The Photoperiodic Entrainment and Induction of Reproductive Rhythms in Male Blackheaded Buntings (Emberiza Melanocephala)

Groups of photosensitive, unstimulated or stimulated, male blackheaded buntings were subjected to photoregimes of 15 hr of green light of three intensities and 9 hr of dark per day. In some groups green light was interrupted with 90 min of bright fluorescent light at different times in the subjective day. While gonads did not develop or regressed in some groups, birds in others behaved as if exposed to long daylengths. The results besides suggesting the involvement of endogenous circadian rhythm during initiation and maintenance of gonadal growth indicate that the reproductive rhythms are entrained and induced by environmental photoperiod.     

Wavelength Dependency of Light-Induced Effects on Photoperiodic Clock in the Migratory Blackheaded Bunting (Emberiza melanocephala)

The effects of light wavelength on photoperiodic clock were determined in the migratory male blackheaded bunting (Emberiza melanocephala). We constructed an action spectrum for photoperiodic induction (body fattening, gain in body mass, and gonadal recrudescence) by exposing birds for 4.5 weeks to 13 h light per day (L:D = 13:11 h) of white (control), blue (450 nm), or red (640 nm) color at irradiances ranging from 0.028 to 1.4 W m^?2. The threshold light irradiance for photoinduction was about 10-fold higher for blue, compared to red and white light. Phase-dependent effects of light wavelength on the photoperiodic clock were further examined in the next two sets of skeleton photoperiods (SKPs). In the first set of SKPs, birds were exposed for four weeks to asymmetrical light periods (L:D:L:D = 6:6:1:11 h) at 0.25 ± 0.01 W m^?2; two light periods applied were of the same (450 nm: blue:blue, B:B; 640 nm, red:red, R:R) or different (blue:red, B:R or red:blue, R:B) wavelengths, or of white:white (W:W, controls). Photoperiodic induction occurred under R:R and B:R, but not under B:B and R:B light conditions; the W:W condition induced an intermediate response. The second set of SKPs used symmetrical light periods (L:D:L:D = 1:11:1:11 h), and measured effects also on the activity rhythm. Birds were first exposed to one of the four SKPs (R:R, B:B, R:B, or B:R) for three weeks, subsequently were released into dim constant light (LL_dim; ?0.01 W m^?2, the night light used in an L:D cycle) for two weeks, and then were returned to respective SKPs for another three weeks. Activity was greater in the R:R compared to B:B, and in B:R compared to R:B light condition. Zugunruhe (intense nighttime activity, indicating migratory restlessness in a caged situation) developed under the R:R and B:R, but not the B:B and R:B, light condition. Under LL_dim, all birds free-ran with a period >24 h, the Zugunruhe had a circadian period longer than the daytime activity, and the re-entrainment to SKPs was influenced by the position of light periods relative to circadian phase of the activity rhythm. Photoperiodic induction at the end of 8 weeks was found in the R:R and B:R, but not in B:B, light conditions; in the R:B condition only one bird had initiated testes. Taken together, these results suggest that in the blackheaded bunting, the circadian photoperiodic clock is differentially responsive to light wavelengths; this responsiveness is phase-dependent, and the development of Zugunruhe reflects a true circadian function. Wavelength-dependent response of the photoperiodic clock could be part of an adaptive strategy in evolution of the seasonality in reproduction and migration among photoperiodic species under wild conditions.     

Part-Time Mate Guarding Affects Paternity in Male Reed Buntings (Emberiza schoeniclus)

Male mate guarding by close following has been reported in many socially monogamous bird species and is generally believed to function as a paternity guard. Many aspects of the dynamics and effectiveness of this behavior are still however poorly understood. Here, we describe the temporal variation in mate guarding behavior in male reed buntings (Emberiza schoeniclus) with a particular focus on how males allocate their mating effort between mate guarding and extrapair mating in a context of intense sperm competition. In our highly synchronous study population most males have to balance the simultaneous and mutually exclusive demands of mate guarding and seeking extrapair copulations (EPCs). We found that males frequently switched between guarding their mates and performing intrusions to neighboring territories. Both activities seemed to have significant fitness payoffs, as male mate guarding effort had a positive effect on paternity, and a large fraction of extrapair fertilizations occurred during the days when the sire guarded its own female. The reed bunting is thus an example of how discontinuous or part‐time mate guarding can still be effective in securing paternity. Female reed buntings were not particularly active in initiating EPCs as they never were observed performing extraterritorial forays. We argue that the absence of female‐initiated EPCs is a prerequisite for males to trade mate guarding against seeking EPCs. Otherwise, if females circumvent male mate guarding by timing their EPCs to periods of male absence, males should guard their mates almost continuously or rely on alternative paternity guards.     

