Heritability of corticosterone response and changes in life history traits during selection in the zebra finch

Vertebrates respond to environmental stressors through the neuro-endocrine stress response, which involves the production of glucocorticoids. We have selected independent, duplicate divergent lines of zebra finches for high, low and control corticosterone responses to a mild stressor. This experiment has shown that over the first four generations, the high lines have demonstrated a significant realized heritability of about 20%. However, the low lines have apparently not changed significantly from controls. This asymmetry in response is potentially because of the fact that all birds appear to be showing increased adaptation to the environment in which they are housed, with significant declines in corticosterone response in control lines as well as low lines. Despite the existence of two- to threefold difference in mean corticosterone titre between high and low lines, there were no observed differences in testosterone titre in adult male birds from the different groups. In addition, there were no consistent, significant differences between the lines in any of the life history variables measured – number of eggs laid per clutch, number of clutches or broods produced per pair, number of fledglings produced per breeding attempt, nor in any of egg, nestling and fledgling mortality. These results highlight the fact that the mechanisms that underlie variation in the avian physiological system can be modified to respond to differences between environments through selection. This adds an additional level of flexibility to the avian physiological system, which will allow it to respond to environmental circumstances.

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.

Developmental stress and female mate choice behaviour in the zebra finch

The potential effects of early environmental conditions on adult female mate choice have been largely neglected in studies of sexual selection. Our study tested whether developmental stress affects the mate choice behaviour of female zebra finches, Taeniopygia guttata, when choosing between potential mates. In an experiment manipulating developmental condition, female zebra finches were raised under nutritional stress or control conditions. In adulthood, female preferences were assessed using extensive four-stimulus mate choice trials. Nutritional stress affected growth rates during the period of stress, with experimentally stressed females lighter than controls. During mate choice trials stressed females were almost three times less active than controls and made fewer sampling visits to the stimulus males, although we found no evidence of a direct effect of developmental experience on which males were preferred. Thus, developmental experience had a clear effect on behavioural patterns in a mate choice context. To test whether this effect is specific to a mate choice context, we also investigated the effect of developmental stress on female activity rates in three social contexts: isolation, contact with a conspecific male (a potential mate) and contact with a conspecific female. Here, female activity did not differ between the experimental treatments in any of the social situations. Overall, our findings suggest that environmental conditions during early development can have long-term context-dependent consequences for adult female mate choice behaviour, mediated by changes in activity rates.

The colour of paternity: extra-pair paternity in the wild Gouldian finch does not appear to be driven by genetic incompatibility between morphs

In socially monogamous species, individuals can use extra-pair paternity and offspring sex allocation as adaptive strategies to ameliorate costs of genetic incompatibility with their partner. Previous studies on domesticated Gouldian finches (Erythrura gouldiae) demonstrated a genetic incompatibility between head colour morphs, the effects of which are more severe in female offspring. Domesticated females use differential sex allocation, and extra-pair paternity with males of compatible head colour, to reduce fitness costs associated with incompatibility in mixed-morph pairings. However, laboratory studies are an oversimplification of the complex ecological factors experienced in the wild, and may only reflect the biology of a domesticated species. This study aimed to examine the patterns of parentage and sex-ratio bias with respect to colour pairing combinations in a wild population of the Gouldian finch. We utilized a novel PCR assay that allowed us to genotype the morph of offspring before the morph phenotype develops, and to explore bias in morph paternity and selection at the nest. Contrary to previous findings in the laboratory, we found no effect of pairing combinations on patterns of extra-pair paternity, offspring sex ratio, or selection on morphs in nestlings. In the wild, the effect of morph incompatibility is likely much smaller, or absent, than was observed in the domesticated birds. Furthermore, the previously studied domesticated population is genetically differentiated from the wild population, consistent with the effects of domestication. It is possible that the domestication process fostered the emergence (or enhancement) of incompatibility between colour morphs previously demonstrated in the laboratory.

