Corticosterone in migrating songbirds during endurance flight

The specific role of the glucocorticoid hormone corticosterone in regulating the migratory stages of flight and refueling in free-living migrants is as yet poorly studied, because these stages are difficult to identify in the field. Night-migrating songbirds provide an excellent model to investigate how corticosterone correlates with behavior and physiology because they fly during the night and rest and forage during the day. We measured baseline corticosterone and the adrenocortical response to restraint in 9 free-ranging songbird species: 3 night-migrating species, 3 day-migrating species, and 3 day-migrating irruptive species. Baseline corticosterone of night migrants was higher in birds caught out of nocturnal migration than in birds resting and foraging, and on the same level as in day migrants, suggesting that a rise in circulating corticosterone may facilitate the heightened metabolic processes of active flight, in particular protein breakdown. Stress-induced corticosterone levels increased in both actively flying birds and birds resting and foraging. The increase was highest in landing birds, which are possibly most sensitive to stress when arriving at an unfamiliar place. Migratory endurance flight is thus characterized by corticosterone concentrations that are lower than those associated with acute stressful and life-threatening episodes. In addition, the responsiveness to stress increased with decreasing fat score in a night-migrating species. Corticosterone approaches therefore stressful concentrations only when fat depots are nearly depleted, possibly to promote protein catabolism and to trigger a change in behavior, i.e., a switch to landing and searching for food.     

Effect of endurance flight on haematocrit in migrating birds

The effects of an endurance flight on the haematocrit, the percentage of packed red blood cells per blood volume, were examined within the framework of six possible factors explaining possible changes in the haematocrit. Two approaches were adopted: (1) the haematocrit was studied in four species of passerine birds which landed on an Italian island after having crossed the Mediterranean Sea on their spring migration in a non-stop flight; (2) the haematocrit was evaluated in six individual red knots after a flight of 1, 2, 4 and 10 h in a wind tunnel and the data thus obtained compared with data on resting birds with or without food. In the four passerine species, the haematocrit decreased from 51% in fat birds to 48% in lean birds. In lean birds, the haematocrit dropped from 48% in birds with well-developed breast muscles to 36% in birds with emaciated breast muscles. In the red knots, the haematocrit was dependent on body mass in flying and resting birds. The haematocrit decreased from about 51% pre-flight to about 49% within 1 h of flight and remained at this level for up to 10 h of flight. Taking the results from the passerines and the red knots together, it seems that the haematocrit drops by a few percentage points within 1 h after the onset of flight, decreases very slowly with decreasing body mass and decreases more steeply in very lean birds having entered stage III of fasting. This indicates that dehydration is not an underlying factor in decreased haematocrit because if this were the case we would expect an increase with endurance flight. We found no effect of the presence of blood parasites on haematocrit. With the onset of flight, haemodilution may be adaptive, because it reduces blood viscosity and, thereby, energy expenditure by the heart, or it may be a sign of water conservation as an insurance against the risk of dehydration during long non-stop flights. During endurance flight, a reduction in the haematocrit may be adaptive, in that oxygen delivery capacity is adjusted to the decreased oxygen needs as body mass decreases. A decreasing haematocrit would also allow birds to reduce heart beat frequency and/or heart size, because blood viscosity decreases disproportionally with decreasing haematocrit. However, when energy stores are about to come to an end and birds increase protein breakdown, the haematocrit decreases even further, and birds probably become anaemic due to a reduced erythropoiesis.     

Within night correlations between radar and ground counts of migrating songbirds

ABSTRACT.   Studies comparing numbers of nocturnal migrants in flight with numbers of migrants at stopover sites have produced equivocal results. In 2003, we compared numbers of nocturnal migrants detected by radar to numbers of passerines observed at the Atlantic Bird Observatory in southwestern Nova Scotia, Canada. Numbers of nocturnal migrants detected by radar were positively correlated with numbers of migrants as determined by mist-netting, censuses, and daily estimated totals (daily estimates of birds present based netting and census results and casual observations) the following day. On nights with winds favorable for migration (tailwinds), the peak correlation between ground counts and radar counts the night before occurred just after sunset. On nights with unfavorable winds (headwinds), the correlation increased through the night, with a peak just before sunrise. The patterns of correlation are consistent with a scenario where birds accumulate at the coastline during periods of unfavorable wind, likely because they are not willing to cross a major ecological barrier, the Gulf of Maine. On nights with favorable winds, many birds departed, but some, possibly after testing wind conditions, apparently decided not to cross the Gulf of Maine and returned. Our results suggest that combining data collected using different methods to generate a daily estimated total provides the best estimate of the number of migrants present at a stopover site. Simultaneous studies at multiple locations where different census methods are used, making more effective use of temporal data (both from radar and diurnal counts), will more clearly elucidate patterns of flight behavior by migratory songbirds and the relationship between ground counts and counts of birds aloft.     