太平鸟和灰头wu的代谢产热特征及体温调节

采用封闭式流体压力呼吸仪,测定了在环境温度5、10、15、16、18、20、22、25、26、28和30 ℃时,捕自黑龙江省齐齐哈尔市的太平鸟(Bombycilla garrulus,1♀、7♂)和灰头wu(Emberiza spodocephala,8♂)的代谢率(MR),计算每个温度点的热传导?、MR和C的体重预期值。结果显示：在环境温度(Ta)为5～30 ℃时,太平鸟的体温(Tb)基本维持恒定,平均Tb为(38.20±0.05) ℃;热中性区(TNZ)为18～27 ℃,基础代谢率(BMR)为(2.33±0.47)mL O.2/(g·h);Ta在5～18 ℃,MR与Ta呈负相关,回归方程为：MR=5.46(±0.19)-0.17(±0.01)Ta;Ta在5～22 ℃,C最低且基本保持恒定,平均为(0.13±0.00)mL O.2／(g·h·℃)。5～30 ℃时,灰头wu Tb也基本维持恒定,平均Tb为(38.25±0.05) ℃; TNZ为20～26 ℃,BMR为(4.75±0.18)mL O.2／(g·h),最低C为(0.26±0.00)mL O.2／(g·h· ℃);在5～20 ℃时,MR与Ta的回归方程为：MR=9.54(±0.52)-0.23(±0.03)Ta。两种鸟具有BMR和Tb较高、下临界温度较低和TNZ较宽、热传导稍高的北方地区小型鸟类的代谢特点,通过自身的物理和化学调节(代谢产热)以更好地适应其生存环境。     

太平鸟和灰头鹀的代谢产热特征及体温调节

采用封闭式流体压力呼吸仪,测定了在环境温度5、10、15、16、18、20、22、25、26、28和30℃时,捕自黑龙江省齐齐哈尔市的太平鸟(Bombycillagarrulus,1♀、7♂)和灰头(Emberizaspodocephala,8♂)的代谢率(MR),计算每个温度点的热传导(C)、MR和C的体重预期值。结果显示:在环境温度(Ta)为5~30℃时,太平鸟的体温(Tb)基本维持恒定,平均Tb为(38·20±0·05)℃;热中性区(TNZ)为18~27℃,基础代谢率(BMR)为(2·33±0·47)mLO2/(g·h);Ta在5~18℃,MR与Ta呈负相关,回归方程为:MR=5·46(±0·19)-0·17(±0·01)Ta;Ta在5~22℃,C最低且基本保持恒定,平均为(0·13±0·00)mLO2/(g·h·℃)。5~30℃时,灰头Tb也基本维持恒定,平均Tb为(38·25±0·05)℃;TNZ为20~26℃,BMR为(4·75±0·18)mLO2/(g·h),最低C为(0·26±0·00)mLO2/(g·h·℃);在5~20℃时,MR与Ta的回归方程为:MR=9·54(±0·52)-0·23(±0·03)Ta。两种鸟具有BMR和Tb较高、下临界温度较低和TNZ较宽、热传导稍高的北方地区小型鸟类的代谢特点,通过自身的物理和化学调节(代谢产热)以更好地适应其生存环境。     

Different singing styles in mated and unmated Reed Buntings Emberiza schoeniclus

Ewin described two singing styles, "rapid" and "slow", in the Reed Bunting Emberiza schoeniclus. My research has demonstrated that the prevalence of song styles is related to male mating status. Songs of mated and unmated males differed in the interval structure, number of syllables per song, number of frequency modulations per syllable and number of introductory syllables. Since both singing styles are uttered at the same intensity, it is not likely that they are parts of a continuous range and it is justified to treat them as two different song categories. After losing their mate, males sang like unpaired males and therefore a learning effect can be excluded. Both singing styles were directed to females, the slow one probably only to the mate. The use of these singing styles as a means of territory or mate defence cannot be excluded, but, if so used, it would seem to be of minor significance.     

