[show abstract][hide abstract] ABSTRACT: Artificial light-at-night is known to affect a broad array of behaviours and physiological processes. In urbanized bird species, light-at-night advances important biological rhythms such as daily cycles of activity/rest and timing of reproduction, but our knowledge of the underlying physiological mechanisms is limited. Given its role as chronobiological signal, melatonin is a strong candidate for mediating the effects of light-at-night.
We exposed urban and rural European blackbirds (Turdus merula) to two light treatments equal in photoperiod but with different light intensities at night. The control group was exposed to 0.0001 lux (almost darkness), while the experimental group was exposed to 0.3 lux at night, simulating conditions recorded previously on free-living urban blackbirds. We obtained diel profiles of plasma melatonin for all birds in summer (July) and winter (January), while simultaneously recording locomotor activity. Daily patterns of melatonin concentrations were clearly affected by light-at-night in both seasons. In winter, melatonin concentrations of light-at-night birds were lower in the early and late night than in those of birds kept in darkness. In summer, melatonin concentrations of the light-at-night birds were lower through all night compared to birds kept in darkness. Locomotor activity in light-at-night birds was overall higher than in control individuals, both during the day and at night, and it increased sharply before dawn. In winter, the amount of activity before dawn in the light-at-night group correlated with changes in melatonin from midnight to late night: the greater the decrease in melatonin, the greater the amount of pre-dawn activity. Urban and rural birds responded similarly to light-at-night with respect to melatonin, but differed in their behaviour, with rural birds showing more locomotor activity than urban counterparts.
This study points to reduced melatonin release at night as a potential physiological mechanism underlying the advanced onset of morning activity of urbanized birds. Based on the pattern of melatonin secretion, we suggest that birds responded to light-at-night as if they were exposed to a longer day than birds kept under dark nights.
Frontiers in Zoology 10/2013; 10(1):60. · 3.87 Impact Factor
[show abstract][hide abstract] ABSTRACT: Arctic environments are challenging for circadian systems. Around the solstices, the most important zeitgeber, the change between night and day, is reduced to minor fluctuations in light intensities. However, many species including songbirds nonetheless show clear diel activity patterns. Here we examine the possible physiological basis underlying diel rhythmicity under continuous Arctic summer light. Rhythmic secretion of the hormone melatonin constitutes an important part of the songbird circadian system and its experimental suppression, e.g., by constant light, usually leads to behavioral arrhythmia. We therefore studied melatonin patterns in a free-living migratory songbird, the willow warbler (Phylloscopus trochilus), that maintains diel activity during the Arctic summer. We compared melatonin profiles during late spring and summer solstice in two Swedish populations from the south (58°N) and near the Arctic circle (66°N). We found the northern Swedish population maintained clear diel changes in melatonin secretion during the summer solstice, although peak concentrations were lower than in southern Sweden. Melatonin levels were highest before midnight and in good accordance with periods of reduced activity. The maintenance of diel melatonin rhythmicity under conditions of continuous light may be one of the physiological mechanisms that enables continued functioning of the circadian system.
Hormones and Behavior 02/2013; 56(1):163-168. · 3.74 Impact Factor
[show abstract][hide abstract] ABSTRACT: Circadian clocks are centrally involved in the regulation of daily behavioural and physiological processes. These clocks are synchronized to the 24 h day by external cues (Zeitgeber), the most important of which is the light-dark cycle. In polar environments, however, the strength of the Zeitgeber is greatly reduced around the summer and winter solstices (continuous daylight or continuous darkness). How animals time their behaviour under such conditions has rarely been studied in the wild. Using a radio-telemetry-based system, we investigated daily activity rhythms under continuous daylight in Barrow, Alaska, throughout the breeding season in four bird species that differ in mating system and parental behaviour. We found substantial diversity in daily activity rhythms depending on species, sex and breeding stage. Individuals exhibited either robust, entrained 24 h activity cycles, were continuously active (arrhythmic) or showed 'free-running' activity cycles. In semipalmated sandpipers, a shorebird with biparental incubation, we show that the free-running rhythm is synchronized between pair mates. The diversity of diel time-keeping under continuous daylight emphasizes the plasticity of the circadian system, and the importance of the social and life-history context. Our results support the idea that circadian behaviour can be adaptively modified to enable species-specific time-keeping under polar conditions.
