Search

Photoperiodic control of reproduction in birds is based on two processes, a positive effect leading to gonadal maturation and an inhibitory effect subsequently inducing regression. Nonphotoperiodic cues can modulate photoperiodic control, particularly the inhibitory process. In previous studies of common starlings (Sturnus vulgaris), (1) restriction of food availability to 8 h after dawn had little effect on testicular maturation but dramatically delayed subsequent regression and (2) lower ambient temperature also had little effect during maturation but delayed regression. Could the effects of food restriction and temperature share a common underlying mechanism? Four groups of starlings were kept on a simulated natural cycle in photoperiod in a 2 x 2 factorial experimental design. Two groups were held under an ambient temperature of 16°C, and the other two were held under 6°C.One of each of these groups had food provided ad lib., and in the other two groups access to food was denied 7 h after dawn. In both the ad lib. food groups and the food-restricted groups, lower temperature had little effect on testicular maturation but delayed subsequent regression and molt. In both the 16°Cgroups and the 6°Cgroups, food restriction had no effect on testicular maturation but delayed regression and molt. The daily cycle in body temperature was recorded in all groups when the photoperiod had reached 12L∶12D, the photoperiod at which regression is initiated. In both 6°C groups, nighttime body temperature was lower than in the 16°C groups, a characteristic of shorter photoperiods. In the two ad lib. food groups high daytime temperature was maintained until dusk, whereas in the two food-restricted groups body temperature began to decrease after food withdrawal. Thus, both lower temperature and food restriction delayed regression, as if the photoperiod was shorter than it actually was, and both resulted in daily cycles in body temperature that reflected cycles under shorter photoperiods. This implies that the daily cycle in body temperature is possibly a common pathway through which nonphotoperiodic cues may operate. [ABSTRACT FROM AUTHOR]

Although it is well known that body temperature (Tb) is higher during the day in diurnal birds than at night, no data are available regarding exactly how Tb varies during a 24-h period, how this differs under different photoperiods, and how it responds to a change in photoperiod. This study used implanted temperature loggers in starlings (Sturnus vulgaris) to address these questions. The duration of elevated Tb was directly related to photoperiod, but the amplitude of the daily cycle was significantly greater under shorter photoperiods. Under all photoperiods, Tb started to increase before dawn and continued to increase after dawn; there was no sudden change associated with dawn. In contrast, Tb decreased immediately and rapidly at dusk (significantly by 15 min). The daily cycle in Tb rapidly adjusted to a change in photoperiod. Following an acute increase in photoperiod, Tb increased immediately at the new earlier dawn but did not decrease until the new later dusk. Following a decrease in photoperiod, Tb did not increase after the time of the missed dawn; it only increased after the new later dawn. It decreased at the new earlier dusk. Following transfer to constant darkness, there was a moderate increase in Tb around the missed dawn, but then Tb gradually decreased before the missed dusk to lower values than during the previous night. The results suggest that the daily cycle in Tb is weakly circadian and may be entrained by dusk rather than dawn. [ABSTRACT FROM AUTHOR]

This paper reviews current knowledge of photoperiod control of GnRH-1 secretion and proposes a model in which two processes act together to regulate GnRH1 secretion. Photo-induction controls GnRH1 secretion and is directly related to prevailing photoperiod. Photo-inhibition, a longer term process, acts through GnRH1 synthesis. It progresses each day during daylight hours, but reverses during darkness. Thus, photo-inhibition gradually increases when photoperiods exceed 12 h, and reverses under shorter photoperiods. GnRH1 secretion on any particular day is the net result of these two processes acting in tandem. The only difference between species is their sensitivity to photo-inhibition. This can potentially explain differences in timing and duration of breeding seasons between species, why some species become absolutely photorefractory and others relatively photorefractory, why breeding seasons end at the same time at different latitudes within species, and why experimental protocols sometimes produce results that appear counter to what happens naturally. [ABSTRACT FROM AUTHOR]

Clutch size is a key avian fitness and life history trait. A physiological model for clutch size determination (CSD), involving an anti-gonadal effect of prolactin (PRL) via suppression of luteinizing hormone (LH), was proposed over 20 years ago, but has received scant experimental attention since. The few studies looking at a PRL-based mechanistic hypothesis for CSD have been equivocal, but recent experiments utilizing a pharmacological agent to manipulate PRL in the zebra finch ( Taeniopygia guttata ) found no support for a role of this hormone in clutch size determination. Here, we take a complementary approach by manipulating clutch size through egg removal, examining co-variation in PRL and LH between two breeding attempts, as well as through experimentally-extended laying. Clutch size increased for egg removal females, but not controls, but this was not correlated with changes in PRL or LH. There were also no differences in PRL between egg removal females and controls, nor did PRL levels during early, mid- or late-laying of supra-normal clutches predict clutch size. By uncoupling PRL, LH and clutch size in our study, several key predictions of the PRL-based mechanistic model for CSD were not supported. However, a positive correlation between PRL levels late in laying and days relative to the last egg (clutch completion) provides an alternative explanation for the equivocal results surrounding the conventional PRL-based physiological model for CSD. We suggest that females coordinate PRL-mediated incubation onset with clutch completion to minimize hatching asynchrony and sibling hierarchy, a behavior that is amplified in females laying larger clutches. [ABSTRACT FROM AUTHOR]

Highlights: [•] We examined the role of pre-breeding and breeding PRL and LH in avian clutch size. [•] Pre-breeding LH supports individual differences in ‘reproductive readiness’. [•] PRL and LH were correlated only during the pre-breeding period. [•] Neither PRL nor LH were related to clutch size for control or experimental birds. [•] Our data do not support clutch size determination involving circulating PRL or LH. [ABSTRACT FROM AUTHOR]