Lifetime patterns of pairing success in male Ortolan Buntings Emberiza hortulana

Analyses of lifetime fitness in birds are typically based on estimates of breeding success, in particular the number of offspring fledged. Small and isolated bird populations often have a male‐skewed adult sex ratio, so that male lifetime productivity depends to a large degree on pairing success, but few studies have focused on patterns of lifetime pairing success. The Norwegian population of Ortolan Buntings Emberiza hortulana is strongly male‐skewed, such that in any year about half of all males are unpaired. Pairing success of first‐year males (16–44%) was significantly lower than for older males (52–89%). Lifetime pairing success was correlated with lifespan and was strongly skewed, with a majority of males being paired only once or never, and only 11% paired three or more times despite a stable lifetime annual survival rate of 63%. Males that were paired in one year were more likely to be paired the next year than males that were unpaired in the previous year. The shortage of females caused even the older males to have a substantial probability of becoming unpaired, and 49% of long‐lived males (known as adults for at least 4 years) were unpaired after years in which they were paired. Pairing success in the Ortolan Bunting therefore follows similar age‐related and lifetime patterns in breeding success documented in other species. However, even the older males ran a high risk of not being paired, contrasting with earlier distinctions between pre‐breeding and breeding lifespans. I discuss the importance of knowledge of pairing success for the management of endangered and declining populations.     

Biochronometry of photoperiodically induced fat deposition in a migratory finch, the Blackheaded bunting (Emberiza melanocephala) (Aves)

This investigation attempts to identify the mechanism(s) involved in the fat deposition in a photoperiodic migratory species, the Blackheaded bunting ( Emberiza melanocephala ). Groups of photosensitive male buntings were exposed to resonance, interrupted night, and ultra-short-day light cycles for 35, 42 and 75 days, respectively. Birds were weighed at the beginning and at the end of the experiments. Those exposed to ultra-short-day light cycles were also weighed at critical intervals during the treatment period. Our results indicate that: (a) a light-sensitive rhythm with a period of about 24 hr is involved in the photoperiodic induction of premigratory fattening and weight gain in Blackheaded buntings: (b) buntings possess a bimodal pattern 'of sensitivity to photoperiods that induce fattening, and (c) this endogenous circadian rhythm can be entrained by an ultrashort photophase of 3 h if the latter is coupled with scotophases of specific duration.     

The breeding ecology of Reed Buntings Emberiza schoeniclus in farmland and wetland habitats in lowland England

We studied the nesting and feeding ecology of Reed Buntings Emberiza schoeniclus breeding on farmland and wetland habitats along the Trent Valley in Nottinghamshire, England. Rank and emergent vegetation accounted for most nests and most foraging by provisioning adults. Caterpillars and spiders accounted for 70% of chick invertebrate prey and all broods were fed cereal grain or other vegetable matter. Variation in the abundance of key invertebrate prey across habitats accounted for the foraging preferences of adult buntings. Depredation was the main cause of nest failure, and survival of nests at the egg stage was positively related to the extent of nest concealment. A measure of total brood biomass was positively related to the abundance of key invertebrate prey within 100 m of nests. Rank and emergent vegetation provided Reed Buntings with greater nest concealment and a richer source of invertebrate prey than agricultural habitats such as set-aside, cereals and oilseed rape. The provision of rank and emergent vegetation on farmland is likely to increase the nesting opportunities and productivity of Reed Buntings in agricultural landscapes.     