Short- and long-term consequences of early developmental conditions : a case study on wild and domesticated zebra finches

Divergent selection pressures among populations can result not only in significant differentiation in morphology, physiology and behaviour, but also in how these traits are related to each other, thereby driving the processes of local adaptation and speciation. In the Australian zebra finch, we investigated whether domesticated stock, bred in captivity over tens of generations, differ in their response to a life-history manipulation, compared to birds taken directly from the wild. In a ‘common aviary’ experiment, we thereto experimentally manipulated the environmental conditions experienced by nestlings early in life by means of a brood size manipulation, and subsequently assessed its short- and long-term consequences on growth, ornamentation, immune function and reproduction. As expected, we found that early environmental conditions had a marked effect on both short- and long-term morphological and life-history traits in all birds. However, although there were pronounced differences between wild and domesticated birds with respect to the absolute expression of many of these traits, which are indicative of the different selection pressures wild and domesticated birds were exposed to in the recent past, manipulated rearing conditions affected morphology and ornamentation of wild and domesticated finches in a very similar way. This suggests that despite significant differentiation between wild and domesticated birds, selection has not altered the relationships among traits. Thus, life-history strategies and investment trade-offs may be relatively stable and not easily altered by selection. This is a reassuring finding in the light of the widespread use of domesticated birds in studies of life-history evolution and sexual selection, and suggests that adaptive explanations may be legitimate when referring to captive bird studies.

Artificial ornaments manipulate intrinsic male quality in wild-caught zebra finches (Taeniopygia guttata)

The addition of red and green color bands is a commonly used method for manipulating male attractiveness in the zebra finch (Taeniopygia guttata), providing insight into the study of maternal investment and sexual selection. The addition of artificial ornaments has been assumed to manipulate a females’ perception of the male, rather than affecting intrinsic qualities of the male himself. Here, however, we reveal that the artificial band color worn by a male changes his body mass, condition, and courtship display rate. Males wearing red color bands in aviaries prior to mate-choice trials had a significantly higher song rate during trials than those wearing green color bands, alongside a significant increase in mass change and condition. Male song rate was found to significantly correlate with female preference alongside a female preference for red-banded males. However, male song rate in turn increased when female response was positive, suggesting a social feedback between the interacting birds. Our data suggest the presence of socially mediated feedback mechanisms whereby the artificial increase in attractiveness or dominance of a male directly affects other aspects of his attractiveness. Therefore, housing birds in social groups while manipulating attractiveness can directly influence other male qualities and should be accounted for by future studies.

Sex steroid profiles and pair-maintenance behavior of captive wild-caught zebra finches (taeniopygia guttata)

Here, we studied the life-long monogamous zebra finch, to examine the relationship between circulating sex steroid profiles and pair-maintenance behavior in pairs of wild-caught zebra finches (paired in the laboratory for >1 month). We used liquid chromatography-tandem mass spectrometry to examine a total of eight androgens and progestins [pregnenolone, progesterone, dehydroepiandrosterone (DHEA), androstenediol, pregnan-3,17-diol-20-one, androsterone, androstanediol, and testosterone]. In the plasma, only pregnenolone, progesterone, DHEA, and testosterone were above the limit of quantification. Sex steroid profiles were similar between males and females, with only circulating progesterone levels significantly different between the sexes (female > male). Circulating pregnenolone levels were high in both sexes, suggesting that pregnenolone might serve as a circulating prohormone for local steroid synthesis in zebra finches. Furthermore, circulating testosterone levels were extremely low in both sexes. Additionally, we found no correlations between circulating steroid levels and pair-maintenance behavior. Taken together, our data raise several interesting questions about the neuroendocrinology of zebra finches.

Using an Electronic Monitoring System to Link Offspring Provisioning and Foraging Behavior of a Wild Passerine

Although the costs of parental care are at the foundations of optimal-parental-investment theory, our understanding of the nature of the underlying costs is limited by the difficulty of measuring variation in foraging effort. We simultaneously measured parental provisioning and foraging behavior in a free-living population of Zebra Finches (Taeniopygia guttata) using an electronic monitoring system. We fitted 145 adults with a passive transponder tag and remotely recorded their visits to nest boxes and feeders continuously over a 2-month period. After validating the accuracy of this monitoring system, we studied how provisioning and foraging activities varied through time (day and breeding cycle) and influenced the benefits (food received by the offspring) and costs (interclutch interval) of parental care. The provisioning rates of wild Zebra Finches were surprisingly low, with an average of only one visit per hour throughout the day. This was significantly lower than those reported for this model species in captivity and for most other passerines in the wild. Nest visitation rate only partially explained the amount of food received by the young, with parental foraging activity, including the minimum distance covered on foraging trips, being better predictors. Parents that sustained higher foraging activity and covered more distance during the first breeding attempt took longer to renest. These results demonstrate that in some species matching foraging activity with offspring provisioning may provide a better estimate of the true investment that individuals commit to a reproductive attempt.