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Sarco(endo)plasmic reticulum Ca 2+ -ATPase (SERCA) is a transmembrane pump critical to muscle calcium cycling during contraction, and SERCA has also been proposed as the basis for a non-shivering thermogenesis mechanism in birds. Despite its potential importance to both shivering and non-shivering thermogenesis, the activity of this transporter has rarely been studied in altricial birds, and never during the developmental transition from ectothermy to endothermy. Here, we describe SERCA activity in the pectoralis muscle and heart ventricle of red-winged blackbird ( Agelaius phoeniceus ) nestlings, fledglings and adults. Additionally, using a diet manipulation, we tested the hypothesis that muscle SERCA activity is affected by dietary fatty acid composition, as has been shown in some previous studies. In blackbird hearts, SERCA activity increased throughout development and into adulthood, conspicuously jumping higher just prior to fledging. In pectoralis muscle, SERCA activity increased throughout the nestling period, but then declined after fledging, an effect we attribute to remodeling of the muscle from a primarily heat-generating organ to a primarily force-generating organ. SERCA activity of the pectoralis muscle was correlated with the proportion of linoleic acid in muscle phospholipids when including all ages in the control group. However, in diet-manipulated birds, there was no consistent relationship between SERCA activity and muscle membrane fatty acid composition at any tested age (5-9 days old). It is unclear whether SERCA might be affected by developmental changes in fatty acid composition at younger ages., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Docosahexaenoic acid (DHA) is an important and abundant fatty acid moiety in vertebrate brains. We measured brain phospholipid composition during development in red-winged blackbirds ( Agelaius phoeniceus ), an altricial species that breeds in aquatic habitats. We also manipulated diet by feeding nestlings fish oil or sunflower oil. Finally, we assessed selective uptake of yolk by comparing the yolk fatty acid composition of freshly laid eggs and day-old hatchlings. Relative to other altricial species, blackbirds achieved high DHA in brain phospholipids (20% of phospholipid fatty acids in day-old hatchlings). This was not a result of selective uptake from the yolk, but rather a consequence of a high proportion of DHA in the yolk (2.5% of total lipids) at laying. Our dietary study confirmed that nestling brains are sensitive to fatty acid supply. Red-winged blackbirds may be able to advance cognitive development relative to other altricial species owing to their aquatic maternal diet., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The 'membrane pacemaker' hypothesis proposes a biochemical explanation for among-species variation in resting metabolism, based on the positive correlation between membrane docosahexaenoic acid (DHA) and metabolic rate. We tested this hypothesis using a novel model, altricial red-winged blackbird nestlings, predicting that the proportion of DHA in muscle and liver membranes should increase with the increasing metabolic rate of the nestling as it develops endothermy. We also used a dietary manipulation, supplementing the natural diet with fish oil (high DHA) or sunflower oil (high linoleic acid) to alter membrane composition and then assessed metabolic rate. In support of the membrane pacemaker hypothesis, DHA proportions increased in membranes from pectoralis muscle, muscle mitochondria and liver during post-hatch development. By contrast, elevated dietary DHA had no effect on resting metabolic rate, despite causing significant changes to membrane lipid composition. During cold challenges, higher metabolic rates were achieved by birds that had lower DHA and higher linoleic acid in membrane phospholipids. Given the mixed support for this hypothesis, we conclude that correlations between membrane DHA and metabolic rate are likely spurious, and should be attributed to a still-unidentified confounding variable., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Life-history theory predicts that animals face a trade-off in energy allocation between performing strenuous exercise, such as migratory flight, and mounting an immune response. We experimentally tested this prediction by studying immune function in European starlings, Sturnus vulgaris, flown in a wind tunnel. Specifically, we predicted that constitutive immune function decreases in response to training and, additionally, in response to immediate exercise. We compared constitutive immune function among three groups: (1) 'untrained' birds that were kept in cages and were not flown; (2) 'trained' birds that received flight training over a 15 day period and performed a 1-4 h continuous flight, after which they rested for 48 h before being sampled; and (3) 'post-flight' birds that differed from the 'trained' group only in being sampled immediately after the final flight. A bird in our trained group represents an individual during migration that has been resting between migratory flights for at least 2 days. A bird in our post-flight group represents an individual that has just completed a migratory flight and has not yet had time to recover. Three of our four indicators (haptoglobin, agglutination and lysis) showed the predicted decrease in immune function in the post-flight group, and two indicators (haptoglobin, agglutination) showed the predicted decreasing trend from the untrained to trained to post-flight group. Haptoglobin levels were negatively correlated with flight duration. No effect of training or flight was detected on leukocyte profiles. Our results suggest that in European starlings, constitutive immune function is decreased more as a result of immediate exercise than of exercise training. Because of the recent emergence of avian-borne diseases, understanding the trade-offs and challenges faced by long-distance migrants has gained a new level of relevance and urgency.