Highlights: [•] Maturation starts earlier at lower latitudes; regression starts at the same time. [•] Photoperiodic responses alone can explain earlier gonadal maturation. [•] The mechanism for timing regression remains unclear. [•] Absolute and relative photorefractory species use the same photoperiodic mechanism. [Copyright &y& Elsevier]

In vertebrates, reproductive output often increases with age. Unlike older birds, first-year photoperiodic birds lack experience with the reproductively stimulatory effects of long day lengths (photostimulation). We examined whether age-related differences in annual reproductive development could be partially attributed to previous experience with photostimulation in the photoperiodic house finch (Carpodacus mexicanus). By manipulating photoperiod, we generated 2 groups of first-year females: a photo-experienced group that underwent 1 photoperiodically induced cycle of gonadal development and regression and a photo-naïve group exposed to long days since hatch. We transferred both groups from long to short days and then photostimulated and exposed them to male birdsong prior to sacrifice. Following concurrent photostimulation, both groups exhibited similar plasma luteinizing hormone surges and hypothalamic vasoactive intestinal polypeptide immunoreactivity. In contrast, hypothalamic gonadotropin-releasing hormone immunoreactivity and circulating vitellogenin levels were higher in photo-experienced birds, and yolk deposition occurred in only 2 females, both of which were photo-experienced. Our results demonstrate that photo-experience enhances some aspects of early photo-induced reproductive development and raise the hypothesis that photo-experience may account for at least some age-related variation in reproductive output. [ABSTRACT FROM PUBLISHER]

Abstract: Most animals reproduce seasonally. They time their reproduction in response to environmental cues, like increasing photoperiod and temperature, which are predictive for the time of high food availability. Although individuals of a population use the same cues, they vary in their onset of reproduction, with some animals reproducing consistently early or late. In avian research, timing of reproduction often refers to the laying date of the first egg, which is a key determinant of fitness. Experiments measuring temporal patterns of reproductive hormone concentrations or gonadal size under controlled conditions in response to a cue commonly assume that these proxies are indicative of the timing of egg laying. This assumption often remains untested, with few studies reporting both reproductive development and the onset of laying. We kept in total 144 pairs of great tits (Parus major) in separate climate-controlled aviaries over 4years to correlate pre-breeding plasma luteinizing hormone (LH), prolactin (PRL) and gonadal growth with the timing of laying. Individuals varied consistently in hormone concentrations over spring, but this was not directly related to the timing of gonadal growth, nor with the laying date of the first egg. The timing of gonadal development in both sexes was similarly not correlated with the timing of laying. This demonstrates the female’s ability to adjust the onset of laying to environmental conditions irrespective of substantial differences in pre-laying development. We conclude that stages of reproductive development are regulated by different cues, and therefore egg laying dates need to be studied to measure the influences of environmental cues on timing of seasonal reproduction. [Copyright &y& Elsevier]

Timing of reproduction in temperate-zone birds is strongly correlated with spring temperature, with an earlier onset of breeding in warmer years. Females adjust their timing of egg laying between years to be synchronized with local food sources and thereby optimize reproductive output. However, climate change currently disrupts the link between predictive environmental cues and spring phenology. To investigate direct effects of temperature on the decision to lay and its genetic basis, we used pairs of great tits (Parus major) with known ancestry and exposed them to simulated spring scenarios in climate-controlled aviaries. In each of three years, we exposed birds to different patterns of changing temperature. We varied the timing of a temperature change, the daily temperature amplitude, and the onset and speed of a seasonal temperature rise. We show that females fine-tune their laying in response to a seasonal increase in temperature, whereas mean temperature and daily temperature variation alone do not affect laying dates. Luteinizing hormone concentrations and gonadal growth in early spring were not influenced by temperature or temperature rise, possibly posing a constraint to an advancement of breeding. Similarities between sisters in their laying dates indicate genetic variation in cue sensitivity. These results refine our understanding of how changes in spring climate might affect the mismatch in avian timing and thereby population viability. [ABSTRACT FROM AUTHOR]

Abstract: In avian species that have evolved life-history strategies wherein molt and breeding overlap, there are potential conflicts between the regulatory roles of baseline prolactin and corticosterone in parental care (positive) and moult (negative). We describe seasonal patterns of hormonal secretion, moult, and parental behaviour in sibling species of giant petrels (Macronectes spp.) which begin moult during the incubation/early chick-rearing stage of reproduction. With the exception of male Southern giant petrels (Macronectes giganteus), prolactin secretion and moult in Northern (Macronectes halli) and female Southern giant petrels conformed to those observed in all other avian species, with the initiation of moult coincident with decreases from peak prolactin levels. However, male Southern giant petrels began moulting early in incubation when prolactin was increasing and had not yet peaked, which suggests a requirement of prolactin for incubation behaviour and a dissociation of prolactin from moult. Corticosterone showed little seasonal variation and no relationship with moult. When comparing prolactin, corticosterone, and moult in failed vs. active breeders, we found that failed breeding enabled a more rapid down-regulation of prolactin, thus facilitating a more rapid moult. We present specific examples of the behavioural ecology of giant petrels which we conclude help mediate any potential hormonal conflicts between parental care and moult. [Copyright &y& Elsevier]

