No evidence for melatonin-linked immunoenhancement over the annual cycle of an avian species.

The winter immunoenhancement hypothesis associates long nights and increased exposure to melatonin with enhanced immune function in winter when resource availability is low and the chances of becoming ill are high. Thus, increased exposure to melatonin in the winter could be adaptive for species facing difficult winter conditions. This idea has found some support in studies of resident mammals. In birds, the link between day length and melatonin over the annual cycle is weaker, and contributions of melatonin to seasonal timing are unclear. Furthermore, many species, especially migrants, do not experience the most difficult conditions of their annual cycle in winter. In this study, we tested whether the winter immunoenhancement hypothesis holds in an avian species, the red knot Calidris canutus. We found that melatonin duration and amplitude varied significantly over the annual cycle with the highest values occurring in winter. However, peaks did not correspond to the winter solstice or with annual variation in immune function. Our findings do not support the winter immunoenhancement hypothesis in knots and question whether the idea that immune function should be bolstered in winter can be generalized to systems where winter is not the most difficult time of the year.

Are birds stressed during long-term flights? A wind-tunnel study on circulating corticosterone in the red knot.

During endurance flight most birds do not feed and have to rely on their body reserves. Fat and protein is catabolised to meet the high energetic demands. Even though the hormonal regulation of migration is complex and not yet fully understood, the adrenocortical hormone corticosterone crystallizes to play a major role in controlling physiological traits in migratory birds during flight. However, results from field studies are partially equivocal, not least because data from birds during endurance flight are hard to get and present mostly a momentary shot. A wind-tunnel experiment offered the possibility to measure repeatedly under controlled conditions the effect of long flights on the stress hormone corticosterone. In a long-distance migrating shorebird, the red knot Calidris canutus, we measured plasma concentrations of corticosterone within 3 min and after a restraint time of 30 min directly after 2h and 10h non-stop flights, respectively, and during rest. Baseline corticosterone levels were unchanged directly after the flights, indicating that endurance flight did not affect corticosterone levels. The adrenocortical response to restraint showed the typical rise in birds during rest, while birds after a 2 or 10h flight substantially decreased plasma corticosterone concentrations. We suggest that the negative adrenocortical response to restraint after flight is part of the mechanism to reduce the proteolytic effect of corticosterone to save muscle protein and to avoid muscle damaging effects.

Presence-absence of marine macrozoobenthos does not generally predict abundance and biomass.

Many monitoring programmes of species abundance and biomass increasingly face financial pressures. Occupancy is often easier and cheaper to measure than abundance or biomass. We, therefore, explored whether measuring occupancy is a viable alternative to measuring abundance and biomass. Abundance- or biomass-occupancy relationships were studied for sixteen macrozoobenthos species collected across the entire Dutch Wadden Sea in eight consecutive summers. Because the form and strength of these relationships are scale-dependent, the analysis was completed at different spatiotemporal scales. Large differences in intercept and slope of abundance- or biomass-occupancy relationships were found. Abundance, not biomass, was generally positively correlated with occupancy. Only at the largest scale, seven species showed reasonably strong abundance-occupancy relationships with large coefficients of determination and small differences in observed and predicted values (RMSE). Otherwise, and at all the other scales, intraspecific abundance and biomass relationships were poor. Our results showed that there is no generic relationship between a species' abundance or biomass and its occupancy. We discuss how ecological differences between species could cause such large variation in these relationships. Future technologies might allow estimating a species' abundance or biomass directly from eDNA sampling data, but for now, we need to rely on traditional sampling technology.

Seasonal Time Keeping in a Long-Distance Migrating Shorebird.

Because of the complications in achieving the necessary long-term observations and experiments, the nature and adaptive value of seasonal time-keeping mechanisms in long-lived organisms remain understudied. Here we present the results of a 20-year-long study of the repeated seasonal changes in body mass, plumage state, and primary molt of 45 captive red knots Calidris canutus islandica, a High Arctic breeding shorebird that spends the nonbreeding season in temperate coastal areas. Birds kept outdoors and experiencing the natural photoperiod of the nonbreeding area maintained sequences of life-cycle stages, roughly following the timing in nature. For 6 to 8 years, 14 of these birds were exposed to unvarying ambient temperature (12 °C) and photoperiodic conditions (12:12 LD). Under these conditions, for at least 5 years they expressed free-running circannual cycles of body mass, plumage state, and wing molt. The circannual cycles of the free-running traits gradually became longer than 12 months, but at different rates. The prebreeding events (onset and offset of prealternate molt and the onset of spring body mass increase) occurred at the same time of the year as in the wild population for 1 or several cycles. As a result, after 4 years in 12:12 LD, the circannual cycles of prealternate plumage state had drifted less than the cycles of prebasic plumage state and wing molt. As the onset of body mass gain drifted less than the offset, the period of high body mass became longer under unvarying conditions. We see these differences between the prebreeding and postbreeding life-cycle stages as evidence for adaptive seasonal time keeping in red knots: the life-cycle stages linked to the initiation of reproduction rely mostly on endogenous oscillators, whereas the later stages rather respond to environmental conditions. Postbreeding stages are also prone to carryover effects from the earlier stages.

Avian migration: temporal multitasking and a case study of melatonin cycles in waders.

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.

Reversed optimality and predictive ecology: burrowing depth forecasts population change in a bivalve.

Optimality reasoning from behavioural ecology can be used as a tool to infer how animals perceive their environment. Using optimality principles in a 'reversed manner' may enable ecologists to predict changes in population size before such changes actually happen. Here we show that a behavioural anti-predation trait (burrowing depth) of the marine bivalve Macoma balthica can be used as an indicator of the change in population size over the year to come. The per capita population growth rate between years t and t+1 correlated strongly with the proportion of individuals living in the dangerous top 4 cm layer of the sediment in year t: the more individuals in the top layer, the steeper the population decline. This is consistent with the prediction based on optimal foraging theory that animals with poor prospects should accept greater risks of predation. This study is among the first to document fitness forecasting in animals.

Long flights do not influence immune responses of a long-distance migrant bird: a wind-tunnel experiment.

Heavy physical work can result in physiological stress and suppressed immune function. Accordingly, long-distance migrant birds that fly for thousands of km within days can be expected to show immunosuppression, and hence be more vulnerable to infections en route. The red knot Calidris canutus Linnaeus is a long-distance migrant shorebird. We flew red knots the equivalent of 1500 km over 6 days in a wind tunnel. The humoral and cell-mediated immune responses of the flyers were compared to those of non-flying controls. Humoral immunity was measured as antibody production against injected diphtheria and tetanus antigens, and cell-mediated response as phytohemagglutinin-induced wing-web swelling. Blood corticosterone levels, which may modulate immune function, were measured in parallel. The long flights had no detectable effects on humoral or cell-mediated immune responses, or on corticosterone levels. Thus, flight performance per se may not be particularly stressful or immunosuppressive in red knots. Some birds assigned as flyers refused to fly for extended periods. Before flights started, these non-flyers had significantly lower antibody responses against tetanus than the birds that carried out the full flight program. This suggests that only birds in good physical condition may be willing to take on heavy exercise. We conclude that these long-distance migrants appear well adapted to the work load induced by long flights, enabling them to cope with long flight distances without increased stress levels and suppression of immunity. Whether this also applies in the wild, where the migrating birds may face adverse weather and food conditions, remains to be investigated.