No abstract

Whereas several studies have shown that experimentally increased levels of the androgenic steroid testosterone can affect female behavior, fewer studies have focused on the activational effects of exogenous testosterone on female morphology. With respect to colorful displays in birds, almost exclusively the effects of testosterone manipulation on female carotenoid-based colorations have been studied. Other color types such as structural colors (i.e. UV, blue and violet colors that result from differential light reflection in the nanostructures of the tissue) remain largely unstudied. Here, we investigated the short- and long-term effects of exogenous testosterone on the expression of structural bare-part coloration in female budgerigars, Melopsittacus undulatus. In this parrot species, bare-part coloration is expressed in the cere, a structure over the beak which is brown in females and structural blue in males. We experimentally increased plasma testosterone levels in testosterone-treated females (T-females) compared to controls (C-females) and we performed weekly spectrophotometric measurements of the cere for five weeks after implantation and one measurement after ten weeks. We also estimated the extent to which testosterone masculinized female cere color by comparing the experimental females with untreated males. We found significant effects of testosterone on cere color from week four after implantation onwards. T-females expressed significantly bluer ceres than C-females with higher values for brightness and UV reflectance. T-female cere color, however, remained significantly less blue than in males, while values for brightness and UV reflectance were significantly higher in T-females than in males. Our quantitative results show that exogenous testosterone induces the expression of structural blue color in females but does not strongly masculinize female cere coloration. We provide several potential pathways for the action of testosterone on structural color.

An increasing number of studies indicate that not only females but also males can be selective when choosing a mate. In species exhibiting male or mutual mate choice, females may benefit from being attractive. While male attractiveness is often positively influenced by higher plasma levels of the androgenic hormone testosterone, it has been shown that testosterone can masculinise female behavior and morphology in several bird species, potentially rendering them less attractive. In this study, we investigated whether female budgerigars, Melopsittacus undulatus , suffer from increased plasma testosterone levels through a negative effect on their attractiveness to males. We experimentally increased plasma testosterone levels in testosterone-treated females (T-females) compared to controls (C-females) and allowed males to choose between a T- and a C-female in a two-way choice situation. Although testosterone treatment significantly affected female behavioral and morphological characteristics, males did not show a significant difference in preference between T- and C-females. These results suggest that experimentally increasing testosterone levels in females does not appear to influence male preference during initial mate choice. Our findings indicate that selection for higher levels of testosterone in male budgerigars is probably not constrained by a correlated response to selection causing negative effects on female attractiveness during initial mate choice. Evaluating whether or not a potential constraint may arise from negative testosterone-induced effects on other fitness related traits in females requires further work.

It is well established that in many avian species, prenatal maternal resource allocation varies both between and within clutches and may affect offspring fitness. Differential allocation of maternal resources, in terms of egg weight and yolk composition, may therefore allow the female to adjust brood reduction and to fine-tune reproductive investment in accordance with the expected fitness returns. The adaptive value of such maternal resource allocation is thought to be context-dependent as well as species-specific. We investigated the effects of female preference for her mate on the allocation of prenatal maternal resources in the budgerigar, Melopsittacus undulatus, a monogamous species of parrot that shows an extreme hatching asynchrony. We assessed mate preferences in a two-way preference test and allowed females two breeding rounds: one with the preferred and one with the non-preferred partner. We found no effect of preference on either latency to lay or clutch size, but females mated with the preferred partner laid eggs that contained significantly more yolk. Their eggs also contained significantly more androstenedione but not testosterone. Our results suggest that in this species, female preference may influence maternal resource allocation, and that the functional roles of each androgen in the yolk should be considered separately. In addition, we found a significant effect of laying order on egg and yolk weight as well as on yolk testosterone and androstenedione levels. These measures, however, did not change linearly with the laying order and render it unlikely that female budgerigars compensate for the extreme hatching asynchrony by adjusting within-clutch allocation of prenatal maternal resources.

Studies in several songbird species have shown that treating females with the androgenic steroid hormone testosterone (T) can negatively affect female reproductive behaviors and breeding success. As the effects of T on females appear to be species-specific, it is not clear if similar effects of high T occur in non-songbird species. Here, we studied the effects of T supplementation on female reproductive behavior and oviposition in the budgerigar, Melopsittacus undulatus, a small monogamous parrot species with distinct sex differences in parental behavior. We experimentally increased T concentrations to male-like levels in T-treated females compared to controls and we allowed females to breed. We found no significant effects of treatment on the latency to enter the nestbox but T treatment significantly interfered with oviposition. Our results show that T-treated females were seven times less likely to produce a clutch than control females. As we found that T treatment had a strong inhibitory effect on oviposition, our results indicate that female budgerigars suffer fitness costs from male-like plasma T levels. Therefore, it may be possible that, also in non-songbird species, selection for higher T levels in males is constrained by a correlated response to selection which imposes fitness costs on females in terms of reproduction. Evaluating whether or not this is indeed the case requires further work combining different approaches to the study of the evolution of male and female testosterone levels.