Pesticide exposure affects flight dynamics and reduces flight endurance in bumblebees

The emergence of agricultural land use change creates a number of challenges that insect pollinators, such as eusocial bees, must overcome. Resultant fragmentation and loss of suitable foraging habitats, combined with pesticide exposure, may increase demands on foraging, specifically the ability to collect or reach sufficient resources under such stress. Understanding effects that pesticides have on flight performance is therefore vital if we are to assess colony success in these changing landscapes. Neonicotinoids are one of the most widely used classes of pesticide across the globe, and exposure to bees has been associated with reduced foraging efficiency and homing ability. One explanation for these effects could be that elements of flight are being affected, but apart from a couple of studies on the honeybee (Apis mellifera), this has scarcely been tested. Here, we used flight mills to investigate how exposure to a field realistic (10 ppb) acute dose of imidacloprid affected flight performance of a wild insect pollinator—the bumblebee, Bombus terrestris audax. Intriguingly, observations showed exposed workers flew at a significantly higher velocity over the first ¾ km of flight. This apparent hyperactivity, however, may have a cost because exposed workers showed reduced flight distance and duration to around a third of what control workers were capable of achieving. Given that bumblebees are central place foragers, impairment to flight endurance could translate to a decline in potential forage area, decreasing the abundance, diversity, and nutritional quality of available food, while potentially diminishing pollination service capabilities.     

Bats' conquest of a formidable foraging niche: the myriads of nocturnally migrating songbirds

Along food chains, i.e., at different trophic levels, the most abundant taxa often represent exceptional food reservoirs, and are hence the main target of consumers and predators. The capacity of an individual consumer to opportunistically switch towards an abundant food source, for instance, a prey that suddenly becomes available in its environment, may offer such strong selective advantages that ecological innovations may appear and spread rapidly. New predator-prey relationships are likely to evolve even faster when a diet switch involves the exploitation of an unsaturated resource for which few or no other species compete. Using stable isotopes of carbon and nitrogen as dietary tracers, we provide here strong support to the controversial hypothesis that the giant noctule bat Nyctalus lasiopterus feeds on the wing upon the multitude of flying passerines during their nocturnal migratory journeys, a resource which, while showing a predictable distribution in space and time, is only seasonally available. So far, no predator had been reported to exploit this extraordinarily diverse and abundant food reservoir represented by nocturnally migrating passerines.     

Pointed wings, low wingloading and calm air reduce migratory flight costs in songbirds

Migratory bird, bat and insect species tend to have more pointed wings than non-migrants. Pointed wings and low wingloading, or body mass divided by wing area, are thought to reduce energy consumption during long-distance flight, but these hypotheses have never been directly tested. Furthermore, it is not clear how the atmospheric conditions migrants encounter while aloft affect their energy use; without such information, we cannot accurately predict migratory species' response(s) to climate change. Here, we measured the heart rates of 15 free-flying Swainson's Thrushes (Catharus ustulatus) during migratory flight. Heart rate, and therefore rate of energy expenditure, was positively associated with individual variation in wingtip roundedness and wingloading throughout the flights. During the cruise phase of the flights, heart rate was also positively associated with wind speed but not wind direction, and negatively but not significantly associated with large-scale atmospheric stability. High winds and low atmospheric stability are both indicative of the presence of turbulent eddies, suggesting that birds may be using more energy when atmospheric turbulence is high. We therefore suggest that pointed wingtips, low wingloading and avoidance of high winds and turbulence reduce flight costs for small birds during migration, and that climate change may have the strongest effects on migrants' in-flight energy use if it affects the frequency and/or severity of high winds and atmospheric instability.     

