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1.  Changes in Plasma Testosterone Levels and Brain AVT Cell Number during the Breeding Season in the Green Treefrog 
Brain, Behavior and Evolution  2010;75(4):271-281.
We exposed groups of adult male green treefrogs, Hyla cinerea, to acoustic stimuli (natural chorus or random tones) for seven consecutive nights at three time points during their natural breeding season (May, July, and September) and assessed seasonal changes in plasma androgen levels and number of arginine vasotocin (AVT) immunoreactive cells in the brain over this time period. We also tested whether social cues altered either androgens or AVT-ir cell number or size at each time point. Finally, we analyzed how these factors related to calling behavior. Data were collected over two breeding seasons. Call rate (calls/h) was assessed during the stimulus time (i.e. ‘evoked calling’) and during the remainder of the day (‘spontaneous calling’). Plasma hormone levels were measured at the end of the acoustic treatment when brains were collected for immunocytochemistry. Circulating androgen levels declined over the breeding season. Males exposed to chorus sounds, however, had higher androgen levels than males exposed to tones. AVT-ir cell number increased across the breeding season in the nucleus accumbens but not the amygdala, anterior preoptic area, or magnocellular preoptic area, and soma size decreased in the nucleus accumbens as cell number increased. Social stimulation had no significant influence on either AVT-ir cell measure. Evoked call rate was higher in males exposed to natural chorus sounds compared to those exposed to random tones, but did not change during the season. In contrast, spontaneous call rate was higher at the beginning of the breeding season compared to the end, and unlike evoked calling was correlated with circulating androgen levels across all treatments and time points. AVT-ir soma size was positively correlated with both evoked and spontaneous calling. These results suggest that social exposure can prolong the elevation of gonadal hormones in the bloodstream, thus mitigating or slowing the seasonal decline of such hormones. In contrast, social exposure does not affect the seasonal pattern of AVT-ir cell number or soma size. The reciprocal relationship between social cues and hormones and the subsequent effect on behavior may provide hidden benefits to animals engaging in social interactions. However, unlike steroid hormone levels, the seasonal change in AVT-ir cell number and size is not counteracted by social stimulation.
doi:10.1159/000316084
PMCID: PMC3202922  PMID: 20664184
Amphibian; Androgens; Arginine vasotocin; Communication; Seasonality
2.  Current research in amphibians: Studies integrating endocrinology, behavior, and neurobiology 
Hormones and behavior  2005;48(4):440-450.
Amphibian behavioral endocrinology has focused on reproductive social behavior and communication in frogs and newts. Androgens and estrogens are critical for the expression of male and female behavior, respectively, and their effects are relatively clear. Corticosteroids have significant modulatory effects on the behavior of both sexes, as does the peptide neuromodulator arginine vasotocin in males, but their effects and interactions with gonadal steroids are often complex and difficult to understand. Recent work has shown that the gonadal hormones and social behavior are mutually reinforcing: engaging in social interactions increases hormone levels just as increasing hormone levels change behavior. The reciprocal interactions of hormones and behavior, as well as the complex interactions among gonadal steroids, adrenal steroids, and peptide hormones have implications for the maintenance and evolution of natural social behavior, and suggest that a deeper understanding of both endocrine mechanisms and social behavior would arise from field studies or other approaches that combine behavioral endocrinology with behavioral ecology.
doi:10.1016/j.yhbeh.2005.06.001
PMCID: PMC2581512  PMID: 16023646
Frogs; Newts; Androgen; Corticosteroids; AVT; Communication; Reproduction

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