Proceedings of the Royal Society B: Biological Sciences 01/2013; 280(1764):20131016. · 5.68 Impact Factor
[show abstract][hide abstract] ABSTRACT: Seasonal recurrence of biological processes (phenology) and its relationship to environmental change is recognized as being of key scientific and public concern, but its current study largely overlooks the extent to which phenology is based on biological time-keeping mechanisms. We highlight the relevance of physiological and neurobiological regulation for organisms' responsiveness to environmental conditions. Focusing on avian and mammalian examples, we describe circannual rhythmicity of reproduction, migration and hibernation, and address responses of animals to photic and thermal conditions. Climate change and urbanization are used as urgent examples of anthropogenic influences that put biological timing systems under pressure. We furthermore propose that consideration of Homo sapiens as principally a 'seasonal animal' can inspire new perspectives for understanding medical and psychological problems.
Proceedings of the Royal Society B: Biological Sciences 01/2013; 280(1765):20130016. · 5.68 Impact Factor
[show abstract][hide abstract] ABSTRACT: Circadian rhythms are ubiquitous in many organisms. Animals that are forced to be active around the clock typically show reduced performance, health and survival. Nevertheless, we review evidence of animals showing prolonged intervals of activity with attenuated or nil overt circadian rhythms and no apparent ill effects. We show that around-the-clock and ultradian activity patterns are more common than is generally appreciated, particularly in herbivores, in animals inhabiting polar regions and habitats with constant physical environments, in animals during specific life-history stages (such as migration or reproduction), and in highly social animals. The underlying mechanisms are diverse, but studies suggest that some circadian pacemakers continue to measure time in animals active around the clock. The prevalence of around-the-clock activity in diverse animals and habitats, and an apparent diversity of underlying mechanisms, are consistent with convergent evolution. We suggest that the basic organizational principles of the circadian system and its complexity encompass the potential for chronobiological plasticity. There may be trade-offs between benefits of persistent daily rhythms versus plasticity, which for reasons still poorly understood make overt daily arrhythmicity functionally adaptive only in selected habitats and for selected lifestyles.
Proceedings of the Royal Society B: Biological Sciences 01/2013; 280(1765):20130019. · 5.68 Impact Factor
[show abstract][hide abstract] ABSTRACT: Daily schedules of many organisms, including birds, are thought to affect fitness. Timing in birds is based on circadian clocks that have a heritable period length, but fitness consequences for individuals in natural environments depend on the scheduling of entrained clocks. This chronotype, i.e., timing of an individual relative to a zeitgeber, results from interactions between the endogenous circadian clock and environmental factors, including light conditions and ambient temperature. To understand contributions of these factors to timing, we studied daily activity patterns of a captive songbird, the great tit (Parus major), under different temperature and light conditions. Birds were kept in a light (L)-dark (D) cycle (12.5 L:11.5 D) at either 8°C or 18°C with ad libitum access to food and water. We assessed chronotype and subsequently tested birds at the same temperature under constant dim light (LL(dim)) to determine period length of their circadian clock. Thermal conditions were then reversed so that period length was measured under both temperatures. We found that under constant dim light conditions individuals lengthened their free-running period at higher temperatures by 5.7 ± 2.1 min (p = .002). Under LD, birds kept at 18°C started activity later and terminated it much earlier in the day than those kept under 8°C. Overall, chronotype was slightly earlier under higher temperature, and duration of activity was shorter. Furthermore, individuals timed their activities consistently on different days under LD and over the two test series under LL(dim) (repeatability from .38 to .60). Surprisingly, period length and chronotype did not show the correlation that had been previously found in other avian species. Our study shows that body clocks of birds are precise and repeatable, but are, nonetheless, affected by ambient temperature. (Author correspondence: firstname.lastname@example.org ).
