Food availability, energetic constraints and reproductive development in a wild seasonally breeding songbird

1. In many organisms, food availability is a proximate cue that synchronizes seasonal development of the reproductive system with optimal environmental conditions. Growth of the gonads and secondary sexual characteristics is orchestrated by the hypothalamic–pituitary–gonadal (HPG) axis. However, our understanding of the physiological mechanisms by which food availability modulates activity of the HPG axis is limited. 2. It is thought that many factors, including energetic status, modulate seasonal reproductive activation. We tested the hypothesis that food availability modulates the activity of the HPG axis in a songbird. Specifically, we food‐restricted captive adult male Abert's Towhees Melozone aberti for 2 or 4 weeks during photoinduced reproductive development. A third group (control) received ad libitum food throughout. We measured multiple aspects of the reproductive system including endocrine activity of all three levels of the HPG axis [i.e. hypothalamic gonadotropin‐releasing hormone‐I (GnRH‐I), plasma luteinizing hormone (LH) and testosterone (T)], and gonad morphology. Furthermore, because gonadotropin‐inhibitory hormone (GnIH) and neuropeptide Y (NPY; a potent orexigenic peptide) potentially integrate information on food availability into seasonal reproductive development, we also measured the brain levels of these peptides. 3. At the hypothalamic level, we detected no effect of food restriction on immunoreactive (ir) GnRH‐I, but the duration of food restriction was inversely related to the size of ir‐GnIH perikarya. Furthermore, the number of ir‐NPY cells was higher in food‐restricted than control birds. Food restriction did not influence photoinduced testicular growth, but decreased plasma LH and T, and width of the cloacal protuberance, an androgen‐sensitive secondary sexual characteristic. Returning birds to ad libitum food availability had no effect on plasma LH or T, but caused the cloacal protuberance to rapidly increase in size to that of ad libitum‐fed birds. 4. Our results support the tenet that food availability modulates photoinduced reproductive activation. However, they also suggest that this modulation is complex and depends upon the level of the HPG axis considered. At the hypothalamic level, our results are consistent with a role for the GnIH and NPY systems in integrating information on energetic status. There also appears to be a role for endocrine function at the anterior pituitary gland and testicular levels in modulating reproductive development in the light of energetic status and independently of testicular growth

Reproductive endocrinology of the Wandering Albatross Diomedea exulans in relation to biennial breeding and deferred sexual maturity

The reproductive endocrinology of the Wandering Albatross Diomedea exulans was studied at South Georgia to investigate the potential endocrine correlates of biennial breeding and of the acquisition of sexual maturity. Gonads of breeding birds and of known‐age immature birds of both sexes were examined by laparoscopy throughout the period that they were at the nest site. Blood samples, subsequently analysed to determine concentrations of luteinizing hormone (LH), prolactin, progesterone, testosterone and oestradiol‐17/i, were obtained from samples of breeding birds of both sexes at regular intervals from first arrival until the chicks fledged nearly a year later. Before laying in December, breeding birds had mature testes and ovarian follicles and high concentrations of LH, prolactin and sex steroids. Gonadal regression and a rapid drop in hormone levels (except for LH in females) occurred in early incubation (January). Testes (and follicles to a lesser extent) enlarged in mid‐incubation, coinciding with high levels of LH and increases in prolactin and testosterone. Gonads finally regressed completely near hatching time. LH, prolactin and testosterone remained at low levels throughout chick rearing (April to November), but females had several periods of active progesterone and oestradiol secretion, and progesterone was detectable in males only late in the chick‐rearing period. Although some changes in hormone levels are difficult to explain, the patterns are fairly typical of temperate birds. The persistence of progesterone secretion in both female breeders and non‐breeding ‘immature’ birds is viewed as part of a mechanism inhibiting an ovary from becoming vitellogenic. Although testis size and testosterone concentrations increased with age in immature males (of ages 4–10 years), birds of 5 years and older are probably physiologically mature, even though breeding does not start until they are 7 years of age and only half an age group has bred by an age of 11 years. Immature females (of age 4–7 years) had undeveloped follicles, very low oestradiol concentrations but high progesterone levels, providing further support for the role of this hormone in inhibiting gonadotropin secretion. The condition of the female is therefore probably decisive in determining when a pair first attempts to breed but it is unknown what factors initiate normal ovarian development

The Effect of Pinealectomy on Plasma Levels of Gonadotrophins and Growth Hormone in the Pigeon (Columba livid)

peer reviewe

The endocrine basis of deferred sexual maturity in the wandering albatross, Diomedea exulans. L.

1 Earlier work with albatrosses led to the hypothesis that a critical element influencing ovarian development each year is whether the ovary responds to rising gonadotrophin secretion by secreting progesterone or oestradiol. In the former case no vitellogenesis occurs, the ovary does not mature and no egg-laying occurs. In the latter case ovarian growth and ovulation occurs. This `switch' between progesterone and oestradiol secretion may be important not only for the breeding frequency of adult birds but also prepubertally, influencing the onset of sexual maturity. This paper tests some of the underlying endocrine responses experimentally using known-age (4-11 years old) wandering albatrosses, Diomedea exulans L., at South Georgia. 2 Experiments using birds before and after bouts of sexual display discounted the possibility that high progesterone levels were an artefact of behavioural interaction. 3 Examination of the pituitary-ovarian axis, by injection of ovine luteinizing hormone (LII) and LH-releasing hormone, confirmed that the ovary does secrete progesterone, and that only progesterone is produced in immatures, unlike breeding birds in which oestradiol is also secreted. 4 In terms of endocrine status, female D. exulans were divisible into three main groups: younger (7 years) and last-time non-breeders. These categories were consistent with behavioural data from the same population. Thus the earliest age of breeding is 7 years, display rates increase after `physiological maturity' is attained but decrease once stable pairs are formed and attendance ashore prior to maturity does not influence the time taken to start breeding

Changes in pituitary, ovarian, and testicular activity in harbour seals (Phoca vitulina) in relation to season and sexual maturity

This study examined pituitary sensitivity to exogenous gonadotrophin-releasing hormone (GnRH) in relation to time of year and changes in plasma progesterone, testosterone, luteinising hormone (LH), and follicle-stimulating hormone (FSH) concentrations during the annual cycle of adult and juvenile harbour seals (Phoca vitulina vitulina). There was no significant difference in seasonal changes in reproductive hormone levels between six captive individuals and seals caught from the wild (n = 80). Based on results from both the captive and the wild individuals, the plasma progesterone concentration was elevated in late gestation, whereas it declined at the equivalent time in nonpregnant females. The progesterone concentration was also elevated during the phase of delayed implantation, but there was no difference between pregnant and nonpregnant seals. In captive seals given exogenous GnRH, the LH concentration peaked 10-20 min after injection of GnRH, and the magnitude of the LH response depended upon the dose up to ~120 ng·kg-1. FSH did not respond to GnRH in the time course of the experiment. Pituitary sensitivity to GnRH was greatest in the summer and autumn and lowest in the winter and spring. Seasonal changes in pituitary sensitivity were apparent in both adults and juveniles. There was evidence that three of the four juvenile seals attained puberty during the study. We conclude that the seasonal patterns of pituitary sensitivity and plasma hormone concentration in harbour seals are consistent with a reproductive cycle under photoperiodic control and with the general mechanisms underlying photoperiodic control of seasonal reproduction in higher vertebrates