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The adrenocorical stress-response of Black-legged Kittiwake chicks in relation to dietary restrictions

In this study we examined hormonal responses of Black-legged Kittiwake ( Rissatridactyla ) chicks to experimental variations in energy content and nutritional quality (low or high lipid to protein ratio, LPR) of their food. Starting at the age of 10 days, chicks were fed either high or low LPR fish at 30, 50, 70 and 100% of ad libitum energy intake. After 20 days of treatment, chicks were exposed to a standardized acute handling and restraint stress protocol, where a baseline sample was taken immediately after taking a chick from the nest, and three additional blood samples were taken at intervals up to 50 min. Testosterone and corticosterone titres in plasma were measured via radioimmunoassay. We found that baseline testosterone levels were not significantly affected by the experimental treatments. Food-restricted chicks had elevated baseline and acute stress-induced levels of corticosterone compared to chicks fed ad libitum. An elevation of circulating levels of corticosterone in energetically stressed individuals was further magnified by low nutritional quality of food. Baseline and acute stress-induced corticosterone levels of chicks were negatively correlated with their fat reserves. We conclude that the physiological condition of Black-legged Kittiwake chicks can be assessed reliably by measuring circulating levels of corticosterone. We discuss short- and long-term effects of elevated corticosterone secretion in food-stressed nest-bound chicks.

Journal of Comparative Physiology B: Biochemical, ↗

Understanding behavioral responses of fish to pheromones in natural freshwater environments

There is an abundance of experimental studies and reviews that describe odorant-mediated behaviors of fish in laboratory microcosms, but research in natural field conditions has received considerably less attention. Fish pheromone studies in laboratory settings can be highly productive and allow for controlled experimental designs; however, laboratory tanks and flumes often cannot replicate all the physical, physiological and social contexts associated with natural environments. Field experiments can be a critical step in affirming and enhancing understanding of laboratory discoveries and often implicate the ecological significance of pheromones employed by fishes. When findings from laboratory experiments have been further tested in field environments, often different and sometimes contradictory conclusions are found. Examples include studies of sea lamprey (Petromyzon marinus) mating pheromones and fish alarm substances. Here, we review field research conducted on fish pheromones and alarm substances, highlighting the following topics: (1) contradictory results obtained in laboratory and field experiments, (2) how environmental context and physiological status influences behavior, (3) challenges and constraints of aquatic field research and (4) innovative techniques and experimental designs that advance understanding of fish chemical ecology through field research.

Journal of Comparative Physiology A↗

Fatty acid signatures of stomach oil and adipose tissue of northern fulmars (Fulmarus glacialis) in Alaska: Implications for diet analysis of Procellariiform birds

Procellariiforms are unique among seabirds in storing dietary lipids in both adipose tissue and stomach oil. Thus, both lipid sources are potentially useful for trophic studies using fatty acid (FA) signatures. However, little is known about the relationship between FA signatures in stomach oil and adipose tissue of individuals or whether these signatures provide similar information about diet and physiology. We compared the FA composition of stomach oil and adipose tissue biopsies of individual northern fulmars ( N = 101) breeding at three major colonies in Alaska. Fatty acid signatures differed significantly between the two lipid sources, reflecting differences in dietary time scales, metabolic processing, or both. However, these signatures exhibited a relatively consistent relationship between individuals, such that the two lipid sources provided a similar ability to distinguish foraging differences among individuals and colonies. Our results, including the exclusive presence of dietary wax esters in stomach oil but not adipose tissue, are consistent with the notion that stomach oil FA signatures represent lipids retained from prey consumed during recent foraging and reflect little metabolic processing, whereas adipose tissue FA signatures represent a longer-term integration of dietary intake. Our study illustrates the potential for elucidating short- versus longer-term diet information in Procellariiform birds using different lipid sources.

