Telomere length reflects reproductive effort indicated by corticosterone levels in a long-lived seabird.

Telomere length (TL) is a candidate biomarker of ageing and phenotypic quality, but little is known of the (physiological) causes of TL variation. We previously showed that individual common terns Sterna hirundo with high reproductive success had short telomeres independent of age, and this pattern was particularly strong in the longer telomeres of the within-individual TL distribution. To test whether this relation can be attributed to effects of reproductive effort, we investigated baseline corticosterone in relation to reproductive success (number of fledglings) and TL. In this context, we assume that variation in baseline corticosterone can be interpreted as index of energy expenditure and allostatic load. Males with higher corticosterone levels during incubation, compared between and within individuals, achieved higher reproductive success and had shorter telomeres. The effect on telomeres was more pronounced in corticosterone measured later in incubation and in the longer telomeres of the within-individual TL distribution. Female corticosterone level during incubation was neither related to reproductive success nor to TL. That we observed these effects only in males mirrors different parental roles during reproduction in the common tern, where males do most of the chick provisioning. The negative association between reproductive success and TL suggests individual differences in reproductive effort as reflected in, or mediated by, baseline corticosterone. We see this result as a promising step towards unravelling the physiological causes of variation in TL and the costs of reproduction.

Evaluation of Two Miniplate Systems and Figure-of-eight Bandages for Stabilization of Experimentally Induced Ulnar and Radial Fractures in Pigeons ( Columba livia ).

Although plate fixation has advantages over other fixation methods for certain indications, it is rarely used in avian surgery, especially in birds that weigh less than 1000 g. Exceptionally small plating systems for these birds are required, which are relatively expensive and difficult to insert. To study avian fracture healing after repair using miniplates, we evaluated 2 steel miniplate systems in 27 pigeons ( Columba livia ) divided into 4 groups. In each pigeon, the left ulna and radius were transected and the ulna was repaired with a bone plate. In groups A and B, a 1.3-mm adaption plate was applied without and with a figure-of-eight bandage; in groups C and D, a 1.0-mm compression plate was applied without and with a bandage, respectively. Healing was evaluated with radiographs after 3, 14, and 28 days; flight tests were conducted after 14, 21, and 28 days; and the wing was macroscopically examined after euthanasia of birds on day 28. Fractures healed without bending or distortion of the plate in all 27 birds, and no significant differences in healing were found between treatment groups. At the end of the study, 23 pigeons (85.2%) showed good or very good flight ability. Results show the 1.3-mm adaption plate and the 1.0-mm compression plate meet the requirements for avian osteosynthesis and can be recommended for fracture repair of the ulna or other long bones in birds weighing less than 500 g. The application of a figure-of-eight bandage might be beneficial in fracture healing.

Predicting reproductive success from hormone concentrations in the common tern (Sterna hirundo) while considering food abundance.

In birds, reproductive success is mainly a function of skill or environmental conditions, but it can also be linked to hormone concentrations due to their effect on behavior and individual decisions made during reproduction. For example, a high prolactin concentration is required to express parental behaviors such as incubation or guarding and feeding the young. Corticosterone level, on the other hand, is related to energy allocation or stress and foraging or provisioning effort. In this study, we measured individual baseline prolactin and corticosterone between 2006 and 2012 in breeding common terns (Sterna hirundo) using blood-sucking bugs. Reproductive parameters as well as prey abundance on a local and a wider scale were also determined during this period. Baseline prolactin and corticosterone varied significantly between years, as did breeding success. At the individual level, prolactin was positively and corticosterone was negatively linked to herring and sprat abundance. At the population level, we also found a negative link between corticosterone and prey abundance, probably reflecting overall foraging conditions. High prolactin during incubation was mainly predictive of increased hatching success, potentially by supporting more constant incubation and nest-guarding behavior. It was also positively linked to a lesser extent with fledging success, which could indicate a high feeding rate of young. Corticosterone concentration was positively related to high breeding success, which may be due to increased foraging activity and feeding of young. In general, our study shows that baseline prolactin and corticosterone levels during incubation can predict reproductive success, despite the presence of an interval between sampling and hatching or fledging of young.

Mothers under stress? Hatching sex ratio in relation to maternal baseline corticosterone in the common tern (Sterna hirundo).

