Steroids and Reproductive Biology in the Blotched Blue-tongued Lizard, Tiliqua nigrolutea

Abstract

This thesis documents the annual profiles of the primary reproductive steroids testosterone (T), 17beta-oestradiol (E2) and progesterone (P4), in the reproductive cycles of male and female blue-tongued lizards, Tiliqua nigrolutea. Data collected from a large captive population over three consecutive reproductive seasons are included. Reproductive cycles are discussed in the context of other viviparous squamate reptiles, while a broader comparative approach is used to consider patterns of steroid biosynthesis and peripheral metabolism. The annual patterns of circulating concentrations of T, E2 and P4 have been characterised for both sexes. In males, peak plasma T (10.9 +/- 3.00 ng ml-1) and E2 (778.0 +/- 120.00 pg ml-1) concentrations occur coincident with late spermatogenesis and observations of mating, respectively. Plasma P4 concentrations remain basal (< 1.2 ng ml-1) throughout the annual reproductive cycle. In females, increasing plasma E2 concentrations (275.2 +/- 33.87 pg ml-1 - 715.1 +/- 106.68 pg ml-1) are associated with vitellogenesis and plasma T peaks (6.3 +/- 0.63 ng ml-1) in the mating and peri-ovulatory period. In pregnant females, plasma P4 concentrations are elevated for the first two thirds of gestation, peaking in the second trimester at 12.7 +/- 1.27 ng ml-1 and falling rapidly prior to parturition. Concurrently, plasma P4 concentrations in non-reproductively active adult females remain basal (1 - 2 ng ml-1) throughout the year. There is good circumstantial evidence for a multiennial reproductive cycle in females. Parturition occurs late in the active season, presumably leaving little time for females to store sufficient fat reserves to become vitellogenic in the following spring: reproductive opportunities are effectively missed in at least one year following a reproductive effort. Observed reproductive behaviours, including agonistic male - male interactions, mating, and parturition, are documented. An investigation of gonadal steroid biosynthetic pathways in this viviparous squamate is presented. This compares variation in the relative contributions of the delta-4 and delta-5 steroidogenic pathways according to sex and reproductive condition. The delta-4 pathway predominates in both sexes, aligning this species phylogenetically with other reptiles. However, there are clear differences between sexes and with changing reproductive condition in the patterns of production of pathway intermediates and end-products. Additionally, detection of a possibly novel polar steroid as a major end-product of steroid biosynthesis in both sexes is reported. Peripheral (extragonadal) metabolism of T and E2 in a number of reproductively relevant steroid target tissues is compared at times of year chosen to represent three clearly distinctive reproductive conditions in each sex. There are differences both between sexes, between tissue types and with changing reproductive condition in the relative proportions of steroid conjugates and non-conjugated derivatives produced. Biosynthetic pathway activity and peripheral steroid metabolism both appear to be plastic in response to changing reproductive condition in Tiliqua nigrolutea. With a comprehensive database of information about the reproductive endocrinology and physiology of Tiliqua nigrolutea, this species is now available as a model to further examine selected aspects of the steroid hormone control of reproductive physiology and behaviour in a cool temperate, viviparous reptile.