Entrainment to Light of Circadian Activity Rhythms in Tench (Tinca tinca)

The present article analyzes locomotor activity rhythms in Tinca tinca. To that end, three different experiments were conducted on 24 animals (20 g body weight) kept in pairs in 60‐liter aquaria fitted with infrared sensors connected to a computer to continuously record fish movements. The first experiment was designed to study the endogenous circadian clock under free‐running conditions [ultradian 40:40 min LD pulses and constant dark (DD)] and after shifting the LD cycle. Our results demonstrate that tench has a strictly nocturnal activity pattern, an endogenous rhythm being evident in 45.8% of the fish analyzed. The second experiment was conducted to test the influence of different photoperiods (LD 6:18, 12:12, 18:6, and 22:2) on locomotor activity, the results showing that even under an extremely long photoperiod, tench activity is restricted to dark hours. The third experiment examined the effect of light intensity on locomotor activity rhythms. When fish were exposed to decreasing light intensities (from 300:0 lux to 30:0, 3:0, and 0.3:0 lux) while maintaining a constant photoperiod (LD 12:12), the highest percentage of locomotor activity was in all cases associated with the hours of complete darkness (0 lux). In short, our results clearly show that (a) tench is a species with a strictly nocturnal behavior, and (b) daily activity rhythms gradually entrain after shifting the LD cycle and persist under free‐running conditions, pointing to their circadian nature. However, light strongly influences activity rhythms, since (c) the length of the active phase is directly controlled by the photophase, and (d) strictly nocturnal behavior persists even under very dim light conditions (0.3 lux). The above findings deepen our knowledge of tench behavior, which may help to optimize the aquacultural management of this species, for example, by adjusting feeding strategies to their nocturnal behavior.     

The Influence of Light Wavelength on Phase-Dependent Responsiveness of the Photoperiodic Clock in Migratory Blackheaded Bunting

We investigated the phase-dependent effects of light wavelength on photoperiodic clock in the migratory blackheaded bunting. Two experiments were performed, employing a skeleton paradigm (6 hours light : 6 hours darkness : 1 hour light : 11 hours darkness; 6L : 6D : 1L : 11D) at 37 ± 2 lux intensity. In the experiment 1, both 6 and 1 h light pulses were given at the same wavelength, 500 nm (green) or 650 nm (red). A group exposed to both pulses of white light served as control. In the experi-ment 2, the two light pulses were given at two different wavelengths, 6 h at 500 nm (green) and 1 h at 640 nm (red) in one group or vice-versa in the other. There was almost no photoinduction when both light pulses in experiment 1, or 1 h light pulse in experiment 2, were green. On the other hand, birds fattened and testes recrudesced when both the light pulses in experiment 1, or 1 h light pulse in experiment 2, were red. Birds receiving both pulses of white light in experiment 1 showed an intermediate response. Taken together, these results indicate that the photoperiodic clock in buntings is differentially responsive at its various circadian phases to different light wavelengths.     

The Influence of Light Wavelength on Phase-Dependent Responsiveness of the Photoperiodic Clock in Migratory Blackheaded Bunting

We investigated the phase-dependent effects of light wavelength on photoperiodic clock in the migratory blackheaded bunting. Two experiments were performed, employing a skeleton paradigm (6 hours light : 6 hours darkness : 1 hour light : 11 hours darkness; 6L : 6D : 1L : 11D) at 37 ± 2 lux intensity. In the experiment 1, both 6 and 1 h light pulses were given at the same wavelength, 500 nm (green) or 650 nm (red). A group exposed to both pulses of white light served as control. In the experi-ment 2, the two light pulses were given at two different wavelengths, 6 h at 500 nm (green) and 1 h at 640 nm (red) in one group or vice-versa in the other. There was almost no photoinduction when both light pulses in experiment 1, or 1 h light pulse in experiment 2, were green. On the other hand, birds fattened and testes recrudesced when both the light pulses in experiment 1, or 1 h light pulse in experiment 2, were red. Birds receiving both pulses of white light in experiment 1 showed an intermediate response. Taken together, these results indicate that the photoperiodic clock in buntings is differentially responsive at its various circadian phases to different light wavelengths.     