Birds use change in daylength during the year to time events during their annual cycles. Individual Eurasian siskins Carduelis spinus can breed and winter in widely separated areas in different years. Birds at different latitudes will experience different changes in photoperiod. So how does latitude affect photoperiodic control? Our aim in this study was to find whether Siskins caught from the wild in Britain and exposed to different photoperiodic regimes, typical of widely separated latitudes, would differ in the subsequent timing and duration of their moults and associated processes. Siskins were caught in late February and early March, and initially kept outside on natural photoperiods. From the spring equinox (21 March), they were divided into three groups kept under photoperiodic regimes that simulated latitudes 40°, 55° and 70°N respectively. All three groups showed highly significant subsequent changes in body mass, fat scores and cloacal protuberance size. Moult of the primary feathers started during June - August (mean 9 July), and lasted 61-99 days (mean 75 days). Birds that started to moult late in the season had shorter moult durations. All individuals showed lower mass and fat levels during moult than before or after moult. Crucially, there were no significant differences in the timing of these events between the three photoperiodic groups. Apparently these birds did not use prevailing absolute photoperiod or the prevailing rate of change in photoperiod to time moult-related seasonal events, but used instead some other feature of the annual photoperiod cycle or some form of interval timer linked to photoperiod. [ABSTRACT FROM AUTHOR]

Abstract: In a study on starlings (Sturnus vulgaris) kept on a simulated annual cycle in photoperiod, temperature had no effect on the timing or rate of testicular maturation but high temperature resulted in an advance in the timing of testicular regression and molt (). This study asks whether the earlier gonadal regression in response to higher temperature represents a central neuroendocrine response to temperature, and secondly, whether prolactin plays a role in the earlier regression. Castrated starlings were kept on a simulated annual cycle of photoperiod at either 8 or 18°C. Circulating LH and prolactin concentrations were measured and the progress of the post-nuptial molt was recorded as an external indicator of the development of photorefractoriness. Additionally plasma prolactin was measured in samples taken from intact male and female starlings in the 2005 study. In castrated birds, LH concentrations decreased three weeks earlier at 18°C. These birds also showed the same three week advance in molt as males and females in the earlier study. This demonstrates that the advance in regression caused by higher temperatures probably results from a central neuroendocrine mechanism, i.e., an advance in photorefractoriness, rather than an effect at the level of the gonads. Temperature had a highly significant effect on the changes in prolactin – peak prolactin occurred three weeks earlier at 18°C. However, there was no clear consistent significant difference in prolactin between the two temperatures in advance of the onset of photorefractoriness, so the advance in photorefractoriness may not be mediated by prolactin. The higher temperature resulted in a significantly earlier decrease in prolactin and this may be causally related to the advance in molt. [Copyright &y& Elsevier]

Abstract: Avian breeding seasons vary in length and in the degree of asymmetry with respect to the annual cycle in photoperiod to suit species-specific food resources. Asymmetry is the result of photorefractoriness. The degree of photorefractoriness, absolute or relative, is related to the length and asymmetry of the breeding season. Absolute photorefractoriness is associated with a marked decrease in hypothalamic cGnRH-I. However, during the initiation of absolute photorefractoriness there is a transient period during which the gonads regress in advance of the decrease in cGnRH-I, and this stage may be analogous to relative photorefractoriness. Photoinduced prolactin secretion has an inhibitory modulatory role during the initiation of absolute photorefractoriness, but is unlikely to be the only factor involved, while a possible role for avian gonadotrophin inhibitory hormone is not established. The first stage in the termination of photorefractoriness is the resumption of cGnRH-I synthesis. The major environmental cue driving gonadal maturation, and the transitions between the photosensitive state and photorefractoriness is the annual cycle in photoperiod. A range of non-photoperiodic cues may also play a role: social cues, climatic factors (temperature, rainfall, etc.), food availability and nutritional state. There is considerable evidence that these cues can influence gonadal maturation and the timing of egg-laying. There is some evidence that non-photoperiodic cues (certainly temperature and possibly social cues and food availability) can affect the timing of the onset of photorefractoriness, but no evidence that they can influence the time of the end of photorefractoriness. [Copyright &y& Elsevier]

Abstract: Despite the importance of molt to birds, very little is known about its environmental or physiological control. In starlings Sturnus vulgaris, and other species, under both natural conditions and experimental regimes, gonadal regression coincides with peak prolactin secretion. The prebasic molt starts at the same time. The aim of this series of experiments was to keep starlings on photo-schedules that would challenge the normally close relationship between gonadal regression and molt, to determine how closely the start of molt is associated with gonadal regression and/or associated changes in prolactin concentrations. In one series of experiments, photosensitive starlings were moved from a short photoperiod, 8h light per day (8L), to 13 or 18L, and from 13 to 18L or 13 to 8L during testicular maturation. Later, photorefractory birds under 13L that had finished molting were moved to 18L. In another series of experiments, photorefractory starlings were moved from 18 to 8L for 7 weeks, 4 weeks, 2 weeks, 1 week, 3 days, 1 day, or 0 days, before being returned to 18L. There was no consistent relationship between photoperiod, or the increase in photoperiod, and the timing of the start of molt. Nor was there a consistent relationship with gonadal regression and the start of molt–molt could be triggered in the absence of a gonadal cycle. However, there was always an association between the start of molt and prolactin. In all cases where molt was induced, there had been an earlier increase in prolactin. However, the timing of molt was related to the time of peak prolactin, not the magnitude of that peak. This relationship between peak prolactin and the start of molt could explain the normally close relationship between the end of breeding activity and the start of molt. [Copyright &y& Elsevier]