Chronobiology International 08/2012; 29(8):1062-71. · 4.35 Impact Factor
[show abstract][hide abstract] ABSTRACT: At higher latitudes, most organisms use the periodic changes in day length to time their annual life cycle. At the equator, changes in day length are minimal, and it is unknown which cues organisms use to synchronize their underlying circannual rhythms to environmental conditions. Here, we demonstrate that the African stonechat (Saxicola torquatus axillaris)-an equatorial songbird-can use subtle solar cues for the annual timing of postnuptial moult, a reliable marker of the circannual cycle. We compared four groups that were kept over more than 3 years: (i) a control group maintained under constant equatorial day length, (ii) a 12-month solar time group maintained under equatorial day length, but including a simulation of the annual periodic change in sunrise and sunset times (solar time), (iii) a 14-month solar time group similar to the previous group but with an extended solar time cycle and (iv) a group maintained under a European temperate photoperiod. Within all 3 years, 12-month solar time birds were significantly more synchronized than controls and 14-month solar time birds. Furthermore, the moult of 12-month solar time birds occurred during the same time of the year as that of free-living Kenyan conspecifics. Thus, our data indicate that stonechats may use the subtle periodic pattern of sunrise and sunset at the equator to synchronize their circannual clock.
Proceedings of the Royal Society B: Biological Sciences 05/2012; 279(1742):3527-34. · 5.68 Impact Factor
[show abstract][hide abstract] ABSTRACT: Timing "in the real world" must cope with the temporal complexity of natural environments. Extreme examples for the resultant "multitasking" are migratory birds, which precisely time movements to remote areas. New field technologies highlight temporal accuracy, while captivity studies emphasize underlying programs and plasticity of schedules. After reviewing these findings, we focus on waders, which undertake spectacular long-distance migrations, have robust circannual clocks, and cope with diel, tidal, and polar environments. To explore features that may facilitate such multitasking, we speculated that melatonin amplitudes are low and damped during seasons when entrainment to subtle Zeitgebers occurs. We measured melatonin profiles under European daylength in two species with different ecologies and found low-amplitude melatonin cycles that changed over the year. Annual patterns neither fully supported our hypothesis, nor simply reflected daylight availability. While migratory birds are inspiring models for chronobiology, mechanistic understanding of their multitasking is still poor.
Progress in brain research 01/2012; 199:457-79. · 4.19 Impact Factor
[show abstract][hide abstract] ABSTRACT: Timing is essential, but circadian clocks, which play a crucial role in timekeeping, are almost unaddressed in evolutionary ecology. A key property of circadian clocks is their free-running period length (τ), i.e. the time taken for a full cycle under constant conditions. Under laboratory conditions, concordance of τ with the ambient light-dark cycle confers major fitness benefits, but little is known about period length and its implications in natural populations. We therefore studied natural variation of circadian traits in a songbird, the great tit (Parus major), by recording locomotor activity of 98 hand-raised, wild-derived individuals. We found, unexpectedly, that the free-running period of this diurnal species was significantly shorter than 24 h in constant dim light. We furthermore demonstrate, to our knowledge for the first time in a wild vertebrate, ample genetic variation and high heritability (h(2) = 0.86 ± 0.24), implying that period length is potentially malleable by micro-evolutionary change. The observed, short period length may be a consequence of sexual selection, as offspring from extra-pair matings had significantly shorter free-running periods than their half-siblings from within-pair matings. These findings position circadian clocks in the 'real world' and underscore the value of using chronobiological approaches in evolutionary ecology. Evolutionary ecologists study variation and its fitness consequences, but often have difficulties relating behavioural variation to physiological mechanisms. The findings presented here open the possibility that properties of internal, circadian clocks affect performance in traits that are relevant to fitness and sexual selection.
