Artificial rain and cold wind act as stressors to captive molting and non-molting European starlings (Sturnus vulgaris)

Free-roaming animals continually cope with changes in their environment. One of the most unpredictable environmental phenomena is weather. Being able to respond to weather appropriately is crucial as it can be a threat to survival. The stress response, consisting of increases in heart rate and release of glucocorticoids, is an important mechanism by which animals cope with stressors. This study examined behavioral, heart rate, and corticosterone responses of captive European starlings (Sturnus vulgaris) to two aspects of weather mimicked under controlled conditions, a subtle (3 °C) decrease in temperature and a short, mild bout of rain. Both decreased temperature and exposure to rain elicited increases in heart rate and corticosterone in non-molting starlings. Molt is an important life history stage in birds that affects feather cover and may require a different response to weather-related stressors. We repeated the experiment in molting starlings and found increases in heart rate in response to rain and cold wind. However, the hypothalamic–pituitary–adrenal (HPA)-axis was suppressed during molt, as molting starlings did not increase corticosterone release in response to either stimulus. These data suggest these stimuli induce increased allostatic load in starlings, and that animals may adjust their response depending on the life-history stage.     

Combined effects of molt and chronic stress on heart rate, heart rate variability, and glucocorticoid physiology in European Starlings

Molt is an important life-history stage in avian species, but little is known about the effects of chronic stress during this period. Three weeks after the onset of molt, captive European starlings (Sturnus vulgaris) were exposed to 18 days of chronic stress, induced with four 30-minute randomized stressors presented daily. Birds showed no chronic-stress-induced changes in heart rate or heart rate variability when measured either during the middle of the day or at night. These data suggest that chronic stress did not alter the balance between sympathetic and parasympathetic nervous system regulation of cardiovascular function, which contrasts with data from an earlier study indicating that chronic stress profoundly alters cardiovascular function in non-molting starlings. Additionally, there was a significant increase in restraint-induced corticosterone secretion the first week of chronic stress that subsequently returned to pre-chronic-stress levels by the second week of exposure. The attenuated corticosterone response again contrasts with data from non-molting starlings that showed significant decreases in corticosterone responses. Consequently, the resistance to cardiovascular and corticosterone changes indicates that the physiological changes induced by chronic stress are greatly attenuated in molting birds. Overall, the data suggest that molt requires a degree of physiological stability that must be protected, so that if a bird is exposed to chronic stress during this life-history stage, molt takes priority.     

Captive European starlings (Sturnus vulgaris) in breeding condition show an increased cardiovascular stress response to intruders

European starlings (Sturnus vulgaris) alter their physiology and behavior between seasons, becoming territorial during the spring/summer and flocking during the fall/winter. We used captive male starlings in breeding (photostimulated to 18L : 6D) and nonbreeding (11L : 13D) conditions to determine whether changing physiology and behavior alters their reaction to crowding. One or five intruders entered a resident's cage without human disturbance. A subcutaneous heart rate transmitter recorded cardiovascular output in residents. Corticosterone and testosterone were measured in plasma samples taken before and after the intrusion. While corticosterone concentrations did not change, heart rate changed significantly, indicating that these responses can be regulated independently. Long-day birds showed a significantly elevated heart rate response to the single-bird intrusion compared to short-day birds. Whereas five intruders elicited an identical peak response in both groups, long-day birds also demonstrated an equivalent response to one intruder. In addition, one intruder induced longer elevation in heart rate for long-day birds. Male starlings in breeding condition, therefore, demonstrate an increased sensitivity to additional conspecifics. This seasonal shift in response suggests that a higher tolerance for intrusion (i.e., considering a nearby starling as less stressful) may facilitate flocking behavior, while a lower tolerance may aid in territoriality.     