Exposure to organophosphate (OP) pesticides can occur in free-living mammals in treated areas. Risk to nontarget animals from OPs usually is assessed with acute exposure data, but exposure of wild animals is likely to be intermittent and chronic. We compared the effects of single or repeated (hourly and daily) exposure to dimethoate on acetylcholinesterase (AChE) activity in laboratory mice to assess the suitability of standard laboratory tests for assessing risk. Mice were exposed either to a single dose (10 or 30 mg/kg) or to short-term repeated (three hourly doses of 10 mg/kg) intraperitoneal doses of dimethoate, and brain and serum AChE activity were measured. No significant difference was found in the degree of inhibition of AChE activity following acute and short-term repeated exposure. In a second experiment, mice were given three daily doses of 10 or 20 mg/kg of dimethoate, and both AChE activity and hepatic cytochrome P450 enzyme activity were measured. Daily exposure resulted in a dose-dependent decline in brain and serum AChE activity, and inhibition increased progressively with successively repeated exposures. However, this effect was relatively small compared to the effect of dose. Cytochrome P450 enzyme activity (CYP2B) was inhibited in the dimethoate-dosed mice. Our results indicate that acute dose-response toxicity studies are suitable models for predicting the likely occurrence of adverse effects from either short- or longer-term exposure of wild mammals to anticholinesterase compounds. Likely differences in exposure pattern between the laboratory and the natural environment are unlikely to bias the predictive power of these studies significantly. [ABSTRACT FROM AUTHOR]

We previously reported evidence of West Nile virus (WNV) circulation in UK birds, probably introduced by migratory birds from overseas. We now demonstrate WNV-specific seroconversion in sentinel chickens raised on an English farm. Maternal neutralizing antibodies to WNV in hatchlings declined within three weeks. During the following months, healthy chickens developed WNV neutralizing antibodies that were confirmed by immunoblotting and indirect immunofluorescence tests using WNV antigens. The proportion of seropositive chickens was higher for WNV than for Usutu virus or Sindbis virus. Attempts to isolate infectious virus or to detect viral RNA in the sera, failed. [ABSTRACT FROM AUTHOR]

Abstract: In many bird species, males exhibit territorial aggression outside the breeding season, when testosterone concentrations are low and may not regulate territorial behaviors. The hormonal regulation of aggression at this time of year has only been studied in passerine birds. Here, we investigated the role of testosterone in the regulation of aggression in a non-passerine bird, the red grouse Lagopus lagopus scoticus. Male red grouse are aggressive in early spring when breeding starts, in autumn when they establish territories, and sporadically through much of the winter. We first describe seasonal variations in plasma testosterone concentrations and in the size of males'' sexual ornaments, their red combs, which relates to aggressiveness. Testosterone concentrations and comb size were correlated. Both increased in autumn to a peak in October, and then increased again in spring, to a greater peak in early April. Secondly, we experimentally investigated the effects of testosterone, and of an anti-androgen (flutamide) used in combination with an aromatase inhibitor (ATD), on autumn territorial behavior. Males were treated with either empty implants, as controls (C-males), testosterone implants (T-males), or with flutamide and ATD implants (FA-males). One month after implanting, both T- and FA-males had higher concentrations of testosterone than C-males. Comb size, aggressive call rate, and response to playbacks of territorial call all significantly increased in T-males. However, the increase in testosterone in FA-males did not increase comb size or aggressive behavior. In the following spring, after the content of implants was used, FA-males had significantly lower testosterone than C-males, and had a reduced seasonal increase in comb size. The results suggest that testosterone plays a significant role in regulating red grouse aggressive behavior in autumn. However, the observation that flutamide and ATD treatment did not reduce territorial behavior, suggests that estradiol may also be involved in the regulation of non-breeding aggression. [Copyright &y& Elsevier]

In European Starlings (Sturnus vulgaris), mass of new primary feathers increases linearly with time for most of the duration of molt (Dawson 2003). Here, we had two aims: (1) to confirm that mass of new primary feathers increases linearly in other species, and (2) to use that linear increase as the basis of a system to score the progress of molt. Increase in length of primary feathers during molt was recorded for captive Eurasian Magpies (Pica pica), Carrion Crows (Corvus corone), European Greenfinches ( Carduelis chloris), and Eurasian Bullfinches (Pyrrhula pyrrhula). Wing shape differs among those species. Feather lengths during molt were converted into mass by using information on final feather length and mass and the distribution of mass from tip to base of feather. Rate of increase in total mass of new primary feathers was largely linear in all four species. A molt scoring method is described in which individual feather scores are weighted to account for the contribution of each particular primary feather's mass toward total primary-feather mass. When the method was tested on eight captive starlings, the increase in mass-corrected molt score was almost linear, unlike the increase shown by the standard scoring system, which exaggerated molt rate during the early part of molt and underestimated it later. In the four species studied here, mass-corrected molt score likewise closely tracked the actual increase in mass, unlike the standard molt score. Because it is based on feather mass, the method presented here is of greater physiological and energetic relevance than the standard method. Because the mass-corrected score increases more linearly with time, it has the additional advantage of enabling less complicated, and potentially more accurate, estimations of molt start dates and molt durations. [ABSTRACT FROM AUTHOR]

In many species of birds, the end of the breeding season is caused by the development of absolute photorefractoriness, characterised by spontaneous gonadal regression under a long photoperiod followed by a period during which gonadal maturation cannot be stimulated by further photostimulation. It has been suggested that during the dissipation of photorefractoriness under short photoperiods, birds become relatively photorefractory—that there is a gradual decrease in critical photoperiod. If true, birds should begin to respond to a very long photoperiod before they respond to a moderately long photoperiod. To test this, I transferred photorefractory castrated starlings (Sturnus vulgaris) from a long photoperiod of 18 h of light per day (18L:6D) to 8L:16D for 10, 20, or 30 days, before transferring them to either 20L:4D or to 13L:11D. There was a gradual increase in response (increase in plasma LH concentration) the longer birds had been on 8L:16D before transfer to the longer photoperiods, but there was no difference in the timing of these responses between birds moved to 20L:4D or to 13L:11D. Therefore, starlings do not pass through a period of relative photorefractoriness during the dissipation of absolute photorefractoriness. [Copyright &y& Elsevier]