Proceedings of the Royal Society B: Biological Sciences 11/2010; 277(1698):3335-42. · 5.68 Impact Factor
[show abstract][hide abstract] ABSTRACT: Phenology refers to the periodic appearance of life-cycle events and currently receives abundant attention as the effects of global change on phenology are so apparent. Phenology as a discipline observes these events and relates their annual variation to variation in climate. But phenology is also studied in other disciplines, each with their own perspective. Evolutionary ecologists study variation in seasonal timing and its fitness consequences, whereas chronobiologists emphasize the periodic nature of life-cycle stages and their underlying timing programmes (e.g. circannual rhythms). The (neuro-) endocrine processes underlying these life-cycle events are studied by physiologists and need to be linked to genes that are explored by molecular geneticists. In order to fully understand variation in phenology, we need to integrate these different perspectives, in particular by combining evolutionary and mechanistic approaches. We use avian research to characterize different perspectives and to highlight integration that has already been achieved. Building on this work, we outline a route towards uniting the different disciplines in a single framework, which may be used to better understand and, more importantly, to forecast climate change impacts on phenology.
Philosophical Transactions of The Royal Society B Biological Sciences 10/2010; 365(1555):3113-27. · 6.23 Impact Factor
[show abstract][hide abstract] ABSTRACT: With progressively faster global change, shifts in phenology, and distributional ranges are reported for an increasing number of species. The success of organisms at coping with novel seasonal conditions depends on the mechanisms that determine their schedules. Species that rely on fixed schedules and those that time their activities by predictive cues may be particularly constrained in their ability to accommodate changes. The present study examines rigid scheduling and its implications for breeding in captivity in an avian model taxon, the Stonechat (Saxicola torquata). Within their extensive breeding range, Stonechats differ geographically in migratory behavior and reproduce and molt under a wide range of daylengths (10-17 h). Stonechats time their activities by programs that involve circannual rhythms and photoperiodism. The study reports reproductive timing of four taxa (central European, Irish, Siberian, and Kenyan), relates it to laydates in the field, and investigates modifying influences of housing conditions and of social context. Reproductive consequences of timing programs were then tested by crossbreeding of taxa with different schedules. The study revealed persistent, population-specific breeding windows in captivity. Resident Stonechats from equatorial Kenya synchronized their reproductive cycles with the European summer, presumably in response to local photoperiod, and bred at similar times as northern migrants. In all other taxa schedules matched those in the field, but were timed slightly earlier in captivity and advanced by indoor keeping conditions. Influences of social context were negligible. In pairs with clutches, testes regressed slightly later than in pairs without clutches, but presence of a mate per se had no influence on breeding cycles. Accordingly, crossbreeding Stonechats were predicted to have limited capacity to adjust schedules to those of their mates. This prediction was tested by crossbreeding of single-clutched Siberian long-distance migrants with multiple-clutched European short-distance migrants. Males and females of both taxa retained their characteristic breeding schedules, regardless of their mate's activities. This led to dramatic loss of reproductive success in the population with the longer breeding season, European Stonechats. Siberian Stonechats were unable to profit from the presence of a sexually active mate, but they suffered no disadvantage from crossbreeding. In a changing world, inherited timing programs may severely constrain responses to novel conditions, impose schedule-dependent, asymmetric costs of hybridization, and contribute to directional gene flow or to reproductive isolation.
Integrative and Comparative Biology 11/2009; 49(5):563-79. · 3.02 Impact Factor
[show abstract][hide abstract] ABSTRACT: To anticipate seasonal change, organisms schedule their annual activities by using calendrical cues like photoperiod. The use of cues must be fitted to local conditions because schedules differ between species and habitats. In complete absence of temporal information, many species show persistent circannual cycles that are synchronised, but not driven, by photoperiod. The contribution of circannual rhythms to timing under natural photoperiodic conditions is still unclear. In a suite of experiments, we examined timing in two closely related songbirds (Siberian and European stonechats) that inhabit similar latitudes but differ in seasonal behaviour. Under a more continental climate, Siberian stonechats breed later, moult faster and migrate further than European stonechats. We tested hypotheses for seasonal timing mechanisms by comparing the birds under constant and naturally changing daylengths. The taxa retained characteristic reproductive and moult schedules and hybrids behaved roughly intermediately. Based on their distinct circannual cycles, we expected European and Siberian stonechats to differ in photoperiodic responses at a given time of year. We found that the taxa responded, as predicted, in opposite ways to photoperiodic simulations as experienced on different migration routes. The findings indicate that circannual rhythms reflect geographically distinct periodic changes in seasonal disposition and cue-response mechanisms. Under natural daylengths, the phase relationship of the underlying circannual rhythm to the external year determines the action of photoperiod. Circannual rhythms are widespread among long-lived species. Accordingly, responses to environmental change, range expansion and novel migration patterns may depend on the particulars of a species' underlying circannual programming.