Effects of predictable and unpredictable food restriction on the stress response in molting and non-molting European starlings (Sturnus vulgaris)

This study tested whether an ethologically relevant stressor, a three-week period of food restriction where food was unavailable for four hours a day, caused chronic stress in molting and non-molting captive European starlings. Although all birds increased weight during the Food Restriction period, only non-molting birds increased food intake. Morning baseline heart rates increased during the Food Restriction period and all birds showed a decrease in heart rate when food was absent from the cage. In non-molting birds, there were no differences in either baseline or stress-induced corticosterone (CORT) concentrations, whereas molting birds showed attenuated baseline CORT, stress-induced CORT, and fecal glucocorticoid metabolite levels over the Food Restriction period. Although several parameters, such as increased morning heart rate, are consistent with chronic stress, the majority of these data suggest that restricting food availability is not chronically stressful. Furthermore, making the timing of food removal less predictable by randomizing when food was removed during the day did not enhance any of the above responses, but did alter the frequency of maintenance and feeding behaviors. In conclusion, starlings appear resistant to developing symptoms of chronic stress from repeated food restriction.     

Behavioral, physiological, and endocrine responses of starlings to acute increases in density

One potential stressor to vertebrates both in the wild and in captivity is the presence of numerous individuals in a confined space. To examine the effects of increased conspecific density in birds, we simultaneously measured cardiac, behavioral, and endocrine responses of European starlings (Sturnus vulgaris) to acute crowding. A cage containing a resident bird was outfitted with a trap door that allowed for the introduction of intruder birds (one, three, or five birds) without human interference. The resident bird was implanted with a subcutaneous heart rate (HR) transmitter, behavior was videotaped through a two-way mirror, and blood samples were taken at the end of each treatment to determine plasma corticosterone (CORT) concentrations. Resident starlings significantly increased both general activity and aggressive behaviors while decreasing preening following the initiation of elevated conspecific density. Intruder starlings increased feeding, drinking, and aggressive pecking rates, but postintrusion feeding rates decreased as intruder number increased. Preening decreased in both residents and intruders following the intrusion. HR increased in the resident starlings at the time of intruder introduction, with an increase in the magnitude of this response directly correlating with increasing intruder number. The CORT response to increased density was dependent on social role (resident or intruder), since increasing density did not alter CORT levels in resident birds, but resulted in elevated CORT 30 min following the five-intruder introduction in the intruder birds. Together, these data suggest that increased conspecific density is a significant acute stressor in starlings which is capable of inducing aggression in both residents and intruders. Furthermore, it elicits different responses from different physiological and behavioral systems, and behavioral responses such as feeding and general activity may be density-dependent. The data specifically illustrate that cardiac and behavioral activation can be independent of CORT release, and the CORT response of starlings to increased conspecific density is dependent on social role and degree of the increase in density.     

Exposure to chronic stress downregulates corticosterone responses to acute stressors

We used captive European starlings (Sturnus vulgaris) to test whether corticosterone responses differed in birds held under normal laboratory conditions or conditions of chronic stress. Surprisingly, both basal corticosterone concentrations and corticosterone responses to acute stress were significantly reduced when birds were chronically stressed. To determine the mechanism underlying this reduced response, animals under both conditions were injected with lactated Ringer's solution (control), adrenocorticotropin (ACTH), arginine vasotocin (AVT), or dexamethasone (DEX). ACTH increased corticosterone concentrations above stress-induced levels in both cases, although maximum responses were lower in chronically stressed birds. AVT did not augment the corticosterone response under nonchronically stressed conditions, but it did under chronically stressed conditions. DEX reduced maximal corticosterone concentrations in both cases. Neither ovine nor rat corticotropin-releasing factor (CRF) altered normal stress responses. These data indicate that changes in responsiveness of the hypothalamic-pituitary-adrenal axis to ACTH and AVT serve to downregulate corticosterone responses during chronic stress. Furthermore, these data lead to the following hypothesis: ACTH output from the pituitary limits maximum corticosterone concentrations under normal conditions, but reduced AVT release from the hypothalamus regulates lower corticosterone concentrations under chronic stress conditions.     