Alaska↗

Increased daylength stimulates plasma growth hormone and gill Na+, K+ and -ATPase Atlantic salmon (Salmo salar )

Atlantic salmon juveniles reared at constant temperature (9–10°C) were exposed to four photoperiod treatment and sampled every 2 weeks from January through May. Fish reared under normal photoperiod exhibited eight-and three fold increases in plasma growth hormone and gill Na+, K+-ATPase activity, respectively, between January and April. Fish exposed to abrupt increases in daylength (LD 15:9) in February or March responded with earlier increases in plasma growth hormone and gill Na+, K+-ATPase activity, and earlier decreases in condition factor relative to fish in the normal photoperiod group. Fish maintained under short daylength (LD 9:15) from January to May exhibited delayed and muted increases in plasma growth hormone and gill Na+, K+-ATPase activity. Plasma thyroxine exhibited a 2.5-fold increase from February to late March in the normal photoperiod group, was generally lower in the LD 9:15 group, but exhibited no obvious response to abrupt increases in daylength. There was an increase in plasma 3,5,3′-triiodo-l-thyronine with time in all groups (43–80%) but no significant response to photoperiod. Plasma levels of somatostatin-25 were highest in the LD 9:15 group, but there was no detectable response to increased daylength in any of the photoperiod treatments. The results indicate that plasma growth hormone is responsive to increased daylength and may be causally related to subsequent increases in gill Na+, K+-ATPase.

Journal of Comparative Physiology B: Biochemical, ↗

Auditory evoked potentials in the West Indian Manatee (Sirenia: Trichechus manatus )

Potentials evoked by clicks and tone pips were recorded by fine wires inserted extracranially in four West Indian manatees ( Trichechus manatus ) in air. Sounds were delivered via padded ear phones. Averaging a few thousand trials at 20/s reveals early peaks at N5.4 (‘vertex’ negativity to a frontal reference, at 5.4 ms), P7.6, N8.8, P9.5 — probably equivalent to waves IV and VII of the typical mammalian auditory brainstem response (ABR). Averaging 100 trials at <4/s suffices to reveal a complex sequence of later peaks including N25, P80, N150 and P190; consistent smaller peaks are visible when several hundred trials are averaged. Using tone pips with a rise and fall time of 2–5 ms the carrier frequency becomes important. Evoked potential wave forms are not the same at different frequencies, bringing out the fact that frequency is not a scalar that can be compensated for by intensity. Therefore the method was not used to obtain audiograms; however the largest EPs occur in the range of 1–1.5 kHz. EPs are found up to 35 kHz; almost no evoked potential is discernible at 40 kHz but the undistorted intensity available was limited. This is in reasonable agreement with the theoretical expectation for the upper limit of behavioral hearing from Heffner and Masterton based on head size and aquatic medium. Among several ear phone placements, that over the external auditory meatus was the most effective, but only slightly so. The external canal is presumably fluid or tissue filled and sound enters over a large area. Comparing data for two species on the most effective range of frequencies and the power spectra of their vocalizations, T. manatus is lower than T. inunguis in both respects. The results show the utility and limitations of the method of recording extracranial evoked potentials to sounds, especially for large and valuable animals under makeshift conditions.

Journal of Comparative Physiology A↗

The orientation and navigation of juvenile alligators: evidence of magnetic sensitivity

Displaced juvenile alligators, Alligator mississipiensis , were released on land in a 9 m diameter dodecagonal arena to test their ability to orient in the absence of terrestrial landmarks. Navigational ability seemed to improve with age. When displaced along a fairly direct route yearlings (age 7–14 months) compensated for their displacement, moving in the direction from the arena to their home sites. When displaced by a circuitous route, yearlings failed to compensate for their displacement, exhibiting instead simple compass orientation in a direction that would have returned them to water had they been released on land near the site where they were captured. The older juveniles were oriented in a homeward direction under all displacement and test conditions. The latter animals may have been using geomagnetic map information to select their homeward directions as the errors in their homeward bearings correlated with small deviations in the geomagnetic field's dip angle at the time of the test (1980 r s =−0.6047, P =0.0131, all tests r s = −0.4652, P =0.0084). This effect appeared to depend on a very short-term assessment of geomagnetic conditions, as values measured 20 min before or 30 min after the tests began did not correlate with the directions the animals moved. The older juveniles appeared to use magnetically quiet hours on the night of their capture as the baseline from which to measure the geomagnetic deviations that occurred at the time of the arena test. The magnitude of the magnetic effect in the older animals suggests that the geomagnetic information may have been used to perform a ‘map’ step, as small fluctuations in dip angle correlated with much larger deviations in homeward bearings. In addition, the compass-oriented yearlings and the seemingly route-based behavior of the homeward-oriented yearlings did not appear to be influenced by geomagnetic conditions. These findings have many parallels in results obtained from bird orientation studies, providing evidence that navigation may share a common basis in different vertebrate groups.