[show abstract][hide abstract] ABSTRACT: Endogenous circannual clocks are found in many long-lived organisms, but are best studied in mammal and bird species. Circannual clocks are synchronized with the environment by changes in photoperiod, light intensity and possibly temperature and seasonal rainfall patterns. Annual timing mechanisms are presumed to have important ultimate functions in seasonally regulating reproduction, moult, hibernation, migration, body weight and fat deposition/stores. Birds that live in habitats where environmental cues such as photoperiod are poor predictors of seasons (e.g. equatorial residents, migrants to equatorial/tropical latitudes) rely more on their endogenous clocks than birds living in environments that show a tight correlation between photoperiod and seasonal events. Such population-specific/interspecific variation in reliance on endogenous clocks may indicate that annual timing mechanisms are adaptive. However, despite the apparent adaptive importance of circannual clocks, (i) what specific adaptive value they have in the wild and (ii) how they function are still largely untested. Whereas circadian clocks are hypothesized to be generated by molecular feedback loops, it has been suggested that circannual clocks are either based upon (i) a de-multiplication ('counting') of circadian days, (ii) a sequence of interdependent physiological states, or (iii) one or more endogenous oscillators, similar to circadian rhythms. We tested the de-multiplication of days (i) versus endogenous regulation hypotheses (ii) and (iii) in captive male and female house sparrows (Passer domesticus). We assessed the period of reproductive (testicular and follicular) cycles in four groups of birds kept either under photoperiods of LD 12L:12D (period length: 24h), 13.5L:13.5D (27 h), 10.5L:10.5D (23 h) or 12D:8L:3D:1L (24-h skeleton photoperiod), respectively, for 15 months. Contrary to predictions from the de-multiplication hypothesis, individuals experiencing 27-h days did not differ (i.e. did not have longer) annual reproductive rhythms than individuals from the 21- or 24-h day groups. However, in line with predictions from endogenous regulation, birds in the skeleton group had significantly longer circannual period lengths than all other groups. Birds exposed to skeleton photoperiods experienced fewer light hours per year than all other groups (3285 versus 4380) and had a lower daily energy expenditure, as tested during one point of the annual cycle using respirometry. Although our results are tantalizing, they are still preliminary as birds were only studied over a period of 15 months. Nevertheless, the present data fail to support a 'counting of circadian days' and instead support hypotheses proposing whole-organism processes as the mechanistic basis for circannual rhythms. We propose a novel energy turnover hypothesis which predicts a dependence of the speed of the circannual clock on the overall energy expenditure of an organism.
Philosophical Transactions of The Royal Society B Biological Sciences 02/2008; 363(1490):411-23. · 6.23 Impact Factor
[show abstract][hide abstract] ABSTRACT: Daytime light intensity can affect the photoperiodic regulation of the reproductive cycle in birds. The actual way by which light intensity information is transduced is, however, unknown. We postulate that transduction of the light intensity information is mediated by changes in the pattern of melatonin secretion. This study, therefore, investigated the effects of high and low daytime light intensities on the daily melatonin rhythm of Afro-tropical stonechats (Saxicola torquata axillaris) in which seasonal changes in daytime light intensity act as a zeitgeber of the circannual rhythms controlling annual reproduction and molt. Stonechats were subjected to light conditions simulated as closely as possible to native conditions near the equator. Photoperiod was held constant at 12.25 h of light and 11.75 h of darkness per day. At intervals of 2.5 to 3.5 weeks, daytime light intensity was changed from bright (12,000 lux at one and 2,000 lux at the other perch) to dim (1,600 lux at one and 250 lux at the other perch) and back to the original bright light. Daily plasma melatonin profiles showed that they were linked with changes in daytime light intensity: Nighttime peak and total nocturnal levels were altered when transitions between light conditions were made, and these changes were significant when light intensity was changed from dim to bright. We suggest that daytime light intensity could affect seasonal timing via changes in melatonin profiles.