Behavioural and hormonal effects of social isolation and neophobia in a gregarious bird species, the European starling (Sturnus vulgaris)

Separating gregarious individuals from their group members often results in behavioural and physiological changes, like increased levels of corticosterone. Testosterone and corticosterone, in particular, have been implicated in the response of mammals to novelty. Data in birds are, however, rare. The presence or absence of group members may also influence an individual's response to novel stimuli. We assessed the behaviour and hormonal response of European starlings (Sturnus vulgaris) to a novel object in two different situations and seasons: each starling was tested when separated and when in contact with its group members in May/June (breeding season) and again in September/October (non-breeding season). Starlings are gregarious throughout the year, but as foraging flocks are small during the breeding season and large during the non-breeding season, we assumed that non-breeding starlings would be more affected by social isolation. Overall, starlings had higher levels of corticosterone, lost more body mass, and were more active when they were separated from their group. Isolated individuals, however, did not show a greater neophobic response than individuals in the presence of their group members in either season. Circulating levels of testosterone and corticosterone were higher after a test with novel object than after a test with only the familiar feeding dish in both sexes and seasons. However, control tests for handling effects confirmed only the increase in testosterone. Our study shows that social isolation is stressful for unrelated and unpaired members of a wild flocking bird species and demonstrates that novelty can lead to a rise in testosterone in birds.     

Melatonin,immunity and cost of reproductive state in male European starlings.

The effects of reproductive condition and exogenous melatonin on immune function were investigated in castrated European starlings, Sturnus vulgaris. Photorefractory and photostimulated starlings exposed to long days were implanted with melatonin or with blank capsules. Photostimulated starlings with blank capsules exhibited reduced splenocyte proliferation in response to the T-cell mitogen, concanavalin A, compared with the other long-day birds. Exogenous melatonin prevented the suppression of immune function by photostimulation. Photorefractory starlings, with or without melatonin implants, exhibited enhanced immune function compared with photostimulated starlings implanted with blanks. This enhancement was not mediated by endogenous melatonin, but appeared to be related to changes in reproductive state. In addition to the traditional costs of reproduction in birds (e.g. raising of young), there may be a cost of the reproductive state of starlings (i.e. whether they are photorefractory or photostimulated). These data are, we believe, the first to indicate a direct effect of reproductive state on immune function that is independent of both photoperiod (i.e., changes in the duration of melatonin secretion) and gonadal steroids.     

Corticosterone and insulin interact to regulate glucose and triglyceride levels during stress in a bird

Captive European starlings (Sturnus vulgaris) were exposed to the stress of handling and restraint while corticosterone, glucose, and triglyceride concentrations were monitored in blood plasma. In saline-injected controls, basal samples were taken within 3 min of disturbance with subsequent samples taken at 40, 70, and 150 min. This was repeated at two times during the daily cycle (day and night) on two different photoperiods: short and long days. During both photoperiods, corticosterone concentrations approximately tripled (compared with a sixfold increase in free-living starlings) and triglyceride concentrations decreased 25-45% in response to stress at both times of the day, whereas an approximately 25% stress-induced hyperglycemia occurred only at night. Exogenous corticosterone (200 microg), 1.0 or 4.0 IU/kg of insulin, or a combination of corticosterone with each insulin dose was then separately administered to alter the above responses. Insulin did not affect corticosterone or triglyceride concentrations but resulted in a dose-dependent hypoglycemia of 10-40%. Injected corticosterone resulted in supraphysiological corticosterone concentrations (three- to fivefold higher than normal), yet it did not affect the already altered plasma glucose or triglyceride concentrations. This suggests that glucose output and triglyceride decreases were already maximal in response to handling and restraint. However, the low glucose concentrations resulting from exogenous insulin returned to basal quicker with exogenous corticosterone but only during the day. No response to either hormone showed photoperiodic differences. These data suggest that corticosterone's role in metabolism changes to meet varying energetic demands throughout the day.     

Increased energy expenditure but decreased stress responsiveness during molt

Baseline and stress-induced corticosterone (CORT), heart rate (fH), and energy expenditure were measured in eight captive European starlings Sturnus vulgaris during and following a prebasic molt. The fH and oxygen consumption (V O2 ) were measured simultaneously across a range of heart rates, and energy expenditure (kJ/d) was then calculated from data. Energy expenditure and fH were strongly and positively correlated in each individual. Baseline fH and energy expenditure were significantly higher during molt. Molting starlings expended 32% more energy over 24 h than nonmolting birds, with the most significant increase (60%) occurring at night, indicating a substantial energetic cost to molt. Furthermore, the cardiac and metabolic responses to stress during molt were different than during nonmolt. Birds were subjected to four different 30-min acute stressors. The fH and CORT responses to these stressors were generally lower during molt. Although restraint caused a 64% increase in daily energy expenditure during nonmolt, no other stressor caused a significant increase in energy expenditure. Overall, our data suggest that molt is not only energetically expensive but that it also alters multiple stress response pathways. Furthermore, most acute stressors do not appear to require a significant increase in energy expenditure.     