Influence of weather conditions on the flight of migrating black storks

This study tested the potential influence of meteorological parameters (temperature, humidity, wind direction, thermal convection) on different migration characteristics (namely flight speed, altitude and direction and daily distance) in 16 black storks (Ciconia nigra). The birds were tracked by satellite during their entire autumnal and spring migration, from 1998 to 2006. Our data reveal that during their 27-day-long migration between Europe and Africa (mean distance of 4100 km), the periods of maximum flight activity corresponded to periods of maximum thermal energy, underlining the importance of atmospheric thermal convection in the migratory flight of the black stork. In some cases, tailwind was recorded at the same altitude and position as the birds, and was associated with a significant rise in flight speed, but wind often produced a side azimuth along the birds'' migratory route. Whatever the season, the distance travelled daily was on average shorter in Europe than in Africa, with values of 200 and 270 km d⁻¹, respectively. The fastest instantaneous flight speeds of up to 112 km h⁻¹ were also observed above Africa. This observation confirms the hypothesis of thermal-dependant flight behaviour, and also reveals differences in flight costs between Europe and Africa. Furthermore, differences in food availability, a crucial factor for black storks during their flight between Europe and Africa, may also contribute to the above-mentioned shift in daily flight speeds.     

Concealed by darkness: interactions between predatory bats and nocturnally migrating songbirds illuminated by DNA sequencing

Recently, several species of aerial‐hawking bats have been found to prey on migrating songbirds, but details on this behaviour and its relevance for bird migration are still unclear. We sequenced avian DNA in feather‐containing scats of the bird‐feeding bat Nyctalus lasiopterus from Spain collected during bird migration seasons. We found very high prey diversity, with 31 bird species from eight families of Passeriformes, almost all of which were nocturnally flying sub‐Saharan migrants. Moreover, species using tree hollows or nest boxes in the study area during migration periods were not present in the bats’ diet, indicating that birds are solely captured on the wing during night‐time passage. Additional to a generalist feeding strategy, we found that bats selected medium‐sized bird species, thereby assumingly optimizing their energetic cost‐benefit balance and injury risk. Surprisingly, bats preyed upon birds half their own body mass. This shows that the 5% prey to predator body mass ratio traditionally assumed for aerial hunting bats does not apply to this hunting strategy or even underestimates these animals’ behavioural and mechanical abilities. Considering the bats’ generalist feeding strategy and their large prey size range, we suggest that nocturnal bat predation may have influenced the evolution of bird migration strategies and behaviour.     

Optimal strategies for insects migrating in the flight boundary layer: mechanisms and consequences

Directed aerial displacement requires that a volant organism'sairspeed exceeds ambient wind speed. For biologically relevantaltitudes, wind speed increases exponentially with increasedheight above the ground. Thus, dispersal of most insects isinfluenced by atmospheric conditions. However, insects thatfly close to the Earth's surface displace within the flightboundary layer where insect airspeeds are relatively high. Overthe past 17 years, we have studied boundary-layer insects byfollowing individuals as they migrate across the Caribbean Seaand the Panama Canal. Although most migrants evade either droughtor cold, nymphalid and pierid butterflies migrate across Panamanear the onset of the rainy season. Dragonflies of the genusPantala migrate in October concurrently with frontal weathersystems. Migrating the furthest and thereby being the most difficultto study, the diurnal moth Urania fulgens migrates between Centraland South America. Migratory butterflies and dragonflies arecapable of directed movement towards a preferred compass directionin variable winds, whereas the moths drift with winds over water.Butterflies orient using both global and local cues. Consistentwith optimal migration theory, butterflies and dragonflies adjusttheir flight speeds in ways that maximize migratory distancetraveled per unit fuel, whereas the moths do not. Moreover,only butterflies adjust their flight speed in relation to endogenousfat reserves. It is likely that these insects use optic flowto gauge their speed and drift, and thus must migrate wheresufficient detail in the Earth's surface is visible to them.The abilities of butterflies and dragonflies to adjust theirairspeed over water indicate sophisticated control and guidancesystems pertaining to migration.     

Evolution of nocturnal flight calls in migrating wood-warblers: apparent lack of morphological constraints

Many migratory songbirds produce flight calls that vary, sometimes strikingly, among species in duration, frequency, modulation, and pattern, and little is known about what factors are responsible for this variation. Negative correlations between body mass or bill length and song frequencies are well documented in birds, but no studies have examined these associations for flight calls. We explored relationships between flight call and song frequencies and body mass or bill length in 33 species of wood-warblers. Phylogenetically controlled and uncontrolled analyses showed no significant relationship between body mass or bill length and flight call frequency; however, we found significant differences between flight call and song frequencies and a significant relationship between maximum frequencies of flight calls and songs. Our findings suggest that factors other than body mass and bill length are responsible for variation in flight call frequencies in wood-warblers. We suggest that different ecological and atmospheric properties might play important roles in selection for flight call frequencies.     

