Effects of the antiandrogens cyproterone acetate and flutamide on male reproductive behavior in a lizard (Anolis sagrei)

This study examined the effects of the antiandrogens cyproterone acetate (CA) and flutamide (F) on male reproductive behavior in the lizard Anolis sagrei. Reproductively active males were implanted with subcutaneous pellets of either CA, F, or placebo (P). Pellets delivered CA and F at a constant rate of 0.1 mg/day. Three weeks after implantation, males were tested with stimulus males and two days later with stimulus females. Cyproterone acetate inhibited aspects of male aggressive and sexual behavior, and reduced testis weight and size of the renal sex segment. Plasma testosterone (T) levels in CA-treated males were not significantly different than those of P-treated males. Flutamide did not inhibit aggressive or sexual behavior, but did decrease testis weight as well as the size of the renal sex segment. Plasma T levels were significantly higher in F-treated males than in P-treated males. These data suggest that CA, an antiandrogen with antigonadotropic activity, may be used to inhibit reproductive behavior in male lizards.     

Effects of corticosterone treatment on male aggressive behavior in a lizard (Anolis sagrei)

Social dominance in several species of lizards appears related to adrenal gland activity. The purpose of this study was to test whether the adrenal hormone corticosterone (CS) would inhibit the aggressive behavior of males of the lizard Anolis sagrei. Reproductively active males were randomly assigned to either CS or placebo (P) treatment groups or to an untreated stimulus male group. Both CS and P were administered in pellet form. Corticosterone pellets were designed to release approximately 4.8 micrograms of CS/day. Initially (Day 1), treatment males were implanted subcutaneously with a single pellet of either CS or P. After 1 week (Day 8), behavior tests were staged between treatment males and untreated stimulus males. The behavior of treatment males in response to stimulus males was videotaped. Significantly fewer CS-treated males than P-treated males approached the stimulus males, erected a crest, or showed biting behavior. Corticosterone-treated males also displayed significantly less frequently than did P-treated males and gave fewer displays of a type associated with dominance. Mean combined testis weight at the end of the experiment was significantly less in CS-treated males than in P-treated males and spermatogenic activity in CS-treated males was reduced. Plasma testosterone in CS-treated males was nondetectable, whereas P-treated males had normal levels of this steroid. These results indicate that CS can inhibit male aggressive behavior and testicular function in a lizard and support the hypothesis that the behavior changes observed in subordinate animals following defeat in agonistic encounters may in part be due to increased plasma CS levels.     

Copulatory behaviour of the lizard Anolis sagrei: alternation of hemipenis use

The pattern of use of the hemipenes, the two independently functioning intromittent organs of squamate reptiles, was investigated in the male brown anole, Anolis sagrei. Males demonstrated a significant tendency to use the left and right hemipenes alternately when mated daily for 7 days with different females. The duration of copulation differed significantly among males and was independent of the hemipenis used. Hemipenis use was also examined in two successive copulations separated by different time intervals. After an intercopulatory interval of up to 3 h, males showed a significant tendency to alternate in the use of their hemipenes. This trend was still present following an intercopulatory interval of 24–26 h, but was not detectable following an interval of 72–74 h. Thus, alternation of hemipenis use in male brown anoles declines as the interval between copulations increases, suggesting that males adjust their copulatory posture in relation to the length of the intercopulatory interval.     

The role of the anterior hypothalamus-preoptic area in the regulation of male reproductive behavior in the lizard, Anolis carolinensis: lesion studies.

Bilateral radiofrequency lesions of the anterior hypothalamic-preoptic area (AH-POA) abolish male Courtship and Agonistic behaviors in both intact and castrated/androgen-treated Anolis carolinensis. Control lesions caudal and dorsal to the AH-POA have no effect on these behaviors. Lesions rostral to the AH-POA also cause a significant decline in the behavior of intact animals. Histological examination of testes showed that lesions including and rostral to the AH-POA cause testicular collapse and regression to initial stages of spermatogenesis, indicating interruption of gonadotropin secretion. Castration and subcutaneous implantation of androgen restores Courtship and Agonistic behaviors in Rostral and Caudal Control groups to prelesion levels, while AH-POA lesioned animals remain at their low behavioral levels. Neither AH-POA or control lesions cause significant changes in animals' body weight.     