[show abstract][hide abstract] ABSTRACT: The urge of captive birds to migrate manifests itself in seasonally occurring restlessness, termed "Zugunruhe." Key insights into migration and an endogenous basis of behavior are based on Zugunruhe of migrants but have scarcely been tested in nonmigratory birds. We recorded Zugunruhe of African stonechats, small passerine birds that defend year-round territories and have diverged from northern migrants at least 1 million years ago. We demonstrate that Zugunruhe is a regular feature of their endogenous program, one that is precisely timed by photoperiod. These results extend ideas of programs for periodic movement to include nonmigratory birds. Such programs could be activated when movements become necessary, in line with observed fast changes and high flexibility of migration. Attention to Zugunruhe of resident birds promises new insights into diverse and dynamic migration systems and enhances predictions of avian responses to global change.
[show abstract][hide abstract] ABSTRACT: The day lengths to which migratory birds are exposed depend on the timing and course of their journey. While winter day length
is known to influence vernal events, it is not clear if birds also use day length during the spring migration as a temporal
cue. We addressed this question by exposing captive stonechats (Saxicola torquata) to two different photoperiodic simulations of spring migration routes, following common winter conditions. One group experienced
day lengths of the regular (“fast”) migration, and the other group, a “slow”, or more southerly originating, route. The resulting
small, temporary differences in day length had lasting effects on the birds. The groups differed in migratory restlessness
during and following exposure to different day lengths. “Slow” migrants continued nocturnal activity longer than “fast” migrants.
Furthermore, all activities of the ensuing breeding season were delayed in the “slow” migrants, indicating a phase shift in
their underlying annual rhythm. “Slow” migrants delayed terminating their reproductive stage by regressing testes and the
cloacal protuberance later than the “fast” migrants. Molt started and ended later in “slow” migrants, but the duration of
the molt was unaffected by spring day length. Finally, “fast” migrants resumed nightly restlessness earlier than “slow” migrants
in late summer. These results demonstrate that Zugunruhe (migratory restlessness) and reproductive windows are not set exclusively
during winter but can be modified by day length cues during the spring migration. Because migration modifies the day length
exposure of birds, migration routes can have carry-over effects on the timing of breeding season events, including the completion
of molt and initiation of autumnal nocturnal activity.
Zugvögel beeinflussen durch ihre Zugroute und Zugzeit die Tageslänge, in der sie sich befinden. Während experimentell belegt
ist, dass die Wintertageslänge das saisonale Verhalten im Frühling beeinflusst, ist bisher unbekannt, ob Vögel auch während
der Frühjahrszugzeit photoperiodische Zeitinformationen nutzen. Um diese Frage zu beantworten, haben wir Schwarzkehlchen (Saxicola torquata) unter den Lichtbedingungen von zwei verschiedenen Zugrouten untersucht. Die Vögel hatten unter einheitlichen Bedingungen
überwintert. Anschließend erlebte eine Gruppe die Lichtbedingungen ihres regulären (“schnellen“) Zugweges, die andere die
eines “langsameren“, oder weiter südlich beginnenden Zugweges. Die damit verbundenen geringen Unterschiede in der Tageslänge
zeigten nachhaltige Wirkung. Vögel der beiden Gruppen unterschieden sich in Zugunruhe während und nachdem sie unterschiedliche
Tageslängen erlebten. “Langsame” Zieher setzten ihre nächtliche Aktivität länger fort als “schnelle" Zieher. Darüber hinaus
verzögerten sich bei den “langsamen" Ziehern alle Aktivitäten der anschließenden Brutsaison. Dies deutet auf eine Phasenverschiebung
der zugrunde liegenden Jahresrhythmen hin. “Langsame" Zieher verzögerten den Abschluss ihrer Brutphase, indem sie Hoden und
Kloakenprotuberanz später zurückentwickelten als “schnelle" Zieher. Die Mauser begann und endete später als bei “schnellen"
Ziehern, dauerte aber gleich lang an. Schließlich nahmen “schnelle" Zieher die nächtliche Unruhe früher wieder auf als “langsame"
Zieher. Diese Ergebnisse zeigen, dass der Brutzeitraum von Zugvögeln nicht allein im Winter, sondern auch während des Frühjahrszugs
von der Tageslänge beeinflusst wird. Somit können sich Zugzeit und Zugroute auf das anschließende zeitliche Verhalten bis
zu Mauserende und Beginn der Herbstzugunruhe auswirken.
