Temperature is arguably the most important abiotic factor influencing the embryonic development in ectothermic species. Incubation temperature has demonstrated effects on offspring phenotypes in ectotherms, including traits such as sex, size, shape, colouration and post-hatch growth and survival. However, in endotherms the influence temperature has on development is relatively unexplored due to the narrow range of temperatures that embryonic endotherms are exposed to during develop. Megapode birds utilize environmental heat sources to incubate their eggs and therefore provide a potential model to test how temperature influences embryonic development and offspring phenotypes in endotherms. I used the Australian Brush-turkey (Alectura lathami), a megapode bird that incubates its eggs in mounds of soil and leaf litter to investigate the effects of temperature on embryonic development and chick morphology. Previous reports of Australian Brush-turkey incubation mound temperatures relied on spot measurements and theoretical modeling and thus have not provided a comprehensive examination of the range of temperatures Brush-turkey embryos are exposed to and how these might vary during the course of incubation. Therefore to examine the range of temperatures experienced by developing embryos I continuously recorded the temperature of eggs and mound material at naturally occurring positions within incubation mounds over the full developmental period. As in previous studies I found the average incubation temperature to be about 34°C, however egg temperatures typically fluctuated more than previously reported or predicted from modeling. The thermal tolerance of Brush-turkeys is remarkable compared to non-megapode birds, with embryos developing successfully despite prolonged exposure to sub-optimal temperatures over the range 25-40°C. I also demonstrated that the incubation period was negatively correlated with mean incubation temperature. To simplify the examination of temperature effects on embryonic development, constant temperature artificial incubation of Brush-turkey eggs was used to determine influence of incubation temperature on the energetics of embryonic development and the sex ratio, morphology and chemical composition of chicks. Because initial investigation of mound temperatures determined the mean incubation temperature in Brush-turkeys to be 34°C this was used as the preferred temperature for constant temperature incubation with 32°C and 36°C representing low and high temperatures respectively. Previously, the sex ratio of Brush-turkey chicks at hatching was shown to be temperature dependent. A thermally sensitive period early in development resulted in more females hatching from high temperature and more males hatching from low temperatures with an equal ratio at the preferred temperature. Using molecular sexing techniques to determine the sex of both failed embryos and chicks that hatched, I established that at laying the sex ratio of eggs was 50:50, and that temperature-dependent sex-biased embryo mortality was the mechanism behind the skewed sex ratio of chicks hatching from non-preferred temperatures. Low incubation temperature increased female embryonic mortality and high incubation temperature increased male embryonic mortality. This represents a novel mechanism operating to alter sex ratios in a bird species and offers an unparalleled system to explore sex allocation theory. It is well established that temperature influences the rate of development and the morphology of offspring in reptilian species. Also, in a previous study using artificially incubated Brush-turkey eggs, temperature was found to affect the mass of chicks but not their size (linear dimensions). This finding suggests that at different incubation temperatures the amount of yolk converted into tissue during embryonic development is influenced by incubation temperature. I tested this hypothesis by incubating eggs at different constant temperatures and found high incubation temperatures produce chicks with lighter yolk-free bodies and heavier residual yolks but similar linear dimensions compared to chicks hatching from lower temperatures. Because eggs incubated at low temperatures have longer incubation periods, I hypothesized the proportion of lipid in the yolk-free body would be higher in chicks emerging from eggs incubated at low temperature because more time is available for the conversion of yolk to fat bodies during embryonic development. This hypothesis was not supported as the composition of yolk-free chicks (total water, lipid, protein and ash) was not temperature dependent. A previous study in Malleefowl (Leipoa ocellata), another megapode bird, found that the total energetic cost of production was influenced by incubation temperature. Such that embryos developing at low temperatures required 72% more energy than embryos developing at high temperatures. However these findings were contrary to expectation from studies of reptilian incubation where the energetic cost of development is independent of temperature. Therefore I tested the hypothesis that the total energetic cost of development is temperature dependent in the Australian Brush-turkey. I used bomb calorimetry to measure the energy content of freshly laid eggs and of chicks (both the yolk-free body and residual yolk) that had hatched from eggs incubated 32oC, 34oC and 36oC. I found that the total energy content of chicks at hatching was greater in chicks emerging from eggs incubated at 34oC and 36oC compared to eggs incubated at 32oC. My thesis work demonstrated that incubation temperature is more variable for Brush-turkey embryos than for non-megapode birds and that even a small difference in temperature can have important effects on chick sex ratios, morphology and energy reserves. I have shown that incubation under artificial constant temperature conditions can significantly alter the developmental trajectories and phenotypic outcomes for chicks. In addition to laboratory based work, future studies should continue to examine how embryonic development and chick attributes are influenced by temperatures experienced under natural incubation conditions. Furthermore, investigation is required to determine how incubation temperature induced differences in hatchling phenotypes influence the post-hatch grow and fitness of chicks.
| Identifer | oai:union.ndltd.org:ADTP/282233 |
| Creators | Yvonne Eiby |
| Source Sets | Australiasian Digital Theses Program |
| Detected Language | English |
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