Sex ratio of progeny should be balanced if costs and benefits of rearing sons and daughters are equal. However, shifts in sex ratio have been demonstrated across bird species and it was suggested that females are able to adjust the primary sex ratio. One possible mechanism is the glucocorticoid corticosterone which rises under stressful conditions and can be deposited into egg yolk by mothers. We analysed primary sex ratio of common terns Sterna hirundo from 2006 to 2008 and related it to maternal baseline corticosterone level, laying date and year. Therefore, we took 101 blood samples of 71 breeding females via blood sucking bugs, a method with negligible stress for the birds. Sex ratio did not differ from parity in any of the analysed years, which were characterized by poor food availability and breeding success. Only within 1 year there was a tendency for more females in the last hatched chick. Neither corticosterone level nor laying date or year showed an influence on hatching sex ratio. The negative result concerning primary sex ratio and maternal baseline corticosterone level might suggest conditions to be good enough for mothers to prevent them from depositing high levels of corticosterone into eggs.

Why do experienced birds reproduce better? Possible endocrine mechanisms in a long-lived seabird, the common tern.

The influence of age or breeding experience on reproductive success in vertebrates is well known but physiological mechanisms came into focus just recently. Assessing hormone levels could allow insights into these mechanisms and reproductive strategies in long-lived species. Prolactin and corticosterone are two hormones involved in breeding decisions: high prolactin values are necessary for expressing breeding behavior whereas corticosterone is related to activity or stress. We analyzed baseline prolactin and corticosterone under field conditions in common terns (Sterna hirundo) between 2006 and 2010. We took 760 blood samples of 346 known birds 9-14 days after their clutch completion, obtained via blood-sucking bugs (Dipetalogaster maximus), a non-invasive method with negligible stress for the birds. Many individuals were sampled repeatedly during the study period allowing investigation of hormone change on individual level. Prolactin levels increased during the early breeding career, which was confirmed on individual level, whereas corticosterone levels increased mostly in experienced birds, more pronounced in males. Low hormone levels during the first years of breeding could indicate a reduced ability of the endocrine system to secret hormones or it might express a down-regulation to limit parental expenditure. Higher corticosterone values of males could be related to increased foraging activity. Amongst the oldest birds, prolactin seemed to increase in males but not in females. This possible consequence of female senescence might be compensated by high values of male mates. Body mass showed only a weak positive correlation with prolactin level and no correlation with corticosterone concentration.

Regulation of breeding behavior: do energy-demanding periods induce a change in prolactin or corticosterone baseline levels in the common tern (Sterna hirundo)?

Hormones are involved in reproductive decisions, linking environmental cues and body condition and adapting behavior. Mass loss is often accompanied by decreased prolactin and increased corticosterone concentrations, influencing incubation and brooding behavior and ultimately triggering nest desertion. Using blood-sucking bugs (Dipetalogaster maxima), we measured baseline prolactin, corticosterone, and ketone body values in incubating common terns (Sterna hirundo) between 2006 and 2009 during energy-demanding periods: 50 pairs were sampled hungry (after an incubation bout) and again fed (after foraging). In a second approach, we sampled 57 other pairs (experienced and inexperienced birds) three times over their individual breeding period, because reproduction, especially chick rearing, is a very energy-demanding process. In line with the common physiological pattern of fasting, we found significantly lower baseline prolactin values in hungry terns, which were negatively related to mass loss over the incubation bout, whereas corticosterone and ketone body levels were marginally increased. Compared to that in the incubation phase, the prolactin level dropped after hatching of chicks in inexperienced birds, perhaps indicating lower parental expenditure. Corticosterone, on the other hand, increased after hatching in males, probably linked to higher foraging activity, as males mainly deliver food during the first days. These energy-demanding periods clearly influenced hormones and ketone bodies, maybe reinforced by the low energy margin of this small seabird species, but energy reserves were not depleted to a level affecting behavior or reproductive success.

Is the additional effort of renesting linked to a hormonal change in the common tern?

The production of and care for a replacement clutch can bear costs in terms of future reproduction or survival. However, renesting is quite common among seabirds and can contribute considerably to individual fitness. Prolactin and corticosterone are two hormones involved in the mediation of breeding behavior and, as they are linked to body condition or effort, it is of interest if these hormone values change during a second demanding breeding phase within a year. We compared baseline prolactin and corticosterone between the first and the renesting attempt in common terns (Sterna hirundo) on individual level. Therefore, in addition to control birds, 37 breeders were sampled during incubation of their first and their replacement clutch in 2008 and 2009. Blood samples were taken non-invasively by blood-sucking bugs. Prolactin level was lower during the renesting period, especially in birds which abandoned their clutch afterwards, whereas corticosterone did not change. Excluding the deserting birds, the reduced prolactin level was not linked to minor success, but could be related to seasonal processes. The control group of late laying common terns showed comparably low prolactin values, but increased corticosterone concentrations. Renesting individuals exhibited higher prolactin during incubation of their first clutch than non-renesting birds, probably indicating their higher quality. The fact that terns still have relatively high prolactin and low corticosterone values during renesting might confirm their higher quality and suggests that they are able to meet the costs of a second demanding breeding period without being considerably stressed.