Journal of Comparative Physiology A↗

A comparison of auditory brainstem responses across diving bird species

There is little biological data available for diving birds because many live in hard-to-study, remote habitats. Only one species of diving bird, the black-footed penguin ( Spheniscus demersus ), has been studied in respect to auditory capabilities (Wever et al., Proc Natl Acad Sci USA 63:676&ndash;680, 1969). We, therefore, measured in-air auditory threshold in ten species of diving birds, using the auditory brainstem response (ABR). The average audiogram obtained for each species followed the U-shape typical of birds and many other animals. All species tested shared a common region of the greatest sensitivity, from 1000 to 3000 Hz, although audiograms differed significantly across species. Thresholds of all duck species tested were more similar to each other than to the two non-duck species tested. The red-throated loon ( Gavia stellata ) and northern gannet ( Morus bassanus ) exhibited the highest thresholds while the lowest thresholds belonged to the duck species, specifically the lesser scaup ( Aythya affinis ) and ruddy duck ( Oxyura jamaicensis ). Vocalization parameters were also measured for each species, and showed that with the exception of the common eider ( Somateria mollisima ), the peak frequency, i.e., frequency at the greatest intensity, of all species' vocalizations measured here fell between 1000 and 3000 Hz, matching the bandwidth of the most sensitive hearing range.

Journal of Comparative Physiology A↗

Seasonal changes in atmospheric noise levels and the annual variation in pigeon homing performance

Repeated releases of experienced homing pigeons from single sites were conducted between 1972 and 1974 near Cornell University in upstate New York and between 1982 and 1983 near the University of Pittsburgh in western Pennsylvania, USA. No annual variation in homing performance was observed at these sites in eastern North America, in contrast to results from a number of similar experiments in Europe. Assuming pigeons home using low-frequency infrasonic signals (~0.1–0.3 Hz), as has been previously proposed, the annual and geographic variability in homing performance within the northern hemisphere might be explained, to a first order, by seasonal changes in low-frequency atmospheric background noise levels related to storm activity in the North Atlantic Ocean, and by acoustic waveguides formed between the surface and seasonally reversing stratospheric winds. In addition, increased dispersion among departure bearings of test birds on some North American release days was possibly caused by infrasonic noise from severe weather events during tornado and Atlantic hurricane seasons.

Journal of Comparative Physiology A↗

A reinterpretation of “Homing pigeons’ flight over and under low stratus” based on atmospheric propagation modeling of infrasonic navigational cues

Pigeons flying above temperature inversion and related low-stratus layers appear to lack important navigational cues, and a reinterpretation of Wagner’s 1978 study suggests that these cues are low-frequency acoustic signals (infrasound). Wagner released homing pigeons above opaque stratus over the Swiss Plateau to determine whether they could locate their loft beneath it. Birds above the clouds appeared lost, while those that descended beneath them returned home directly. Atmospheric propagation modeling of infrasonic waves virtually transmitted from the loft area shows that these signals would have been ducted beneath the inversion layer, and would not have reached the release sites above it. The absence of homeward infrasonic cues above temperature inversions could explain the disorientation of Wagner’s birds, especially if such signals are the predominant cues used by pigeons to home. The possible generation of infrasonic navigational signals in the loft area and recent queries concerning the infrasound navigational “map” hypothesis are also discussed.

Journal of Comparative Physiology A↗

Avian navigation: Comparing the olfactory navigational “map” and the infrasound direction-finding hypotheses to aeronautics

Animal navigation has long been a fascinating but bewildering subject. Humans and animals might well share similar navigational strategies because they developed within the same physical environments. A “map-and-compass” model has been proposed to explain the two-step avian navigational process, but the “map” step has remained elusive. Although scalar values from bicoordinate geomagnetic or atmospheric olfactory gradients have been considered foundational to the avian map, neither has proved convincing engendering decades of controversy. The olfactory map, and an alternative infrasound direction-finding (IDF) hypothesis, are discussed in this review. The olfactory map hypothesis currently requires extensive stable gradients of trace-odor ratios, but such gradients are highly unlikely within a turbulent and rapidly mixed lower atmosphere. The IDF hypothesis, on the other hand, postulates a two-step navigational model analogous to the maritime and aeronautical radio direction-finding technique. This review was also written to encourage further investigation, and direct testing, of the acoustic navigational process. The IDF hypothesis, at present, appears the better explanation of observed avian navigational behavior and accuracy within the atmosphere’s physical environment.