Physical and physiological costs of ectoparasitic mites on host flight endurance

1. Dispersal is essential for locating mates, new resources, and to escape unfavourable conditions. Parasitism can impact a host's ability to perform energetically demanding activities such as long‐distance flight, with important consequences for gene flow and meta‐population dynamics. 2. Ectoparasites, in particular, can adversely affect host flight performance by diminishing flight aerodynamics and/or by inflicting physiological damage while feeding on host tissue. 3. Experimental flight assays were conducted using two fruit fly‐mite systems: Drosophila nigrospiracula (Patterson and Wheeler) – Macrocheles subbadius (Berlese) and D. hydei (Sturtevan) – M. muscaedomesticae (Scopoli). Flies that are burdened by mites are expected to exhibit lower flight endurance compared to uninfected flies. 4. The results show that the presence of mites (attached) significantly decreased flight endurance by 57% and 78% compared to uninfected D. nigrospiracula and D. hydei, respectively. The physiological damage caused by M. subbadius was revealed through a 53% decline in flight time among previously infected flies (mites removed just prior to flight assay). Surprisingly, the presumably phoretic M. muscaedomesticae also caused a 62% reduction in flight endurance among previously infected D. hydei. 5. These results suggest a strong deleterious effect of ectoparasitic mites on host flight performance, mediated by a reduction in flight aerodynamics and damage to host physiology. Adverse effects on host flight and/or dispersal may have broad implications for gene flow, population genetic structure, and local adaptation in both host and parasite meta‐populations.     

Substrate utilization during endurance exercise in men and women after endurance training

We investigated the effect of endurance training on whole body substrate, glucose, and glycerol utilization during 90 min of exercise at 60% peak O2 consumption (VO2(peak)) in males and females. Substrate oxidation was determined before and after 7 wk of endurance training on a cycle ergometer, with posttesting performed at the same absolute (ABS, W) and relative (REL, VO2(peak)) intensities. [6,6-2H]glucose and [1,1,2,3,3-2H]glycerol tracers were used to calculate the respective substrate tracee flux. Endurance training resulted in an increase in VO2(peak) for both males and females of 17 and 22%, respectively (P < 0.001). Females demonstrated a lower respiratory exchange ratio (RER) both pretraining and posttraining compared with males during exercise (P < 0.001). Glucose rate of appearance (R(a)) and rate of disappearance (R(d)) were not different between males and females. Glucose metabolic clearance rate (MCR) was lower at 75 and 90 min of exercise for females compared with males (P < 0.05). Glucose R(a) and R(d) were lower during exercise at both ABS and REL posttraining exercise intensities compared with pretraining (P < 0.001). Females had a higher exercise glycerol R(a) and R(d) compared with males both pre- and posttraining (P < 0.001). Glycerol R(a) was not different at either the ABS or REL posttraining exercise intensities compared with pretraining. We concluded that females oxidize proportionately more lipid and less carbohydrate during exercise compared with males both pre- and posttraining, which was cotemporal with a higher glycerol R(a) in females. Furthermore, endurance training resulted in a decrease in glucose flux at both ABS and REL exercise intensities after endurance exercise training.     

Flight strategies of migrating raptors; a comparative study of interspecific variation in flight characteristics