A comparison of the annual cycles in testicular size and moult in captive European starlings Sturnus vulgaris during their first and second years

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.     

The LH-RH system of the male European starling: photoperiod induces changes to a possible multifunctional peptide system

In many birds reproduction is triggered by long daylengths but, paradoxically, continued exposure to long days leads to photorefractoriness and a complete shut down of the reproductive system. As these effects are thought to be mediated through the secretion of LH-RH, immunocytochemical techniques were used to investigate changes in the LH-RH system when European starlings were exposed to different photoperiods. Starlings exposed to 11L:13D and with mature testes show strong immunostaining both of LH-RH perikarya and fibers. Photosensitive short-day (8L:16D) starlings with undeveloped testes show an almost identical distribution of strongly immunoreactive perikarya but with less dense fibre staining. However, long-day (18L:6D) photorefractory starlings with fully regressed testes, show a profound reduction in LH-RH immunostaining. Perikarya have the same distribution but show a much reduced intensity of staining and fibers had almost entirely disappeared from all regions of the brain. Preliminary observations on the ultrastructure of immunocytochemically identified LH-RH neurones are also reported.     

Effects of Birds Ingesting Heterodera rostochiensis Cysts on Viability of Eggs and Larvae

Rate of passage through the digestive systems and effects of ingestion on viability of contents of cysts of Heterodera rostochiensis were determined in feeding trials with pigeons, thrushes, starlings, cowbirds, sparrows, and quails. Depending upon species of birds, 12-82% of the cysts ingested passed through the digestive system within 0.5 h. Pigeons required 6 h for complete evacuation. All other birds cmnpletely evacuated ingested cysts from their digestive systems within 3 h. Contents of cysts were nonviable if they were retained in the digestive system of starlings for more than 1.5 h, pigeons more than 1 h, or other birds more than 0.5 h. Cyst contents were nonviable if they remained in contact with excreta from cowbirds or quails for 4 h, thrushes for 96 h, or other species for 72 h after passage. Viability of contents of cysts was inversely related to exposure to excreta-filtrate concentration. Larvae failed to emerge from cysts that were exposed to a 25% concentration of excreta filtrate from starlings, 50% concentration from pigeons or thrushes, or 100% concentration of excreta filtrates from each of the other species. Cysts that were subjected to 44 C (avg. body temperature of cowbirds) for more than 3.5 h were nonviable. Cysts that passed through birds and collected with excrement on polyethylene or soil produced no infective larvae on potato.     

Juveniles exposed to embryonic corticosterone have enhanced flight performance

Exposure to maternally derived glucocorticoids during embryonic development impacts offspring phenotype. Although many of these effects appear to be transiently 'negative', embryonic exposure to maternally derived stress hormones is hypothesized to induce preparative responses that increase survival prospects for offspring in low-quality environments; however, little is known about how maternal stress influences longer-term survival-related performance traits in free-living individuals. Using an experimental elevation of yolk corticosterone (embryonic signal of low maternal quality), we examined potential impacts of embryonic exposure to maternally derived stress on flight performance, wing loading, muscle morphology and muscle physiology in juvenile European starlings (Sturnus vulgaris). Here we report that fledglings exposed to experimentally increased corticosterone in ovo performed better during flight performance trials than control fledglings. Consistent with differences in performance, individuals exposed to elevated embryonic corticosterone fledged with lower wing loading and had heavier and more functionally mature flight muscles compared with control fledglings. Our results indicate that the positive effects on a survival-related trait in response to embryonic exposure to maternally derived stress hormones may balance some of the associated negative developmental costs that have recently been reported. Moreover, if embryonic experience is a good predictor of the quality or risk of future environments, a preparative phenotype associated with exposure to apparently negative stimuli during development may be adaptive.     