Evolutionary implications of phenotypic plasticity in the hindlimb of the lizard Anolis sagrei

Abstract.— Species of Anolis lizards that use broad substrates have long legs, which provide enhanced maximal sprint speed, whereas species that use narrow surfaces have short legs, which permit careful movements. We raised hatchling A. sagrei in terraria provided with only broad or only narrow surfaces. At the end of the experiment, lizards in the broad treatment had relatively longer hindlimbs than lizards in the narrow treatment. These results indicate that not only is hindlimb length a plastic trait in these lizards, but that this plasticity leads to the production of phenotypes appropriate to particular environments. Comparison to hindlimb lengths of other Anolis species indicates that the range of plasticity is limited compared to the diversity shown throughout the anole radiation. Nonetheless, this plasticity potentially could have played an important role in the early stages of the Caribbean anole radiation.     

The role of the basal hypothalamus in the regulation of reproductive behavior in the lizard, Anolis carolinensis: lesion studies

Bilateral radiofrequency lesions in the anterior and posterior basal hypothalamus decreased courtship and agonistic behaviors in both intact, sexually active, and castrated, androgen treated male Anolis carolinensis. Intact males receiving lesions in the anterior basal hypothalamus had atrophied testes, aspermia, and decreased epithelial cell height of the renal sex segment. Lesions of the posterior basal hypothalamus had no effect on testicular activity or the development of accessory organs. All animals demonstrating behavioral changes had lesion destruction in the ventromedial nucleus and the accompanying periventricular system. It is concluded that the basal hypothalamus in male A. carolinensis is involved both in the regulation of reproductive behavior and pituitary function.     

Differential admixture shapes morphological variation among invasive populations of the lizard Anolis sagrei

The biological invasion of the lizard Anolis sagrei provides an opportunity to study evolutionary mechanisms that produce morphological differentiation among non-native populations. Because the A. sagrei invasion represents multiple native-range source populations, differential admixture as well as random genetic drift and natural selection, could shape morphological evolution during the invasion. Mitochondrial DNA (mtDNA) analyses reveal seven distinct native-range source populations for 10 introduced A. sagrei populations from Florida, Louisiana and Texas (USA), and Grand Cayman, with 2-5 native-range sources contributing to each non-native population. These introduced populations differ significantly in frequencies of haplotypes from different native-range sources and in body size, toepad-lamella number, and body shape. Variation among introduced populations for both lamella number and body shape is explained by differential admixture of various source populations; mean morphological values of introduced populations are correlated with the relative genetic contributions from different native-range source populations. The number of source populations contributing to an introduced population correlates with body size, which appears independent of the relative contributions of particular source populations. Thus, differential admixture of various native-range source populations explains morphological differences among introduced A. sagrei populations. Morphological differentiation among populations is compatible with the hypothesis of selective neutrality, although we are unable to test the hypothesis of interdemic selection among introductions from different native-range source populations.     

Effects of ATD on male sexual behavior and androgen receptor binding: a reexamination of the aromatization hypothesis

The aromatization hypothesis asserts that testosterone (T) must be aromatized to estradiol (E2) to activate copulatory behavior in the male rat. In support of this hypothesis, the aromatization inhibitor, ATD, has been found to suppress male sexual behavior in T-treated rats. In our experiment, we first replicated this finding by peripherally injecting ATD (15 mg/day) or propylene glycol into T-treated (two 10-mm Silastic capsules) or control castrated male rats. In a second experiment, we bilaterally implanted either ATD-filled or blank cannulae into the medial preoptic area (MPOA) of either T-treated or control castrated male rats. With this more local distribution of ATD, a lesser decline in sexual behavior was found, suggesting that other brain areas are involved in the neurohormonal activation of copulatory behavior in the male rat. To determine whether in vivo ATD interacts with androgen or estrogen receptors, we conducted cell nuclear androgen and estrogen receptor binding assays of hypothalamus, preoptic area, amygdala, and septum following treatment with the combinations of systemic T alone. ATD plus T, ATD alone, and blank control. In all four brain areas binding of T to androgen receptors was significantly decreased in the presence of ATD, suggesting that ATD may act both as an androgen receptor blocker and as an aromatization inhibitor. Competitive binding studies indicated that ATD competes in vitro for cytosol androgen receptors, thus substantiating the in vivo antiandrogenic effects of ATD. Cell nuclear estrogen receptor binding was not significantly increased by exposure to T in the physiological range. No agonistic properties of ATD were observed either behaviorally or biochemically. Thus, an alternative explanation for the inhibitory effects of ATD on male sexual behavior is that ATD prevents T from binding to androgen receptors.     