Journal of Comparative Physiology A↗

Intermittent fasting during winter and spring affects body composition and reproduction of a migratory duck

We compared food intake, body mass and body composition of male and female black ducks (Anas rubripes) during winter (January-March). Birds were fed the same complete diet ad libitum on consecutive days each week without fasting (control; nine male; nine female) or with either short fasts (2 day.week-1; nine male; nine female), or long fasts (4 day.week-1; eleven male; twelve female). We continued treatments through spring (March-May) to measure the effect of intermittent fasts on body mass and egg production. Daily food intake of fasted birds was up to four times that of unfasted birds. Weekly food intake of males was similar among treatments (364 g.kg-1.week-1) but fasted females consumed more than unfasted females in January (363 g.kg-1.week-1 vs. 225 g.kg-1.week-1). Although both sexes lost 10-14% body mass, fasted females lost less mass and lipid than unfasted females during winter. Total body nitrogen was conserved over winter in both sexes even though the heart and spleen lost mass while the reproductive tract and liver gained mass. Intermittent fasting increased liver, intestinal tissue and digesta mass of females but not of males. Fasting delayed egg production in spring but did not affect size, fertility or hatching of the clutch. Females on long fasts were still heavier than controls after laying eggs. Thus black ducks combine flexibility of food intake with plasticity of digestive tract, liver and adipose tissue when food supply is interrupted during winter. Females modulate body mass for survival and defer reproduction when food supply is interrupted in spring.

Journal of Comparative Physiology B: Biochemical, ↗

Do geese fully develop brood patches? A histological analysis of lesser snow geese (Chen caerulescens caerulescens) and Ross's geese (C. rossii)

Most birds develop brood patches before incubation; epidermis and dermis in the brood patch region thicken, and the dermal connective tissue becomes increasingly vascularized and infiltrated by leukocytes. However, current dogma states that waterfowl incubate without modifications of skin within the brood patch region. The incubation periods of lesser snow geese (Chen caerulescens caerulescens; hereafter called snow geese) and Ross's geese (C. rossii) are 2-6 days shorter than those of other goose species; only females incubate. Thus, we hypothesized that such short incubation periods would require fully developed brood patches for sufficient heat transfer from incubating parents to eggs. We tested this hypothesis by analyzing the skin histology of abdominal regions of snow and Ross's geese collected at Karrak Lake, Nunavut, Canada. For female snow geese, we found that epidermis and dermis had thickened and vascularization of dermis was 14 times greater, on average, than that observed in males (n=5 pairs). Our results for Ross's geese (n=5 pairs) were more variable, wherein only one of five female Ross's geese fully developed a brood patch. Our results are consistent with three hypotheses about brood patch development and its relationship with different energetic cost-benefit relationships, resulting from differences in embryonic development and body size. ?? Springer-Verlag 2006.

Journal of Comparative Physiology B: Biochemical, ↗

Data logging of body temperatures provides precise information on phenology of reproductive events in a free-living arctic hibernator

Precise measures of phenology are critical to understanding how animals organize their annual cycles and how individuals and populations respond to climate-induced changes in physical and ecological stressors. We show that patterns of core body temperature ( T b ) can be used to precisely determine the timing of key seasonal events including hibernation, mating and parturition, and immergence and emergence from the hibernacula in free-living arctic ground squirrels ( Urocitellus parryii ). Using temperature loggers that recorded T b every 20 min for up to 18 months, we monitored core T b from three females that subsequently gave birth in captivity and from 66 female and 57 male ground squirrels free-living in the northern foothills of the Brooks Range Alaska. In addition, dates of emergence from hibernation were visually confirmed for four free-living male squirrels. Average T b in captive females decreased by 0.5–1.0°C during gestation and abruptly increased by 1–1.5°C on the day of parturition. In free-living females, similar shifts in T b were observed in 78% ( n = 9) of yearlings and 94% ( n = 31) of adults; females without the shift are assumed not to have given birth. Three of four ground squirrels for which dates of emergence from hibernation were visually confirmed did not exhibit obvious diurnal rhythms in T b until they first emerged onto the surface when T b patterns became diurnal. In free-living males undergoing reproductive maturation, this pre-emergence euthermic interval averaged 20.4 days ( n = 56). T b -loggers represent a cost-effective and logistically feasible method to precisely investigate the phenology of reproduction and hibernation in ground squirrels.