There is considerable evidence that reproductive success improves with age in birds. It is often suggested that improved performance of older birds is because they are more experienced. Less consideration has been given to the possibility that improvements may be a consequence of age-related changes in reproductive physiology. One factor that consistently changes with age is laying date – first year birds lay later than older birds. In European starlings Sturnus vulgaris , older males begin reproductive activity earlier than first year males and dominate available nest sites. I monitored changes in testicular volume in captive starlings exposed to natural changes in day length and temperature, from their first autumn through the next two annual cycles. Testicular maturation was advanced by 3–4 weeks in birds during their second year compared to the first, and testicular regression occurred about 2 weeks later. The period of full sexual maturation was 50% longer during the second year. The timing of the post-nuptial moult was the same. A possible physiological mechanism to explain this is discussed. The results show that earlier reproductive activity in older birds can be explained, at least in part, by intrinsic physiological mechanisms. This does not preclude additional effects of experience of an individual in the improvement in reproductive performance between first and second year birds. [ABSTRACT FROM AUTHOR]

Abstract Photoperiodic manipulation of young European starlings suggests that their reproductive physiology is incapable of responding to a short photoperiod until they are fully grown. This study aimed to determine whether the lack of response to a short photoperiod is reflected in the daily profile of plasma melatonin concentrations. Five-day-old starlings taken from nest boxes showed a significant (p < 0.0001) rhythm in plasma melatonin concentrations, with high values during night. In nestlings hand-reared from 5 days of age on a long photoperiod (LD 16:8), equivalent to natural photoperiod at the time, the amplitude of the daily rhythm in melatonin increased significantly (p < 0.01) with age until birds were fully grown (20 days old). In nestlings reared on a short photoperiod (LD 8:16), the daily melatonin profile remained almost identical to that of long photoperiod birds until they were fully grown. However, after 20 days old, the duration of elevated nighttime melatonin began to extend to encompass the entire period of darkness. In contrast, fully grown starlings transferred from a long to a short photoperiod had partially adapted to the short photoperiod after 5 days; by 10 days, the daily melatonin profile was identical to that of birds held chronically on a short photoperiod. Thus, consistent with responses of reproductive physiology, the pineal of young birds appears to be incapable of perceiving, or adapting to, a short photoperiod. [ABSTRACT FROM AUTHOR]

This review examines how birds use the annual cycle in photoperiod to ensure that seasonal events—breeding, molt, and song production—happen at the appropriate time of year. Differences in breeding strategies between birds and mammals reflect basic differences in biology. Avian breeding seasons tend to be of shorter duration and more asymmetric with respect to changes in photoperiod. Breeding seasons can occur at the same time each year (predictable) or at different times (opportunistic), depending on the food resource. In all cases, there is evidence for involvement of photoperiodic control, nonphotoperiodic control, and endogenous circannual rhythmicity. In predictable breeders (most nontropical species), photoperiod is the predominant proximate factor. Increasing photoperiods of spring stimulate secretion of gonadotropin-releasing hormone (GnRH) and consequent gonadal maturation. However, breeding ends before the return of short photoperiods. This is the consequence of a second effect of long photoperiods—the induction of photorefractoriness. This dual role of long photoperiods is required to impart the asymmetry in breeding seasons. Typically, gonadal regression through photorefractoriness is associated with a massive decrease in hypothalamic GnRH, essentially a reversal to a pre-pubertal condition. Although breeding seasons are primarily determined by photoperiodic control of GnRH neurons, prolactin may be important in determining the exact timing of gonadal regression. In tropical and opportunistic breeders, endogenous circannual rhythmicity may be more important. In such species, the reproductive system remains in a state of "readiness to breed" for a large part of the year, with nonphotic cues acting as proximate cues to time breeding. Circannual rhythmicity may result from a temporal sequence of different physiological states rather than a molecular or cellular mechanism as in circadian rhythmicity. Avian homologues of mammali... [ABSTRACT FROM AUTHOR]

Breeding periodicity allows organisms to synchronise breeding attempts with the most favourable ecological conditions under which to raise offspring. For most animal species, ecological conditions vary seasonally and usually impose an annual breeding schedule on their populations; sub-annual breeding schedules will be rare. We use a 16-year dataset of breeding attempts by a tropical seabird, the sooty tern (Onychoprion fuscatus), on Ascension Island to provide new insights about this classical example of a population of sub-annually breeding birds that was first documented in studies 60 years previously on the same island. We confirm that the breeding interval of this population has remained consistently sub-annual. By ringing >17000 birds and re-capturing a large sample of them at equivalent breeding stages in subsequent seasons, we reveal for the first time that many individual birds also consistently breed sub-annually (i.e. that sub-annual breeding is an individual as well as a population breeding strategy). Ascension Island sooty terns appear to reduce their courtship phase markedly compared with conspecifics breeding elsewhere. Our results provide rare insights into the ecological and physiological drivers of breeding periodicity, indicating that reduction of the annual cycle to just two life-history stages, breeding and moult, is a viable life-history strategy and that moult may determine the minimum time between breeding attempts. [ABSTRACT FROM AUTHOR]