The effects of an aromatization inhibitor on the reproductive behavior of male zebra finches

Recent evidence indicates that aromatizable androgens are more effective than nonaromatizable androgens in restoring normal levels of sexual behavior in castrated male zebra finches (Poephila guttata). To determine whether the efficacy of treatment with aromatizable androgens, is in part due to their conversion to estrogens, castrated male finches were treated with androstenedione (AE), an aromatizable androgen, and their sexual and aggressive behavior was compared with that of castrates treated with AE plus 1,4,6-androstatriene-3,17-dione (ATD), an aromatization inhibitor. Males treated with AE + ATD showed less courtship activity and less copulatory behavior than AE-treated males, and were unlikely to have nests. Estradiol (E), when given concurrently with AE + ATD, reversed the inhibitory effects of ATD and restored levels of courtship and copulation to those observed in AE-treated males. Only AE- and AE + ATD + E-treated males displayed aggressive behaviors, but the frequency of such behaviors was so low that there were no significant differences across groups. These data affirm the importance of estrogen in the control of reproductive activities in male zebra finches and indicate that aromatization may be an obligatory step for maintaining normal levels of sexual and aggressive behavior.     

Climate‐related environmental variation in a visual signalling device: the male and female dewlap in Anolis sagrei lizards

Animals communicate using a variety of signals that differ dramatically among and within species. The astonishing dewlap diversity in anoles has attracted considerable attention in this respect. Yet, the evolutionary processes behind it remain elusive and have mostly been explored for males only. Here, we considered Anolis sagrei males and females to study signal divergence among populations. First, we assessed the degree of variation in dewlap design (size, pattern and colour) and displays by comparing 17 populations distributed across the Caribbean. Second, we assessed whether the observed dewlap diversity is associated with variation in climate‐related environmental conditions. Results showed that populations differed in all dewlap characteristics, with the exception of display rate in females. We further found that males and females occurring in ‘xeric’ environments had a higher proportion of solid dewlaps with higher UV reflectance. In addition, lizards inhabiting ‘mesic’ environments had primarily marginal dewlaps showing high reflectance in red. For dewlap display, a correlation with environment was only observed in males. Our study provides evidence for a strong relationship between signal design and prevailing environmental conditions, which may result from differential selection on signal efficacy. Moreover, our study highlights the importance of including females when studying dewlaps in an evolutionary context.     

Effects of testosterone treatment and season on the frequency of dewlap extensions during male-male interactions in the lizard Anolis sagrei

Lizards of the genus Anolis extend and retract a large and often brightly colored throat fan called a dewlap. The dewlap in most anoles is a sexually dimorphic structure. It is larger in males than females and males use the dewlap more frequently and in more contexts than do females. In the present study we investigated whether plasma testosterone (T) levels and season affect the frequency of dewlap use in male--male interactions in the brown anole, Anolis sagrei. We manipulated plasma T levels by implanting adult, castrated males with pellets that delivered no T, a moderate dose of T, or a high dose of T. In tests with stimulus males, castrated males that were treated with a moderate or with a high dose of T had a significantly higher frequency of dewlap extensions than did castrated males that received no T. However, the frequency of dewlap extensions in castrated males that received the high dose of T did not differ significantly from castrated males that received the moderate dose of T or from non-castrated control males. Males captured during the breeding season and tested in the laboratory had a significantly higher frequency of dewlap extensions than did males captured in the postbreeding season and tested in the same manner. These results suggest that plasma T levels affect how frequently males of A. sagrei extend their dewlaps in male--male interactions and that seasonal changes in male dewlap use may be due to seasonal differences in plasma T levels.     

