Ruf Thomas, Arnold Walter
Research Institute of Wildlife Ecology, University of Veterinary Medicine, Savoyenstr.1, A-1160 Vienna, Austria.
Am J Physiol Regul Integr Comp Physiol. 2008 Mar;294(3):R1044-52. doi: 10.1152/ajpregu.00688.2007. Epub 2008 Jan 2.
Polyunsaturated fatty acids (PUFAs) can have strong effects on hibernation and daily torpor in mammals. High dietary PUFA contents were found to increase proneness for torpor, decrease body temperatures, prolong torpor bout duration, and attenuate hibernation mass loss. The mechanism by which PUFAs enhance torpor and hibernation is unknown, however. On the basis of a review of the literature, and on reexamining our own data on alpine marmots, we propose that effects on hibernation are not due to PUFAs in general, but to shifts in the ratio of n-6 PUFAs to n-3 PUFAs in membrane phospholipids. Specifically, high ratios of n-6 to n-3 PUFAs increase the activity of the Ca2+-Mg2+ pump in the sarcoplasmic reticulum of the heart (SERCA) and counteract Q10 effects on SERCA activity at low tissue temperatures. Therefore, high n-6 to n-3 PUFA ratios in cardiac myocyte membranes appear to protect the hibernating heart from arrhythmia, which in hypothermic nonhibernators is caused by massive increases in cytosolic Ca2+. The resulting reduced risk of cardiac arrest during hypothermia may explain why increased dietary uptake of n-6 PUFAs, but not of n-3 PUFAs, can strongly enhance the propensity for hibernation, and allows heterotherms to reach lower body temperatures, with associated increased energy savings. Therefore, at least for herbivorous hibernators, such as marmots, linoleic acid (C18:2 n-6)--the dietary source of all n-6 PUFAs--appears to represent a crucial and limited resource in natural environments.
多不饱和脂肪酸(PUFAs)对哺乳动物的冬眠和日常蛰伏有着显著影响。研究发现,高膳食PUFA含量会增加动物进入蛰伏状态的倾向,降低体温,延长蛰伏时间,并减少冬眠期间的体重损失。然而,PUFAs增强蛰伏和冬眠的机制尚不清楚。基于对文献的综述以及对我们自己关于高山旱獭数据的重新审视,我们提出,对冬眠的影响并非普遍归因于PUFAs,而是由于膜磷脂中n-6 PUFAs与n-3 PUFAs比例的变化。具体而言,高比例的n-6与n-3 PUFAs会增加心脏肌浆网(SERCA)中Ca2+-Mg2+泵的活性,并抵消低组织温度下Q10对SERCA活性的影响。因此,心肌细胞膜中高比例的n-6与n-3 PUFAs似乎能保护冬眠心脏免受心律失常的影响,在低温非冬眠动物中,心律失常是由胞质Ca2+大量增加引起的。低温期间心脏骤停风险的降低可能解释了为什么增加膳食中n-6 PUFAs(而非n-3 PUFAs)的摄入量会强烈增强冬眠倾向,并使异温动物能够达到更低的体温,从而节省更多能量。因此,至少对于草食性冬眠动物,如旱獭,亚油酸(C18:2 n-6)——所有n-6 PUFAs的膳食来源——在自然环境中似乎是一种关键且有限的资源。