Birds primarily fuel migratory flights with fat, and the composition of that fat has the potential to affect overall lipid oxidation rates. We measured the whole muscle lipid oxidation rates in extensor digitorum communis muscles from white-throated sparrows (Zonotrichia albicollis Gmelin) incubated for 20 min at 20°C with radiolabeled stearate (18:0), oleate (18:1ω9), or linoleate (18:2ω6). Lipid oxidation rates were ~40% higher with linoleate than oleate (oleate: 36 ± 8.54 μmol CO(2) g(-1) h(-1)), and ~75% lower with stearate compared with oleate, indicating that maximal lipid oxidation rates can indeed be affected by the type of fatty acid supplied to the muscle. Additionally, we investigated the activity of the mitochondrial fatty acid transport-associated enzyme carnitine palmitoyl transferase (CPT) in pectoralis muscles of 5 bird species (Zonotrichia albicollis, Philomachus pugnax, Sturnus vulgaris, Taeniopygia guttata, Passer domesticus). Activity was measured in homogenized samples using various fatty acyl-CoA substrates (16:0, 16:1, 18:0, 18:1ω9, 18:2ω6, 18:3ω3, 18:3ω6, 20:0, 20:4ω6, 22:6ω3) in a spectrophotometric assay. CPT activity increased with the degree of unsaturation and decreased with chain length. CPT activity did not differ between ω3 and ω6 isomers of 18:3, nor was the pattern of CPT substrate preference different between captive white-throated sparrows in a migratory (i.e., displaying Zugunruhe) or non-migratory state. These findings can explain previously observed differences in peak performance induced by dietary fat composition and suggest that lipid supply is limiting to maximal exercise performance in birds.

The extreme energetic demands of avian migration result in various physiological changes that can be observed during the migratory period. However, the degree to which birds alter muscle physiology in advance of migration has been poorly studied. We studied the effects of "migratory" photoperiod and exercise on metabolic enzymes, fatty acid transporter mRNA expression, and muscle phospholipid fatty acid composition in captive white-crowned sparrows (Zonotrichia leucophrys). Ten sparrows were held on short photoperiod (8L:16D) for 58 d then switched to long days (16L:8D) for 3 wk before sampling. Increased nightly activity indicated that the birds were indeed in migratory condition. Another 13 birds were held on short days during the entire experiment, and a subset (5) were exercised for 1 h every other day for the last 2 wk. "Migratory" photoperiod did not change the activities of citrate synthase, carnitine palmitoyl transferase, and 3-hydroxyacyl-CoA dehydrogenase or the expression of FAT/CD36, FABPpm, and H-FABP mRNA in pectoralis muscle, suggesting that these cannot be increased in advance of migratory flight. Docosahexaenoic acid increased in pectoralis muscle phospholipids with exercise but was negatively correlated with catabolic enzyme activity, indicating that the presence of this fatty acid may not aid migratory performance as suggested by other studies.