Neural Correlates of Migration: Activation of Hypothalamic Clock(s) in and out of Migratory State in the Blackheaded Bunting (Emberiza melanocephala)

Background

Many vertebrates distinguish between short and long day lengths using suprachiasmatic nuclei (SCN). In birds particular, the mediobasal hypothalamus (MBH) is suggested to be involved in the timing of seasonal reproduction. This study investigated the response of SCN and MBH to a single long day, and the role of MBH in induction of the migratory phenotype in night-migratory blackheaded buntings.

Methodology/Principal Findings

Experiment 1 immunocytochemically measured c-fos in the SCN, and c-fos, vasoactive intestinal peptide (VIP) and neuropeptide Y (NPY) in the MBH of buntings exposed to a 20 h light period. Long light period induced significantly stronger c-fos expression, measured as number of c-fos-like immunoreactive (c-fos-lir) cells, in MBH, but not in the SCN. Within the MBH, c-fos-lir cells were significantly denser in the inferior hypothalamic nucleus (IH) and infundibular nucleus (IN), but not in the dorsomedial hypothalamus (DMH). IH and IN also had significantly increased number of VIP and NPY labeled cells. DMH had significantly increased number of VIP labeled cells only. Experiment 2 assayed c-fos, VIP and NPY immunoreactivities in the middle of day and night in the MBH of buntings, after seven long days (day active, non-migratory state) and after seven days of Zugunruhe (night active, migratory state) in long days. In the migratory state, the number of c-fos-lir cells was significantly greater only in DMH; VIP-lir cells were denser in all three MBH regions suggesting enhanced light sensitivity at night. The denser NPY-lir cells only in IN in the non-migratory state were probably due to premigratory hyperphagia.

Conclusions/Significance

In buntings, SCN may not be involved in the photoperiod-induced seasonal responses. MBH contains the seasonal clock sensitive to day length. VIP and NPY are parts of the neuroendocrine mechanism(s) involved, respectively, in sensing and translating the photoperiodic message in a seasonal response.     

Decline of Corn Buntings Emberiza calandra on east Scottish study areas in 1989–2007

Capsule Adult numbers on many study areas fluctuated from year to year, but overall showed a large decline, down to extinction on most areas.

Aims To measure changes in summer numbers of adult Corn Buntings in a large sample of study areas holding discrete localized populations.

Methods Singing cocks and hens paired with them were counted in early summer at 30 study areas on farmland from south Angus to central Aberdeenshire. Sixteen populations occurring throughout 1989–95 were called ‘groups’. Counts also included 16 cases where a new ‘offshoot’ appeared in late May or the start of June, not having been seen earlier in that spring or in the previous winter or summer.

Results Following relative stability in 1989–95, large declines occurred in 1995–96, 1998–99 and 2003–04, and decreases far outweighed occasional increases between consecutive summers. The numbers of established groups, new offshoots, and older offshoots all declined with the year. Although containing one to nine founding pairs, new offshoots became extinct later, in six cases after one summer. No new offshoot was seen after 1995.

Conclusions Although 22 study areas held birds in 1989 and 25 in 1990, this fell to 10 in 1999, seven in 2005, and four in 2007 (the only areas that held birds in all years). On all areas combined, cock numbers fell by 83% between 1989 and 2007. Targeted implementation of appropriate agri‐environment measures is urgently needed to halt and reverse such declines, a challenge made all the greater within the last year by rising grain prices, delays to the introduction of new measures, and abolition of EU ‘set‐aside’ as a means for providing food and nesting habitat.     

Photoperiodic Entrainment Patterns in the CO(2) Output of Lemna perpusilla 6746 and of Several Other Lemnaceae

The CO₂ output of Lemna perpusilla 6746 in “skeleton photoperiods” consisting of alternating 10½-hour and 13-hour dark periods separated by ¼-hour illuminations was recorded under stable high and low nitrate conditions. The phase relationship finally attained between light schedule and output is the same regardless of which dark period is given first, but entrainment is more rapid (as is flowering) with an initial 13-hour dark period. In all respects other than bistability—the assumption of two different stable phase relationships depending on the initial dark period—both flowering and the course of CO₂ output conform to Pittendrigh's model derived from Drosophila eclosion rhythms, confirming the view that an endogenous circadian rhythm, or biological clock, underlies the photoperiodic control of flowering in this plant. Experiments with rigorous temperature control show that earlier results with long light exposures were in part due to temperature changes; in consequence, it is clear that entrainment patterns with high nitrate differ even more from those in low nitrate than was previously evident, and not simply by the addition of a “nitrate peak.” Other Lemnaceae tested with a few simple light-dark schedules in both types of media show a variety of responses, with no obvious correlation to photoperiodic response type.     

