Department of Biology, Advanced Facility for Avian Research, University of Western Ontario, 1151 Richmond Street North, London, Ontario N6A5B7, Canada.
Integr Comp Biol. 2010 Sep;50(3):336-45. doi: 10.1093/icb/icq097. Epub 2010 Jul 16.
The metaphor of marathon running is inadequate to fully capture the magnitude of long-distance migratory flight of birds. In some respects a journey to the moon seems more appropriate. Birds have no access to supplementary water or nutrition during a multi-day flight, and they must carefully budget their body fat and protein stores to provide both fuel and life support. Fatty acid transport is crucial to successful non-stop migratory flight in birds. Although fat is the most energy-dense metabolic fuel, the insolubility of its component fatty acids makes them difficult to transport to working muscles fast enough to support the highly aerobic exercise required to fly. Recent evidence indicates that migratory birds compensate for this by expressing large amounts of fatty acid transport proteins on the membranes of the muscles (FAT/CD36 and FABPpm) and in the cytosol (H-FABP). Through endogenous mechanisms and/or diet, migratory birds may alter the fatty acid composition of the fat stores and muscle membranes to improve endurance during flight. Fatty acid chain length, degree of unsaturation, and placement of double bonds can affect the rate of mobilization of fatty acids from adipose tissue, utilization of fatty acids by muscles, and whole-animal performance. However, there is great uncertainty about how important fatty acid composition is to the success of migration or whether particular types of fatty acids (e.g., omega-3 or omega-6) are most beneficial. Migratory bats provide an interesting example of evolutionary convergence with birds, which may provide evidence for the generality of the bird model to the evolution of migration by flight in vertebrates. Yet only recently have attempts been made to study bat migration physiology. Many aspects of their fuel metabolism are predicted to be more similar to those of migrant birds than to those of non-flying mammals. Bats may be distinct from most birds in their potential to conserve energy by using torpor between flights, and in the behavioral and physiological trade-offs they may make between migration and reproduction, which often overlap.
鸟类长途迁徙的飞行,其规模之大,用马拉松长跑来比喻都还远远不够。从某些方面来看,飞向月球似乎更为贴切。鸟类在多日的飞行中无法补充水分或营养,它们必须精心规划体脂和蛋白质的储存,以提供燃料和生命支持。脂肪酸的转运对于鸟类成功的不间断迁徙飞行至关重要。尽管脂肪是能量密度最高的代谢燃料,但由于其组成脂肪酸的不溶性,使得它们难以快速转运到工作肌肉中,无法为需要高度有氧的飞行提供足够的燃料。最近的证据表明,候鸟通过在肌肉(FAT/CD36 和 FABPpm)和细胞质(H-FABP)的膜上大量表达脂肪酸转运蛋白来弥补这一点。通过内源性机制和/或饮食,候鸟可能会改变脂肪储存和肌肉膜中的脂肪酸组成,以提高飞行中的耐力。脂肪酸链长、不饱和程度和双键位置都会影响脂肪组织中脂肪酸的动员速度、肌肉对脂肪酸的利用以及整个动物的表现。然而,脂肪酸组成对于迁徙成功的重要性以及特定类型的脂肪酸(例如 omega-3 或 omega-6)是否最有益仍存在很大的不确定性。迁徙蝙蝠为与鸟类趋同进化提供了一个有趣的例子,这可能为鸟类模型在脊椎动物飞行迁徙进化中的普遍性提供了证据。然而,直到最近才有人试图研究蝙蝠的迁徙生理学。它们的许多燃料代谢方面预计与候鸟更为相似,而与非飞行哺乳动物的相似性则较低。蝙蝠在飞行之间利用蛰伏来保存能量的能力可能与大多数鸟类不同,它们在迁徙和繁殖之间可能会做出行为和生理上的权衡,而这两者往往会重叠。