Wild at heart? : differential maternal investment in wild and domesticated zebra finches (Taeniopygia guttata)
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Over the past twenty years there has been an exponential increase in the investigation of maternal effects. Understanding the adaptive function of maternal allocation strategies is integral to interpreting the evolutionary outcomes of sexual selection. Thus, model animal systems that facilitate experimental manipulation and controlled investigation of the physiological and behavioural mechanisms underlying maternal effects are important to evolutionary biologists. The zebra finch (Taeniopygia guttata) has been used as a model to investigate avian life-history, signalling behaviour, neurophysiology, mate choice, and more recently, maternal effects. However, a potentially influential and rarely addressed problem with this species is the process of domestication. Within this thesis we aimed to both test current predominant maternal allocation hypotheses, but for the first time in both domesticated and wild zebra finches. Chapter 2 develops on earlier work using domesticated zebra finches that has demonstrated differential allocation of maternally derived yolk androgens and antioxidants in eggs dependent on paternal attractiveness. This chapter specifically tests the ratio of these two yolk resources within individual eggs and shows that the balance of androgens to antioxidants varies by offspring sex and paternal attractiveness. Specifically, we found that mothers allocated a smaller androgen to antioxidant ratio to daughters when paired to green ringed (unattractive) males compared to red ringed (attractive) males. This pattern was reversed for sons, where mothers allocated a larger ratio of androgen to antioxidant when paired to red ringed (attractive) compared to green ringed (unattractive) males. We also show that brood sex ratio depended on both female condition and male attractiveness. It is concluded that investigating female allocation of individual resources within egg yolks may lead to incorrect assumptions on offspring fitness consequences, and that individual female state is an important consideration when predicting a resource allocation strategy. Throughout this thesis colour bands are used as a method to manipulate male attractiveness. In chapter 3 the influence of these bands was further tested to elucidate whether they affect male behaviour or quality. Wild birds were used for this chapter as preferences for bands based on colour have only once been demonstrated in wild birds and it was felt this should also be replicated. We confirmed a female preference for males based on colour bands worn in mate choice trials, with red bands preferred over green. Interestingly, we also found that colour of bands worn by males for an extended period in the single sex aviary influenced both their song rate and condition. Males that had worn red bands sang more in mate choice trials than both green banded or un-banded males. In addition red banded males were found to be in significantly better physical condition. These data suggest that earlier experiments in which it has been assumed that colour bands do not manipulate any form of intrinsic male quality should be re-evaluated. The final two data chapters, 4 and 5, return to investigating maternal allocation in response to male attractiveness, but for the first time in wild birds. Chapter 4 presents an experiment that was conducted on a wild, nest box breeding population of birds. Maternal resources allocation was investigated in both an experimental manipulation of male attractiveness, and also by correlating resource allocation with paternal phenotypic traits. A limited sample size meant few conclusions could be drawn from the experimental study, but significant positive correlations were found between both egg size and yolk testosterone (T) concentration and male phenotypic traits. This suggested that wild zebra finches may follow a positive investment strategy but requires further investigation. In chapter 5 experiments were repeated on wild birds that had been brought into captivity, to allow both an improved sample size and further control of influential environmental features. Again, female allocation strategies are tested using colour bands to manipulate male attractiveness, to allow direct comparisons with work on domesticated zebra finches. We found that females laid significantly heavier eggs for attractive compared to unattractive males, supporting the positive investment hypothesis. In addition we found an interaction between offspring size and paternal attractiveness treatment, with daughters of red banded (attractive) males being smaller than sons. This experiment is the first to demonstrate the influence of colour bands on maternal allocation in wild zebra finches and also provides further support for the positive investment hypothesis in this species. The final chapter discusses how overall patterns of female allocation were shown to be similar among wild and domesticated populations. It is concluded that demonstrated variations between populations and/or contexts reported in these studies cannot be explained by inherent differences between wild and domesticated individuals. Thus, the zebra finch remains a robust and reliable model for testing the evolution of avian maternal allocation strategies.
Thesis, PhD Doctor of Philosophy
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