Abstract: Many bird species have advanced their seasonal timing in response to global warming, but we still know little about the causal effect of temperature. We carried out experiments in climate-controlled aviaries to investigate how temperature affects luteinizing hormone, prolactin, gonadal development, timing of egg laying and onset of moult in male and female great tits. We used both natural and artificial temperature patterns to identify the temperature characteristics that matter for birds. Our results show that temperature has a direct, causal effect on onset of egg-laying, and in particular, that it is the pattern of increase rather than the absolute temperature that birds use. Surprisingly, the pre-breeding increases in plasma LH, prolactin and in gonadal size are not affected by increasing temperature, nor do they correlate with the onset of laying. This suggests that the decision to start breeding and its regulatory mechanisms are fine-tuned by different factors. We also found similarities between siblings in the timing of both the onset of reproduction and associated changes in plasma LH, prolactin and gonadal development. In conclusion, while temperature affects the timing of egg laying, the neuroendocrine system does not seem to be regulated by moderate temperature changes. This lack of responsiveness may restrain the advance in the timing of breeding in response to climate change. But as there is heritable genetic variation on which natural selection can act, microevolution can take place, and may represent the only way to adapt to a warming world. [Copyright &y& Elsevier]

Abstract: The Challenge Hypothesis postulates that male vertebrates can respond to social challenges, such as simulated territorial intrusions, by rapidly increasing their concentrations of plasma androgens, such as testosterone (T). This increase may facilitate the expression of aggressive behavior and lead to persistence of this behavior even after withdrawal of the challenge, thus potentially promoting territoriality and the probability of winning future challenges. The scope of the Challenge Hypothesis was tested by exposing free-ranging male Cassin''s Sparrows, Peucaea cassinii, to conspecific song playback (SPB) at the beginning of the vernal nesting season. Exposure to SPB stimulated aggressive behavior but did not influence plasma T. Furthermore, plasma T did not correlate with the duration of exposure to SPB, and the behavioral response to SPB did not differ in males that were challenged a second time shortly after the first challenge. As birds were investigated at a stage of their reproductive cycle when plasma T is presumably seasonally high due to photostimulation, the lack of hormonal response to SPB may have been due to the hypothalamus–pituitary–gonadal axis secreting hormones at maximum rates. This was not the case, however, because administration of gonadotropin-releasing hormone I rapidly stimulated the secretion of luteinizing hormone (LH) and T, and treatment with ovine LH rapidly stimulated T secretion. [Copyright &y& Elsevier]

Hormones mediate major physiological and behavioural components of the reproductive phenotype of individuals. To understand basic evolutionary processes in the hormonal regulation of reproductive traits, we need to know whether, and during which reproductive phases, individual variation in hormone concentrations relates to fitness in natural populations. We related circulating concentrations of prolactin and corticosterone to parental behaviour and reproductive success during both the pre-breeding and the chick-rearing stages in both individuals of pairs of free-living house sparrows, Passer domesticus. Prolactin and baseline corticosterone concentrations in pre-breeding females, and prolactin concentrations in prebreeding males, predicted total number of fledglings. When the strong effect of lay date on total fledgling number was corrected for, only pre-breeding baseline corticosterone, but not prolactin, was negatively correlated with the reproductive success of females. During the breeding season, nestling provisioning rates of both sexes were negatively correlated with stress-induced corticosterone levels. Lastly, individuals of both sexes with low baseline corticosterone before and high baseline corticosterone during breeding raised the most offspring, suggesting that either the plasticity of this trait contributes to reproductive success or that high parental effort leads to increased hormone concentrations. Thus hormone concentrations both before and during breeding, as well as their seasonal dynamics, predict reproductive success, suggesting that individual variation in absolute concentrations and in plasticity is functionally significant, and, if heritable, may be a target of selection. [ABSTRACT FROM AUTHOR]

Presents a study which looked at the effects of varying light intensity on the photoperiodic reproductive response of starlings. Theories on how photoperiodic time measurement occurs; Insight on the methodology used to conduct the study; Discussion based on the results of the study.

Determining the physiological mechanisms underpinning life-history decisions is essential for understanding the constraints under which life-history strategies can evolve. In long-lived species, where the residual reproductive value of breeders is high, adult survival is a key contributor to lifetime reproductive success. We therefore expect that when adult survival is compromised during reproduction, mechanisms will evolve to redirect resources away from reproduction, with implications for reproductive hormones, adult body mass, nest attendance behaviour and breeding success. We investigated whether manipulating corticosterone, to simulate exposure to an environmental stressor, affected the secretion of prolactin and breeding success in the black-legged kittiwake Rissa tridactyla . We used implanted Alzet® osmotic pumps to administer corticosterone to incubating kittiwakes at a constant rate over a period of approximately 8 days. Manipulated birds were compared with sham implanted birds and control birds, which had no implants. There was no significant difference in the body mass of captured individuals at the time of implantation and implant removal. Corticosterone-implanted males showed lower nest attendance during the chick rearing period compared to sham-implanted males; the opposite pattern was found in females. Corticosterone treated birds showed a marginally significant reduction in breeding success compared to sham-implanted individuals, with all failures occurring at least 1 week after implant removal. However, prolactin concentrations at implant removal were not significantly different from initial values. We were unable to measure the profile of change in corticosterone during the experiment. However, our results suggest a delayed effect of elevated corticosterone on breeding success rather than an immediate suppression of prolactin concentrations causing premature failure. [ABSTRACT FROM AUTHOR]