Effect of initial core temperature on hyperthermic hyperventilation during prolonged submaximal exercise in the heat

We investigated whether a core temperature threshold for hyperthermic hyperventilation is seen during prolonged submaximal exercise in the heat when core temperature before the exercise is reduced and whether the evoked hyperventilatory response is affected by altering the initial core temperature. Ten male subjects performed three exercise trials at 50% of peak oxygen uptake in the heat (37°C and 50% relative humidity) after altering their initial esophageal temperature (T(es)). Initial T(es) was manipulated by immersion for 25 min in water at 18°C (Precooling), 35°C (Control), or 40°C (Preheating). T(es) after the water immersion was significantly higher in the Preheating trial (37.5 ± 0.3°C) and lower in the Precooling trial (36.1 ± 0.3°C) than in the Control trial (36.9 ± 0.3°C). In the Precooling trial, minute ventilation (Ve) showed little change until T(es) reached 37.1 ± 0.4°C. Above this core temperature threshold, Ve increased linearly in proportion to increasing T(es). In the Control trial, Ve increased as T(es) increased from 37.0°C to 38.6°C after the onset of exercise. In the Preheating trial, Ve increased from the initially elevated levels of T(es) (from 37.6 to 38.6°C) and Ve. The sensitivity of Ve to increasing T(es) above the threshold for hyperventilation (the slope of the T(es)-Ve relation) did not significantly vary across trials (Precooling trial = 10.6 ± 5.9, Control trial = 8.7 ± 5.1, and Preheating trial = 9.2 ± 6.9 L·min(-1)·°C(-1)). These results suggest that during prolonged submaximal exercise at a constant workload in humans, there is a clear core temperature threshold for hyperthermic hyperventilation and that the evoked hyperventilatory response is unaffected by altering initial core temperature.     

Exogenous and endogenous corticosterone alter feather quality

We investigated how exogenous and endogenous glucocorticoids affect feather replacement in European starlings (Sturnus vulgaris) after approximately 56% of flight feathers were removed. We hypothesized that corticosterone would retard feather regrowth and decrease feather quality. After feather regrowth began, birds were treated with exogenous corticosterone or sham implants, or endogenous corticosterone by applying psychological or physical (food restriction) stressors. Exogenous corticosterone had no impact on feather length and vane area, but rectrices were lighter than controls. Exogenous corticosterone also decreased inter-barb distance for all feathers and increased barbule number for secondaries and rectrices. Although exogenous corticosterone had no affect on rachis tensile strength and stiffness, barbicel hooking strength was reduced. Finally, exogenous corticosterone did not alter the ability of Bacillus licheniformis to degrade feathers or affect the number of feathers that failed to regrow. In contrast, endogenous corticosterone via food restriction resulted in greater inter-barb distances in primaries and secondaries, and acute and chronic stress resulted in greater inter-barb distances in rectrices. Food-restricted birds had significantly fewer barbules in primaries than chronic stress birds and weaker feathers compared to controls. We conclude that, although exogenous and endogenous corticosterone had slightly different effects, some flight feathers grown in the presence of high circulating corticosterone are lighter, potentially weaker, and with altered feather micro-structure.     

Avian influenza A virus susceptibility,infection, transmission,and antibody kinetics in European starlings

Avian influenza A viruses (IAVs) pose risks to public, agricultural, and wildlife health. Bridge hosts are spillover hosts that share habitat with both maintenance hosts (e.g., mallards) and target hosts (e.g., poultry). We conducted a comprehensive assessment of European starlings (Sturnus vulgaris), a common visitor to both urban and agricultural environments, to assess whether this species might act as a potential maintenance or bridge host for IAVs. First, we experimentally inoculated starlings with a wild bird IAV to investigate susceptibility and replication kinetics. Next, we evaluated whether IAV might spill over to starlings from sharing resources with a widespread IAV reservoir host. We accomplished this using a specially designed transmission cage to simulate natural environmental transmission by exposing starlings to water shared with IAV-infected mallards (Anas platyrhynchos). We then conducted a contact study to assess intraspecies transmission between starlings. In the initial experimental infection study, all inoculated starlings shed viral RNA and seroconverted. All starlings in the transmission study became infected and shed RNA at similar levels. All but one of these birds seroconverted, but detectable antibodies were relatively transient, falling to negative levels in a majority of birds by 59 days post contact. None of the contact starlings in the intraspecies transmission experiment became infected. In summary, we demonstrated that starlings may have the potential to act as IAV bridge hosts if they share water with IAV-infected waterfowl. However, starlings are unlikely to act as maintenance hosts due to limited, if any, intraspecies transmission. In addition, starlings have a relatively brief antibody response which should be considered when interpreting serology from field samples. Further study is needed to evaluate the potential for transmission from starlings to poultry, a possibility enhanced by starling’s behavioral trait of forming very large flocks which can descend on poultry facilities when natural resources are scarce.     