Journal of Comparative Physiology B: Biochemical, ↗

Monitoring responses to variation in food supply for a migratory waterfowl: American Black Duck (Anas rubripes) in winter

Wintering Black Ducks ( Anas rubripes ) concentrate in wetlands along the Atlantic coast where natural and anthropogenic disturbances have increased over the last 50 years, a period in which the population of Black Ducks has declined. We studied the sensitivity of Black Ducks to perturbations in food supply that often result from disturbances by storms, predators, and people. In the paper, we characterize the responses of captive Black Ducks to shifts in food quality and availability during winter and apply those measures to a comparison of wild birds. Captive ducks that were fed intermittently (3 consecutive days/week) compensated for fasted days to achieve similar body mass and body fat to control birds that were fed every day on both animal- and plant-based diets. However, birds that were fed intermittently expended 15% more energy each day than controls when both groups were fed (536 vs. 464 kJ/kg 0.75 ), which indicates that variable food supply increases the costs of maintenance and thus reduces the number of birds that can be supported on the same resource of food without interruptions to foraging. Egg production was not affected by diet quality provided in spring or by the frequency of feeding during the preceding winter months. Black Ducks lost body fat through winter in captivity and in the wild. Fat stores of birds in New Jersey were greater than those of birds in Maine (13.3 vs. 8.3% of body mass) in January, which reflected the high energy demands of cold temperatures in Maine. Values for ∂ 15 N were greater in Maine than in New Jersey for both red blood cells and plasma, which indicated a consistent diet of marine invertebrates in Maine. Greater isotopic variation in red blood cells indicated that diets were more diverse in New Jersey than in Maine for both ∂ 15 N (9.7 ± 1.1 vs. 11.2 ± 0.4‰) and for ∂ 13 C (− 15.1 ± 2.2 vs. − 13.8 ± 1.4‰). Plasma ∂ 13 C was enriched over red blood cells in wild birds especially those with low fat stores, which suggested birds with low energy stores were shifting diets. Black Ducks can compensate for disturbances in feeding by increasing intakes if they have access to high quality wetlands where they are able to find abundant food. High energy demands at cold temperatures may constrain fat stores and thus the tolerance of feeding disturbances especially at the northern limits of the winter range. We hypothesize that decreasing variation in diet may indicate an increase in vulnerability to disturbance in winter when body fat is low. Recent efforts to assess and improve habitat quality of Black Ducks could be enhanced by monitoring the body composition and diet of birds to assess their vulnerability to disturbances in food supply and energy demands.

Journal of Comparative Physiology B: Biochemical, ↗

Effects of a temperature rise on melatonin and thyroid hormones during smoltification of Atlantic salmon, Salmo salar

Smoltification prepares juvenile Atlantic salmon ( Salmo salar ) for downstream migration. Dramatic changes characterize this crucial event in the salmon’s life cycle, including increased gill Na + /K + -ATPase activity (NKA) and plasma hormone levels. The triggering of smoltification relies on photoperiod and is modulated by temperature. Both provide reliable information, to which fish have adapted for thousands of years, that allows deciphering daily and calendar time. Here we studied the impact of different photoperiod (natural, sustained winter solstice) and temperature (natural, ~ + 4° C) combinations, on gill NKA, plasma free triiodothyronine (T3) and thyroxine (T4), and melatonin (MEL; the time-keeping hormone), throughout smoltification. We also studied the impact of temperature history on pineal gland MEL production in vitro. The spring increase in gill NKA was less pronounced in smolts kept under sustained winter photoperiod and/or elevated temperature. Plasma thyroid hormone levels displayed day–night variations, which were affected by elevated temperature, either independently from photoperiod (decrease in T3 levels) or under natural photoperiod exclusively (increase in T4 nocturnal levels). Nocturnal MEL secretion was potentiated by the elevated temperature, which also altered the MEL profile under sustained winter photoperiod. Temperature also affected pineal MEL production in vitro, a response that depended on previous environmental acclimation of the organ. The results support the view that the salmon pineal is a photoperiod and temperature sensor, highlight the complexity of the interaction of these environmental factors on the endocrine system of S. salar , and indicate that climate change might compromise salmon’s time “deciphering” during smoltification, downstream migration and seawater residence.