Corticosterone has received considerable attention as the principal hormonal mediator of allostasis or physiological stress in wild animals. More recently, it has also been implicated in the regulation of parental care in breeding birds, particularly with respect to individual variation in foraging behavior and provisioning effort. There is also evidence that prolactin can work either inversely or additively with corticosterone to achieve this. Here we test the hypothesis that endogenous corticosterone plays a key physiological role in the control of foraging behavior and parental care, using a combination of exogenous corticosterone treatment, time-depth telemetry, and physiological sampling of female macaroni penguins (Eudyptes chrysolophus) during the brood-guard period of chick rearing, while simultaneously monitoring patterns of prolactin secretion. Plasma corticosterone levels were significantly higher in females given exogenous implants relative to those receiving sham implants. Increased corticosterone levels were associated with significantly higher levels of foraging and diving activity and greater mass gain in implanted females. Elevated plasma corticosterone was also associated with an apparent fitness benefit in the form of increased chick mass. Plasma prolactin levels did not correlate with corticosterone levels at any time, nor was prolactin correlated with any measure of foraging behavior or parental care. Our results provide support for the corticosterone-adaptation hypothesis, which predicts that higher corticosterone levels support increased foraging activity and parental effort. [ABSTRACT FROM AUTHOR]

Abstract: Chronic stress generally inhibits the activity of the reproductive system. Acute stress also is often inhibitory, but the mechanism involved and its persistence of action once animals are no longer exposed to the stressor are poorly understood. We investigated the effect of capture and restraint stress on plasma testosterone (T), luteinizing hormone (LH), and corticosterone (CORT) in free-ranging male rufous-winged sparrows, Peucaea carpalis. Stress decreased plasma T between 10 and 30min after capture and restraint but did not influence plasma LH, the main hormone that controls T secretion, suggesting that stress did not decrease plasma T by inhibiting LH secretion. The stress-induced decrease in plasma T was associated with elevated plasma CORT, but there was no evidence that these effects were functionally related. Plasma stress-induced T was positively related to plasma initial T measured within 2min of capture. This relationship was, however, complex as plasma T decreased proportionally more in response to stress in sparrows with high than low plasma initial T. The relative sensitivity to the same stressor was, therefore, individually variable and this variation was related to initial plasma T. Birds caught and restrained for 30min, and then released on their breeding territory before recapture up to 6h later, maintained depressed plasma T, indicating that the effect of acute stress on this hormone persists after the stressor removal. These studies provide new information on the effects of acute stress on plasma T in free-ranging birds. In particular, they are among the first to characterize the time course and to describe the persistence of these effects. The findings also contribute to identifying factors that are associated with individual differences in plasma hormone levels. [Copyright &y& Elsevier]

Abstract: Physiological mechanisms mediating carryover effects, wherein events or activities occurring in one season, habitat, or life-history stage affect important processes in subsequent life-history stages, are largely unknown. The mechanism most commonly invoked to explain carryover effects from migration centres on the acquisition and utilization of resources (e.g. body mass, or individual ‘condition’). However, other mechanisms are plausible, e.g. trade-offs reflecting conflict or incompatibility between physiological regulatory systems required for different activities or life-history stages (migration vs. reproduction). Here we show that in female black-browed albatrosses (Thalassarche melanophris) the decision to reproduce or to defer reproduction, made prior to their arrival at breeding colonies after long-distance migration, is associated with condition-related (body mass, hematocrit, hemoglobin concentrations) and hormonal (progesterone, testosterone, estrogen-dependent yolk precursors) traits. In contrast, reproductive success showed little association with condition but showed significant associations with the steroidogenic processes underlying follicle development. Specifically, success was determined by reproductive readiness via differences in steroid hormones and hormone-dependent traits. Successful albatrosses were characterized by high progesterone and high estradiol-dependent yolk precursor levels, whereas failed albatrosses had high testosterone and low yolk precursor levels. Results are discussed with reference to migratory carryover effects and how these can differentially affect the physiologies influencing reproductive decisions and reproductive success. [Copyright &y& Elsevier]

Many seasonal breeders adjust the timing of reproduction in response to year-to-year variations in supplementary environmental cues, amongst which ambient temperature is thought to be most influential. However, it is possible that for species such as the great tit (Parus major L.), phenological cues from sprouting vegetation and the consequent abundance of invertebrate prey, although dependent on temperature, may provide supplementary environmental cues per se. This hypothesis was investigated in breeding pairs of great tits kept in outdoor aviaries. In spring, experimental pairs were provided with access to leafing birch branches and caterpillars as a visual food cue, while control pairs were provided with non-leafing branches. Observations were made on the onset of laying and on concentrations of plasma luteinizing hormone (LH) at regular intervals to monitor changes in reproductive function. The onset of egg laying was not advanced by the presence of leafing branches and caterpillars. LH concentrations increased during the course of the study, but phenological cues did not affect plasma LH levels in females and males. Early spring vegetation, such as the leafing of birch branches, and the appearance of caterpillar prey do not appear to play a significant role in fine-tuning the onset of egg laying in great tits. [ABSTRACT FROM AUTHOR]

Summary 1. Annual variation in the timing of avian reproduction is associated with predictive cues related to ambient temperature. Understanding how these cues affect timing, and estimating the genetic variation in sensitivity to these cues, is essential to predict the micro-evolutionary changes in timing which are needed to adapt to climate change. 2. We carried out a 2-year experiment with great tits Parus major of known genetic background, which were kept in pairs in climate-controlled aviaries with simulated natural photoperiod and exposed to a seasonal change in temperature, where the two treatments differed by 4 °C. We recorded the dates of laying the first and last eggs and timing of moult, as well as physiological proxies associated with reproduction: plasma luteinizing hormone (LH), prolactin, and gonadal size at four-weekly intervals. 3. The temperature treatments did not affect first-egg dates, nor gonadal growth or plasma LH and prolactin concentrations. However, birds terminated egg laying, regressed their testes and started their moult earlier at higher temperatures. 4. There were marked family differences in both the start of egg laying, with sisters from early laying maternal families laying early, and in the termination of laying, indicating that there is heritable variation in sensitivity to cues involved in timing. 5. Our experiment, the first to use genetically related individuals in an experimental design with a natural change in photoperiod and biologically realistic temperature differences, thus shows that genetic adaptation in cue sensitivity is possible, essential for species to be able to adapt to a warming world. [ABSTRACT FROM AUTHOR]