[show abstract][hide abstract] ABSTRACT: Rigid schedules of long-distance migrants could be among candidate traits for adaptive migratory syndromes. This prediction was tested on stonechats, passerines that differ widely in migratory behavior and seasonal schedules. Stonechats in Europe are short-distance migrants and multiclutched, whereas African residents and Siberian long-distance migrants usually raise single broods. In captivity, all subspecies displayed endogenous cycles of reproductive development and molt. The subspecies differed in time afforded to life cycle stages. Under conducive aviary conditions, African stonechats were multibrooded, whereas Siberian stonechats did not add clutches. This difference in flexibility was exclusively related to the length of breeding windows. Stonechats also differed in premigratory preparations. Postjuvenile molt started early in Siberian stonechats, but in European and African stonechats, depended strongly on hatching date. In contrast, all subspecies shortened molt duration at the same rate when hatched from late broods. Plasticity of Zugunruhe timing was identical in Siberian and European subspecies and nearly compensated for hatching late. The stonechat data suggest a refined understanding of temporal plasticity in long-distance migrants. Overall, plasticity was not reduced, but was differently organized. Apparently rigid migrant schedules were related to short breeding cycles and inflexible molt onset. Short windows for breeding and juvenile development could provide safety measures for timely departure. Once molt was initiated, temporal plasticity of long-distance migrants matched that of less migratory conspecifics. In addition to adjusting endogenous programs, stonechats differed in implementing them in the field. Modifying the conditions under which programs are expressed may be an efficient way to enhance seasonal plasticity.
Annals of the New York Academy of Sciences 07/2005; 1046:216-27. · 4.38 Impact Factor
[show abstract][hide abstract] ABSTRACT: Migrant species express wide varieties of movement strategies that closely mirror patterns of resource distribution. Resources can vary predictably in space and time (e.g., seasonal peaks of productivity in temperate regions) or can be unpredictable in one or both dimensions. Evolution of migration strategies and the underlying physiological and behavioral mechanisms are therefore diverse, but similarities have been noted. To understand how migrants respond to environmental cues, we focus on species that represent extremes along a continuum between obligate versus facultative migratory behavior and address the physiological mechanisms regulating these movements. Obligate migrants, which travel on highly predictable schedules, generally rely on the annual change in daylength and circannual rhythms to initiate the vernal and, where known, the fall phases of migration. Facultative species may show similar seasonal preparations in cases where resources are temporally predictable, but appear to rely more heavily upon local supplementary cues in the environment to initiate migrations. In general, the physiological response mechanisms to environmental cues that coordinate migratory movements include perception of the environmental cue and transduction to neuroendocrine and endocrine systems. Clearly, there are huge gaps in our knowledge in this area, but published studies of each of the migratory types suggest striking parallels. Based on a comparison of the physiological aspects of the divergent migratory patterns, we emphasize that the migratory types are not completely distinct. Instead, the differences of migration patterns and responsiveness to cues reflect differences in the relative contributions of common physiological mechanisms that underly the control of movements. A more complete understanding of migratory physiology requires continued descriptive studies, particularly in a greater diversity of model systems, in addition to experimental methods that utilize innovative tracking technology as well as behavioral and physiological innovations to draw attention to different levels of organization. Comprehending the physiological mechanisms underlying migratory patterns will be vital for knowing how migrants respond to current environmental cues and anticipating their reactions to future conditions.