Journal of Comparative Physiology B: Biochemical, ↗

Fatty acid profiles of feeding and fasting bears: Estimating calibration coefficients, the timeframe of diet estimates, and selective mobilization during hibernation

Accurate information on diet composition is central to understanding and conserving carnivore populations. Quantitative fatty acid signature analysis (QFASA) has emerged as a powerful tool for estimating the diets of predators, but ambiguities remain about the timeframe of QFASA estimates and the need to account for species-specific patterns of metabolism. We conducted a series of feeding experiments with four juvenile male brown bears ( Ursus arctos ) to (1) track the timing of changes in adipose tissue composition and QFASA diet estimates in response to a change in diet and (2) quantify the relationship between consumer and diet FA composition (i.e., determine “calibration coefficients”). Bears were fed three compositionally distinct diets for 90–120 days each. Two marine-based diets were intended to approximate the lipid content and composition of the wild diet of polar bears ( U. maritimus ). Bear adipose tissue composition changed quickly in the direction of the diet and showed evidence of stabilization after 60 days. During hibernation, FA profiles were initially stable but diet estimates after 10 weeks were sensitive to calibration coefficients. Calibration coefficients derived from the marine-based diets were broadly similar to each other and to published values from marine-fed mink ( Mustela vison ), which have been used as a model for free-ranging polar bears. For growing bears on a high-fat diet, the temporal window for QFASA estimates was 30–90 days. Although our results reinforce the importance of accurate calibration, the similarities across taxa and diets suggest it may be feasible to develop a generalized QFASA approach for mammalian carnivores.

Journal of Comparative Physiology B: Biochemical, ↗

Na+/HCO3- cotransporter 1 (nbce1) isoform gene expression during smoltification and seawater acclimation of Atlantic salmon

The life history of Atlantic salmon ( Salmo salar ) includes an initial freshwater phase (parr) that precedes a springtime migration to marine environments as smolts. The development of osmoregulatory systems that will ultimately support the survival of juveniles upon entry into marine habitats is a key aspect of smoltification. While the acquisition of seawater tolerance in all euryhaline species demands the concerted activity of specific ion pumps, transporters, and channels, the contributions of Na + /HCO 3 − cotransporter 1 (Nbce1) to salinity acclimation remain unresolved. Here, we investigated the branchial and intestinal expression of three Na + /HCO 3 − cotransporter 1 isoforms, denoted nbce1.1 , -1.2a , and -1.2b . Given the proposed role of Nbce1 in supporting the absorption of environmental Na + by ionocytes, we first hypothesized that expression of a branchial nbce1 transcript ( nbce1.2a ) would be attenuated in salmon undergoing smoltification and following seawater exposure. In two separate years, we observed spring increases in branchial Na + /K + -ATPase activity, Na + /K + /2Cl − cotransporter 1 , and cystic fibrosis transmembrane regulator 1 expression characteristic of smoltification, whereas there were no attendant changes in nbce1.2a expression. Nonetheless, branchial nbce1.2a levels were reduced in parr and smolts within 2 days of seawater exposure. In the intestine, gene transcript abundance for nbce1.1 increased from spring to summer in the anterior intestine, but not in the posterior intestine or pyloric caeca, and nbce1.1 and -1.2b expression in the intestine showed season-dependent transcriptional regulation by seawater exposure. Collectively, our data indicate that tissue-specific modulation of all three nbce1 isoforms underlies adaptive responses to seawater.

Journal of Comparative Physiology B: Biochemical, ↗

Metabolic rates in an anadromous clupeid, the American shad ( Alosa sapidissima )

To assess the energetics of migration in an anadromous fish, adult American shad ( Alosa sapidissima ) were swum in a large respirometer at a range of speeds (1.0–2.3 body lengths (BL) s −1 , 13–24 °C). Metabolic rate (M O2 ) was logarithmically related to swimming speed (Bl s −1 ; r 2 = 0.41, slope = 0.23 ± 0.037) and tailbeat frequency (beats × min −1 ; r 2 = 0.52, slope = 0.003 ± 0.0003). Temperature had a significant effect on metabolic rate ( r 2 = 0.41) with a Q 10 of 2.2. Standard metabolic rate (SMR), determined directly after immobilization with the neuroblocker gallamine triethiodide, ranged from 2.2–6.2 mmolO 2 kg −1 h −1 and scaled with mass (W) such that SMR = 4.0 (±0.03)W 0.695(±0.15) . Comparison of directly determined and extrapolated SMR suggests that swimming respirometry provides a good estimate of SMR in this species, given the differences in basal activity monitored by the two methods. Overall, American shad metabolic rates (M O2 and SMR) were intermediate between salmonids and fast-swimming perciforms, including tunas, and may be a result of evolutionary adaptation to their active pelagic, schooling life history. This study demonstrates variability in metabolic strategy among anadromous fishes that may be important to understanding the relative success of different migratory species under varying environmental conditions.

Journal of Comparative Physiology B: Biochemical, ↗