Light-Dark Entrainment of Circadian Activity Rhythms of the Diurnal Indian Palm Squirrel (Funambulus pennanti)

A recent focus of chronobiological studies has been to establish diurnal models as alternatives to the more frequently used nocturnal rodents. In the present study, light-dark (LD) entrainment characteristics were examined in one diurnal species, the Indian palm squirrel (Funambulus pennanti). Palm squirrels showed strongly diurnal locomotor activity rhythms (? 88 percent) under light-dark (LD) cycles, with activity bimodally distributed during the L phase. In comparison to a dim LD cycle, exposure to a bright LD cycle caused a phase advance in the onset of activity, an increase in daily activity levels and an increase in the duration of activity. Percentage diurnality, however, did not vary between bright and dim LD cycles. Activity rhythms reentrained in significantly fewer days after an 8 hour phase delay of the LD cycle compared to an 8 hour phase advance. In both cases, the direction of reentrainment followed the direction of the LD shift. When exposed to single light pulses (1 hour) presented at the same time each day, 6/7 squirrels entrained. Under a skeletal photoperiod cycle (2 x 1 hour light pulses each day), 6/8 squirrels showed stable entrainment. The remaining squirrels exhibited rhythm splitting, with each component synchronising in an unstable manner with one of the light pulses. Under entrainment to single light pulses and to the skeletal photoperiod cycle, the phase angle of entrainment was negatively correlated with t. Finally, when exposed to a skeletal scotoperiod cycle (2 x 1-hour dark pulses each day), only 3/8 squirrels entrained, while the others free-ran. Two of the entrained squirrels showed spontaneous phase reversals during entrainment. As with other species, the activity rhythm of palm squirrels appears to be controlled by two separate self-sustaining oscillators. The strongly diurnal nature of palm squirrels make them a promising diurnal model for studies examining endogenous and exogenous influences on circadian functioning.     

Light-Dark Entrainment of Circadian Activity Rhythms of the Diurnal Indian Palm Squirrel (Funambulus pennanti)

A recent focus of chronobiological studies has been to establish diurnal models as alternatives to the more frequently used nocturnal rodents. In the present study, light-dark (LD) entrainment characteristics were examined in one diurnal species, the Indian palm squirrel ( Funambulus pennanti ). Palm squirrels showed strongly diurnal locomotor activity rhythms (~ 88 percent) under light-dark (LD) cycles, with activity bimodally distributed during the L phase. In comparison to a dim LD cycle, exposure to a bright LD cycle caused a phase advance in the onset of activity, an increase in daily activity levels and an increase in the duration of activity. Percentage diurnality, however, did not vary between bright and dim LD cycles. Activity rhythms reentrained in significantly fewer days after an 8 hour phase delay of the LD cycle compared to an 8 hour phase advance. In both cases, the direction of reentrainment followed the direction of the LD shift. When exposed to single light pulses (1 hour) presented at the same time each day, 6/7 squirrels entrained. Under a skeletal photoperiod cycle (2 x 1 hour light pulses each day), 6/8 squirrels showed stable entrainment. The remaining squirrels exhibited rhythm splitting, with each component synchronising in an unstable manner with one of the light pulses. Under entrainment to single light pulses and to the skeletal photoperiod cycle, the phase angle of entrainment was negatively correlated with t. Finally, when exposed to a skeletal scotoperiod cycle (2 x 1-hour dark pulses each day), only 3/8 squirrels entrained, while the others free-ran. Two of the entrained squirrels showed spontaneous phase reversals during entrainment. As with other species, the activity rhythm of palm squirrels appears to be controlled by two separate self-sustaining oscillators. The strongly diurnal nature of palm squirrels make them a promising diurnal model for studies examining endogenous and exogenous influences on circadian functioning.