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Seasonal adjustments to muscle size in migratory birds may result from preparatory physiological changes or responses to changed workloads. The mechanisms controlling these changes in size are poorly understood. We investigated some potential mediators of flight muscle size (myostatin and insulin-like growth factor, IGF1) in pectoralis muscles of wild wintering or migrating white-throated sparrows (Zonotrichia albicollis), captive white-throated sparrows that were photoperiod manipulated to be in a `wintering' or `migratory' (Zugunruhe) state, and captive European starlings (Sturnus vulgaris) that were either exercised for 2 weeks in a wind tunnel or untrained. Flight muscle size increased in photo-stimulated `migrants' and in exercised starlings. Acute exercise but not long-term training caused increased expression of IGF1, but neither caused a change in expression of myostatin or its metalloprotease activator TLL1. Photo-stimulated `migrant' sparrows demonstrated increased expression of both myostatin and IGF1, but wild sparrows exhibited no significant seasonal changes in expression of either myostatin or IGF1. Additionally, in both study species we describe several splice variants of myostatin that are shared with distantly related bird species. We demonstrate that their expression patterns are not different from those of the typical myostatin, suggesting that they have no functional importance and may be mistakes of the splicing machinery. We conclude that IGF1 is likely to be an important mediator of muscle phenotypic flexibility during acute exercise and during endogenous, seasonal preparation for migration. The role of myostatin is less clear, but its paradoxical increase in photo-stimulated `migrants' may indicate a role in seasonal adjustments of protein turnover.

Birds rely substantially on fat to fuel migratory flights. The importance of the composition of those fat stores to flight performance is a field of recent interest. Here I review the evidence that dietary lipid fat composition affects exercise in birds. Seasonal changes in adipose composition and diet choice experiments have yet to provide strong evidence that fatty acid composition can affect flight performance. Direct manipulations of dietary fat, however, have been demonstrated to affect exercise performance in both avian and non-avian species. I also describe the major hypotheses for the mechanisms by which dietary fat affects exercise, focusing on the role of fatty acids as oxidative substrates and as structural components of membranes. Evidence is accumulating that fatty acids that are shorter or have more double bonds increase peak performance due to their higher transport rates en route to oxidation. Endurance and efficiency of flight may or may not be affected in similar ways. Other mechanisms requiring further investigation include membrane composition, peroxisome proliferator-activated receptors, and eicosanoid-mediated inflammation. Finally, I develop a theoretical framework for studying the composition of fat stores in migrants, focusing on the tradeoffs between fatty acid transport rates, energy storage, and assimilation during stopover refueling., (Copyright © 2010 Elsevier Inc. All rights reserved.)

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.

The fatty acid composition of muscle membrane phospholipids and fat stores may affect migration performance in birds. The purpose of this study was to investigate seasonal changes in the fatty acid composition of (1) pectoralis muscle phospholipids, (2) intramuscular triglyceride stores and (3) adipose tissue triglycerides in free-living white-throated sparrows (Zonotrichia albicollis). During migratory seasons there was an increase in the n-6:n-3 ratio of muscle membrane phospholipid fatty acids without a change in the proportion of unsaturated fatty acids. This change was driven mainly by an increase in the proportion of 18:2n-6 and a decrease in the proportion of 22:6n-3. An increase in the proportion of 18:2n-6 was also observed in the intramuscular and adipose tissue triglyceride stores during the migratory seasons. These increases in 18:2n-6 were offset by a decrease in 16:0; resulting in an elevated proportion of unsaturated fatty acids and elevated double bond index in both fat stores of migrants. The elevated levels of 18:2n-6 in migrant fat stores indicates a high dietary component of this fatty acid, as white-throated sparrows feed mainly on tree seeds and some insects during migration and may not have access to a diet high in n-3 fatty acids. We suspect that elevated dietary levels of 18:2n-6 also caused the observed increases in the proportion of this fatty acid in muscle phospholipids. Overall, we conclude that seasonal changes in adipose and muscle fatty acid composition are likely attributable to diet more than other factors such as migratory exercise or mitochondrial density.

Dietary polyunsaturated fatty acids (PUFA) can have various effects on animal physiology through their roles as energy, structural, regulatory, and signaling molecules. Of recent interest has been the incorporation of dietary PUFA into muscle membranes as phospholipids, thereby potentially affecting exercise performance by mechanisms such as altered mitochondrial proton leak and membrane-bound protein activity. We first studied the effects of a high-omega6 PUFA diet vs. a high-omega3 PUFA diet on peak metabolic rate (PMR) in white-throated sparrows, and additionally measured mRNA expression of fatty acid transporters and the activity of major oxidative enzymes. Our experiment, thus, allowed a test of the "natural doping" hypothesis. With a simple diet manipulation, the two groups of sparrows diverged significantly in both muscle phospholipid composition and adipose triacylglycerol composition. The high-omega6 sparrows achieved higher PMR without a change in enzyme activity or transporter expression. We then fed sparrows the 2 diets, followed by a food restriction (H omega3RI and H omega6RI treatments). When their adipose stores were exhausted, we fed both groups a common diet of intermediate fatty acid composition. This protocol resulted in the H omega6RI and H omega3RI groups diverging significantly in muscle phospholipid composition, but they had substantially similar adipose stores. PMR did not differ between the H omega6RI and H omega3RI groups. We conclude that muscle phospholipids do not play a major role in affecting exercise performance. The fatty acid composition of stored triacylglycerol may instead affect exercise via the preferential use of particular fatty acids by muscles.

During times of high energy demand, stored fatty acids are mobilized from adipocytes. This mobilization has previously been shown to be a non-random process, with more hydrophilic fatty acids being mobilized most readily. The objectives of this study were to characterize the relative mobilization of fatty acids from adipocytes in two migratory bird species and to investigate possible changes in selective fatty acid mobilization associated with the migratory period. Captive ruffs (Philomachus pugnax) and white-crowned sparrows (Zonotrichia leucophrys) were studied. The sparrows were divided into two treatments: 'winter' (photoperiod 8 h:16 h L:D) and 'migrant' (in which migratory condition was induced with a photoperiodic manipulation of 8 h:16 h L:D, followed by 16 h:8 h L:D). Adipose tissue was removed from ruffs and sparrows and incubated for 90 min after stimulation with epinephrine. The proportions of individual fatty acid species released into the incubation medium were compared with their proportions in the adipocytes to determine relative mobilizations. We found that patterns of relative mobilization in ruffs and sparrows are similar to those of mammals, with shorter chain lengths and more double bonds leading to higher relative mobilization. Seasonal condition in sparrows did not alter this pattern. This pattern of relative mobilization from adipocytes seems to be a general rule amongst birds and mammals and should be considered before inferring functionality about selective retention or mobilization of certain fatty acids. The composition of adipose stores in birds may affect migratory performance; however, our results indicate that patterns of relative mobilization at the adipocytes do not vary with season in migratory birds.