Pollutants as endocrine disrupting chemicals (EDCs) are of worldwide concern, due to their detrimental effects on the physiology and behaviour of wildlife. One major source of EDCs is sewage treatment works which serve as foraging grounds for many bird species. However, the risks of consuming EDCs to wild birds at these sites have been largely overlooked. We have previously quantified contamination levels of invertebrates from within filter beds of sewage treatment works and the intake rates of these invertebrates by wild European starlings feeding at these sites. Our work to date has shown that environmentally-relevant EDC exposure results in reduced immunocompetence, increased song behaviour and neural development in adult starlings. In the present study, we hypothesized that nestling starlings exposed to similar EDC levels from contaminated prey, from parental feeding at sewage treatment works, would show altered growth and physiology. Our findings show that EDC exposure results in reduced growth and immunocompetence in nestling birds. However, there was no effect on corticosterone or haematocrit levels. As growth rates and immunocompetence are likely to be related to survival rates, we suggest that these exposure levels are likely to adversely affect recruitment into the adult population. Synthesis and applications. These results highlight the potential transfer of EDCs between semi-aquatic or aquatic organisms and terrestrial organisms up the food chain at sites of sewage treatment works, broadening the scope of potential routes of exposure. Our findings suggest that birds foraging in these sites are at risk of physiological manipulation from EDCs and reduced body condition as a result. We advocate further work to evaluate the potential for EDCs from sewage treatment works to bioaccumulate and modulate the condition of wild organisms feeding on contaminated prey. Our results suggest that changes in management practices of sewage treatment works are required. We support measures to reduce EDC levels in sewage treatment works and that these sites should be designed to exclude or deter wildlife from foraging on contaminated prey. The management implications of such policy - oriented steps will ensure the health of wildlife foraging at sewage treatment works. [ABSTRACT FROM AUTHOR]

Recent changes in the seasonal timing (phenology) of familiar biological events have been one of the most conspicuous signs of climate change. However, the lack of a standardized approach to analysing change has hampered assessment of consistency in such changes among different taxa and trophic levels and across freshwater, terrestrial and marine environments. We present a standardized assessment of 25 532 rates of phenological change for 726 UK terrestrial, freshwater and marine taxa. The majority of spring and summer events have advanced, and more rapidly than previously documented. Such consistency is indicative of shared large scale drivers. Furthermore, average rates of change have accelerated in a way that is consistent with observed warming trends. Less coherent patterns in some groups of organisms point to the agency of more local scale processes and multiple drivers. For the first time we show a broad scale signal of differential phenological change among trophic levels; across environments advances in timing were slowest for secondary consumers, thus heightening the potential risk of temporal mismatch in key trophic interactions. If current patterns and rates of phenological change are indicative of future trends, future climate warming may exacerbate trophic mismatching, further disrupting the functioning, persistence and resilience of many ecosystems and having a major impact on ecosystem services. [ABSTRACT FROM AUTHOR]

Where life-history stages overlap, there is the potential for physiological conflicts that might be important in mediating carryover effects. However, our knowledge of the specific physiological mechanisms underlying carryover effects remains rudimentary, and specific examples remain rare. Here we show that female macaroni penguins (Eudyptes clirysolophus) initiate vitellogenesis and yolk formation while at sea during return migrations to breeding colonies; yolk formation takes approximately 16 days, but females lay only 714 days after their return. Once on land, Eudyptes penguins show a unique reproductive pattern of extreme egg size dimorphism in which the smaller, first-laid A-egg is 55%-75% of the size of the larger B-egg. We show that the degree of egg size dimorphism is inversely correlated with time between arrival and laying; that is, females that begin reproductive development well in advance of their return produce more dimorphic eggs. Furthermore, late-arriving females that produce the most dimorphic eggs have lower plasma levels of the yolk precursor vitellogenin on arrival; that is, they show lower reproductive "readiness." These data support the hypothesis that extreme egg size dimorphism in Eudyptes penguins is due to a physiological constraint imposed by a migratory carryover effect and argue against small A-eggs having a specific, adaptive function. [ABSTRACT FROM AUTHOR]

Photorefractoriness is the insensitivity of gonadal development to the stimulatory effects of long photoperiods in birds and to the inhibitory effects of short photoperiods in small mammals. Its molecular mechanism remains unknown. Recently, it has been shown that reciprocal expression of thyroid hormone-activating enzyme [type 2 deiodinase (Dio2)] and -inactivating enzyme [type 3 deiodinase (Dio3)] genes in the mediobasal hypothalamus is critical for photoperiodically induced gonadal growth. Since thyroid hormones are required not only for photoinduction, but also for the induction of photorefractoriness, we examined the expression of these genes in relation to photorefractoriness in birds and mammals. Transfer of birds to long photoperiods induced strong expression of Dio2. This was maintained in tree sparrow when they later became photorefractory, but decreased somewhat in quail. In hamsters, transfer to long photoperiods also induced strong expression of Dio2. High values were not maintained under long photoperiods, and, indeed, expression decreased at the same rate as in animals transferred to short photoperiods. There was no renewed expression of Dio2 associated with testicular growth as animals became refractory to short photoperiods. Expression of Dio3 was high under short photoperiods and low under long photoperiods in all the animals examined, except for the short photoperiod-refractory hamsters. Our present study revealed complex regulation of deiodinase genes in the photoinduction and photorefractory processes in birds and mammals. These gene changes may be involved in the regulation of photorefractoriness, as well as photoinduction. [ABSTRACT FROM AUTHOR]