Enzyme function and polymorphism: A test in two Anolis lizard species

Electrophoretic surveys of two lizard species were used to test hypotheses that relate levels of enzyme variability to enzyme function (single-substrate versus multiple-substrate enzymes, regulatory versus nonregulatory enzymes). Anolis roquet behaviorally regulates its body temperature, but its congener A. gundlachi is passive to variations in the thermal environment. As a result, populations of A. gundlachi probably experience the thermal environment as temporally coarse-grained, whereas populations of A. roquet do not. We therefore predicted that A. gundlachi would exhibit greater enzyme heterozygosity than A. roquet and that different enzyme classes would contribute disproportionately to this interspecific difference. The data show (1) that A. gundlachi does have a higher heterozygosity and (2) that this difference appears to result from high levels of heterozygosity at loci coding for multiple-substrate enzymes. The dichotomy between regulatory and nonregulatory enzymes offers no explanation for the variability in heterozygosity among enzyme loci in these species.E.Z. was supported by a grant from the Natural Sciences and Engineering Council of Canada. The study was accomplished while P.E.H. held a postdoctoral fellowship from the Killam Trust of Dalhousie University and a grant from the Research Development Fund of Dalhousie University. The collection of material was made possible by grants (to P.E.H.) from National Science Foundation (DEB 75-16334), the Explorers Club of New York, Sigma Xi, and the Richmond and Anderson Funds of Harvard University. We thank Dr. D. W. Foltz for his help with the calculations.     

Parasites and the evolution of extravagant male characters: Anolis lizards on Caribbean islands as a test of the Hamilton-Zuk hypothesis

Anolis lizards from Puerto Rico (five species from one site), Curaçao and Aruba in the southern Caribbean (2 populations), and 22 populations from 14 islands in the eastern Caribbean were surveyed for blood parasites (two species of Plasmodium and haemogregarines). Literature records for gut helminths from nine of these populations were added to the data set. Dorsal body color and dewlap color of males were also observed and classified into objective classes with no subjective view of showiness. These data were used to test the among-species prediction of the Hamilton-Zuk hypothesis which states that species harboring more harmful parasites over their evolutionary history will be more likely to evolve extravagant sexually dimorphic traits. Critics have noted important shortcomings in previous tests of the prediction; here we corrected for these errors. Parasite loads (prevalence and number of species) and dorsal and dewlap color varied substantially among the populations sampled. However, there was no association of parasite load with color either in a broad analysis or when correcting for phylogenetic relationships among the lizard species.     

A circadian clock measures photoperiodic time in the male lizard Anolis carolinensis

Summary Photoperiod plays an important role in controlling the annual reproductive cycle of the male lizard Anolis carolinensis. The nature of photoperiodic time measurement in Anolis was investigated by exposing anoles to 3 different kinds of lighting paradigms (resonance, T cycles, and night breaks) to determine if photoperiodic time measurement involves the circadian system. Both the reproductive response and the patterns of entrainment of the activity rhythm were assessed. The results show that the circadian system is involved in photoperiodic time measurement in this species and that a discrete photoinducible phase resides in the latter half of the animals' subjective night. Significantly, the ability of the circadian system to execute photoperiodic time measurement is crucially dependent on the length of the photoperiod. Resonance, T cycle and night break cycles utilizing a photoperiod 10–11 h in duration reveal circadian involvement whereas these same cycles utilizing 6 or 8 h photoperiods do not.Abbreviation CRPP circadian rhythm of photoperiodic sensitivity     

Predation by an exotic lizard,Anolis sagrei,alters the ant community structure in betelnut palm plantations in southern Taiwan

Abstract 1. Predators can affect prey directly by reducing prey abundance and indirectly by altering behavioural patterns of prey. From previous studies, there is little evidence that ant community structure is affected by vertebrate predation. 2. Researchers tend to consider the interactions between vertebrate predators and ants to be weak. The present study examined the impact of the exotic invasive lizard, Anolis sagrei, on the ant community structure by manipulating the density of lizards within enclosures. The natural density of A. sagrei in the field was surveyed and used as the stocking density rate in the lizard‐present sub‐enclosures. 3. Before the lizard density was manipulated, there was no difference in the ant diversity between sub‐enclosures. After the lizard density manipulation, the ant diversity in sub‐enclosures with A. sagrei present was significantly different from that of enclosures where the lizards were absent, although the overall ant abundance did not differ significantly. 4. The ant diversity difference was generated by a significant reduction of the ant species Pheidole fervens in sub‐enclosures with A. sagrei present. Such an abundance change might be the result of direct predation by the lizards, or it might be generated by a foraging site shift by this ant. 5. The results of this study thus demonstrated that the invasion of an exotic vertebrate can significantly alter the community structure of ants, perhaps through the combined direct and indirect effects of lizards on ants.