and androgen receptors and are, according to modern high throughput technologies, subjected to gender differences not explainable by gene dose effects by the X chromosome alone. Because sex steroids are lipophilic, it is often thought that approximately 10% of them passively diffuse from plasma to saliva. Indeed, saliva can find use as sample material in sports medicine, pediatrics, veterinary medicine and behavioral sciences. Last but not least, humans and other primates are unique in that they have a reticular zone in their adrenal cortex, which produces dehydroepiandrosterone and androstendione pro-hormones. These are processed in peripheral tissues, not only in female breast and uterus and male prostate, but also in salivary glands by an intracrine enzymatic machinery to active 17b-estradiol, dihydrotestosterone and others, to satisfy and buffer against a constantly changing needs caused by circadian, menstrual, pregnancy and chronobiological hormonal changes in the systemic circulation. Female dominance of Sjö gren's syndrome and certain forms of salivary gland cancer probably reflect these gender-based differences.Oral Diseases (2010) 16, 577-585
This study validated a technique for non-invasive hormone measurements in California killifish Fundulus parvipinnis, and looked for associations between cortisol (a stress hormone) and 11-ketotestosterone (KT, an androgen) release rates and the density or intensity of the trematode parasites Euhaplorchis californiensis (EUHA) and Renicola buchanani (RENB) in wild-caught, naturally infected F. parvipinnis. In experiment 1, F. parvipinnis were exposed to an acute stressor by lowering water levels to dorsal-fin height and repeatedly handling the fish over the course of an hour. Neither parasite was found to influence cortisol release rates in response to this acute stressor. In experiment 2, different F. parvipinnis were exposed on four consecutive days to the procedure for collecting water-borne hormone levels and release rates of 11-KT and cortisol were quantified. This design examined whether F. parvipinnis perceived the water-borne collection procedure to be a stressor, while also exploring how parasites influenced hormone release rates under conditions less stressful than those in experiment 1. No association was found between RENB and hormone release rates, or between EUHA and 11-KT release rates. The interaction between EUHA density and handling time, however, was an important predictor of cortisol release rates. The relationship between handling time and cortisol release rates was negative for F. parvipinnis harbouring low or intermediate density infections, and became positive for fish harbouring high densities of EUHA.
Phenotypic flexibility is essential for organisms to respond to changing environments. Guppies experience light environments that alter their visibility to conspecifics and predators. We used guppies from populations with low rates of predation by fish, but which may be subject to avian predators, to test the hypotheses that conspicuous behaviors and the androgens that mediate them are reduced under high light, and that cortisol levels are increased under high light because the perceived risk is stressful. We found reduced courtship, potentially driven by the reduced female response to courtship, under high light. Aggression and testosterone levels were higher in the absence of females. We found elevated androgen and decreased cortisol levels following social interactions, but no relationship between hormones and behavior, and no influence of light level on hormones. We forward explanations for these results and advocate understanding the flexible response to light environments in a range of guppy populations.
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