Effects of arginine vasotocin (AVT) on the behavioral, cardiovascular, and corticosterone responses of starlings (Sturnus vulgaris) to crowding

Previous studies in European starlings have concluded that conspecific crowding can be a significant stressor that is capable of simultaneously altering behavior, heart rate, and corticosterone (CORT) concentrations. It was hypothesized that the peptide hormone arginine vasotocin (AVT) has a role in the regulation of these three types of responses to crowding. Four male and four female resident starlings were submitted to nine combinations of 3 crowding treatments (0, 1, or 5 intruder starlings) and 3 subcutaneous injections (1, 4 microg AVT, and saline control). Resident starlings were given a treatment injection, their heart rate and behavior were monitored for 30 min, 0, 1, or 5 intruder Starlings were allowed to enter the residents cage, and HR and behavior were monitored for another 30 min. Blood samples were taken before and after all treatments to assess CORT concentrations. Exogenous AVT decreased the frequency of maintenance behaviors (feeding, drinking, preening, and beak wiping), as well as activity in resident starlings. Although aggressive behaviors upright posture, head feather expansion, and pecking) increased during crowding, these increases were significantly attenuated by AVT. Heart rate was significantly lower during these behavioral effects, and the CORT data indicate that the cardiovascular and behavioral effects are not dependent on significant increases in CORT. These data support the hypothesis that AVT's attenuation of general behavior and crowding induced aggression are modulated by a cardiovascular mechanism.     

A comparative study of avian auditory brainstem responses: correlations with phylogeny and vocal complexity, and seasonal effects

We conducted a comparative study of the peripheral auditory system in six avian species (downy woodpeckers, Carolina chickadees, tufted titmice, white-breasted nuthatches, house sparrows, and European starlings). These species differ in the complexity and frequency characteristics of their vocal repertoires. Physiological measures of hearing were collected on anesthetized birds using the auditory brainstem response to broadband click stimuli. If auditory brainstem response patterns are phylogenetically conserved, we predicted woodpeckers, sparrows, and starlings to be outliers relative to the other species, because woodpeckers are in a different Order (Piciformes) and, within the Order Passeriformes, sparrows and starlings are in different Superfamilies than the nuthatches, chickadees, and titmice. However, nuthatches and woodpeckers have the simplest vocal repertoires at the lowest frequencies of these six species. If auditory brainstem responses correlate with vocal complexity, therefore, we would predict nuthatches and woodpeckers to be outliers relative to the other four species. Our results indicate that auditory brainstem responses measures in the spring broadly correlated with both vocal complexity and, in some cases, phylogeny. However, these auditory brainstem response patterns shift from spring to winter due to species-specific seasonal changes. These seasonal changes suggest plasticity at the auditory periphery in adult birds.     

Circannual variations in plasma luteinizing hormone levels in castrated male European starlings (Sturnus vulgaris)

Intact and castrated male European starlings were held for about 2 years in a constant 12-hr photoperiod and constant temperature conditions. At 1- to 2-month intervals, testicular width was measured by laparotomy, and blood samples were taken for analysis of plasma luteinizing hormone (LH). Most of the control birds went through at least one circannual cycle of testicular width and plasma LH concentration. In the castrates, a similar proportion of birds went through circannual LH cycles with periods indistinguishable from those of the controls. It is concluded that the testes and their hormones are not essential components of the mechanism that generates circannual gonadal cycles in male European starlings.