The comparison of flight styles and flight parameters of migrating raptors in Israel revealed the following. (1) Climbing rate in thermal circling did not differ between species, indicating that chiefly the strength of thermal updrafts determined the climbing rate and that morphological features were less relevant. (2) In interthermal gliding, air speed was positively and gliding angle negatively related to the species' average body mass. Heavier species glided faster and had smaller gliding angles. (3) In soaring and gliding flight, cross-country speed relative to the air was positively related to the species' body mass; it was obviously the result of the gliding ability increasing with body mass. (4) Eagles and buzzards used soaring and gliding flight for more than 95% of the observation time. Additional soaring in a straight line whilst gliding was extensively used by the Steppe Eagle Aquila nipalensis, Lesser Spotted Eagle Aquila pomarina and Booted Eagle Hieraætus pennatus and even more frequently by the resident species, the Griffon Vulture Gyps fulvus and Shorttoed Eagle Circaetus gallicus. Smaller species, such as the Levant Sparrowhawk Accipiter brevipes, harriers (Circus sp.) and small falcons (Falco sp.). showed the highest proportion of flapping and gliding flight (9–33%). (5) In a comparison of the flight parameters and proportions of flight styles, a cluster analysis distinguished two main groups: The first consisted of Montagu's Harrier Circus pygargus, Pallid Harrier Circus macrourus, Levant Sparrowhawk and small falcons; their flight behaviour was characterized by both the high proportion of flapping and the low gliding performance. The second group comprised the typical soaring migrants: Steppe Eagle, Lesser Spotted Eagle, Booted Eagle, Steppe Buzzard Buteo buteo vulpinus, Honey Buzzard Pernis apivorus and Egyptian Vulture Neophron percnopterus, and they had very similar flight behaviour and were closely clustered. The Black Kite Milvus migrans and Marsh Harrier Circus aeruginosus were intermediate between typical soarers and flappers. The two resident species, Griffon Vulture and Short-toed Eagle, were grouped separately from the soaring migrants.     

No small feat: microRNA responses during vocal communication in songbirds

Simply hearing the song produced by another bird of the same species triggers the regulation of microRNAs (miRNAs) in high-order auditory parts of the zebra finch brain. Some of the identified miRNAs appear to be unique to birds, possibly to songbirds. These findings, reported in BMC Genomics, highlight the complexities of gene regulation associated with vocal communication and point to possible key regulators of song-triggered gene networks.     

Protein metabolism during endurance exercise

After reviewing all the available results from our laboratory and numerous reports in the literature concerning changes that have occurred in various aspects of protein metabolism during exercise, a number of conclusions can be drawn with some degree of confidence. During exercise, protein synthesis is depressed and this change leaves amino acids available for catabolic processes. The rate of leucine oxidation appears to be increased during exercise, and there is a movement of amino acids, mostly in the form of alanine, from muscle to liver where the rate of gluconeogenesis is increased as a result of exercise. These changes in protein metabolism are probably physiologically significant in at least three ways: amino acid conversion to citric acid cycle intermediates enhances the rate of oxidation of acetyl-CoA generated from glucose and fatty acid oxidation; increased conversion of amino acids to glucose helps to prevent hypoglycemia; oxidation of some amino acids may provide energy for muscular contraction.     

Substrate utilization and flight speed during tethered flight in the locust

Mature laboratory locusts normally exhibit a characteristic pattern of change in flight speed with time. They fly at high speed for the first few minutes, during which carbohydrate forms the major fuel, but then slow to a cruising speed when lipid is used almost exclusively. Locusts flown for 30 min, rested for 2hr, and then reflown, exhibit an identical pattern of flight, even though they oxidise only half the amount of carbohydrate used in the first flight. The injection of adipokinetic hormone before the first flight elicits a low initial flight speed for 10 to 15 min but then the locusts accelerate to a constant higher speed. The injection of hormone before the second flight, when blood lipid levels are already high, reduces the utilization of carbohydrate by the flight muscles dramatically but results in constant high-speed flight.     

Morning flight behavior of nocturnally migrating birds along the western basin of Lake Erie

Many species of birds that normally migrate during the night have been observed engaging in so‐called morning flights during the early morning. The results of previous studies have supported the hypothesis that one function of morning flights is to compensate for wind drift that birds experienced during the night. Our objective was to further explore this hypothesis in a unique geographic context. We determined the orientation of morning flights along the southern shore of Lake Erie's western basin during the spring migrations of 2016 and 2017. This orientation was then compared to the observed orientation of nocturnal migration. Additionally, the orientation of the birds engaged in morning flights following nights with drifting winds was compared to that of birds following nights with non‐drifting winds. The morning flights of most birds at our observation site were oriented to the west‐northwest, following the southern coast of Lake Erie. Given that nocturnal migration was oriented generally east of north, the orientation of morning flight necessarily reflected compensation for accumulated, seasonal wind drift resulting from prevailingly westerly winds. However, the orientation of morning flights was similar following nights with drifting and non‐drifting winds, suggesting that birds on any given morning were not necessarily re‐orienting as an immediate response to drift that occurred the previous night. Given the topographical characteristics of our observation area, the west‐northwest movement of birds in our study is likely best explained as a more complex interaction that could include some combination of compensation for wind drift, a search for suitable stopover habitat, flying in a direction that minimizes any loss in progressing northward toward the migratory goal, and avoidance of a lake crossing.