Pernet Fabrice, Tremblay Réjean, Gionet Chantal, Landry Thomas
Institut de Recherche sur les Zones Côtières, 232B rue de l'Eglise, Shippagan, Nouveau-Brunswick, E8S 1J2, Canada.
J Exp Biol. 2006 Dec;209(Pt 23):4663-75. doi: 10.1242/jeb.02581.
A temperature decrease usually induces an ordering effect in membrane phospholipids, which can lead to membrane dysfunction. Poikilotherms inhabiting eurythermal environments typically counteract this temperature effect by remodeling membrane lipids as stipulated in the homeoviscous adaptation theory (HVA). Hard clams, Mercenaria mercenaria, can suffer high overwintering mortalities in the Gulf of St Lawrence, Canada. The selectively bred M. mercenaria var. notata can have higher overwintering mortalities than the wild species, thus suggesting that the two varieties have different degrees of adaptation to low temperatures. The objective of this study was to investigate the changes in lipid composition of soft tissues in wild and selected hard clams in relation to their metabolic and genetic characteristics. Clams were placed at the northern limit of their distribution from August 2003 to May 2004; they were exposed to a gradual temperature decrease and then maintained at <0 degrees C for 3.5 months. This study is the first to report a major remodeling of lipids in this species as predicted by HVA; this remodeling involved a sequential response of the phospholipid to sterol ratio as well as in levels of 22:6n-3 and non-methylene interrupted dienoic fatty acids. Hard clams showed an increase in 20:5n-3 as temperature decreased, but this was not maintained during overwintering, which suggests that 20:5n-3 may have been used for eicosanoid biosynthesis as a stress response to environmental conditions. Selectively bred hard clams were characterized by a higher metabolic demand and a deviation from Hardy-Weinberg equilibrium at several genetic loci due to a deficit in heterozygote frequency compared with wild clams, which is believed to impose additional stress and render these animals more vulnerable to overwintering mortality. Finally, an intriguing finding is that the lower metabolic requirements of wild animals coincide with a lower unsaturation index of their lipids, as predicted by Hulbert's theory of membranes as pacemakers of metabolism.
温度降低通常会在膜磷脂中引发有序化效应,这可能导致膜功能障碍。栖息在广温环境中的变温动物通常会按照适应性体温调节理论(HVA)的规定,通过重塑膜脂来抵消这种温度效应。硬壳蛤,即商业蛤(Mercenaria mercenaria),在加拿大圣劳伦斯湾可能会遭受较高的越冬死亡率。经过选择性培育的商业蛤变种(M. mercenaria var. notata)的越冬死亡率可能高于野生品种,这表明这两个变种对低温的适应程度不同。本研究的目的是调查野生和经过选择的硬壳蛤软组织中脂质组成的变化及其代谢和遗传特征。2003年8月至2004年5月,将蛤放置在其分布的北界;使其经历逐渐降温,然后在<0摄氏度下维持3.5个月。本研究首次报告了该物种脂质的重大重塑,正如HVA所预测的那样;这种重塑涉及磷脂与固醇比例以及22:6n-3和非亚甲基间断二烯酸脂肪酸水平的顺序反应。随着温度降低,硬壳蛤的20:5n-3增加,但在越冬期间并未维持,这表明20:5n-3可能已用于类花生酸生物合成,作为对环境条件的应激反应。经过选择性培育的硬壳蛤的特征是代谢需求较高,并且在几个基因位点偏离哈迪-温伯格平衡,这是由于与野生蛤相比杂合子频率不足,据信这会带来额外压力并使这些动物更容易遭受越冬死亡。最后,一个有趣的发现是,正如Hulbert的膜作为代谢起搏器的理论所预测的那样,野生动物较低的代谢需求与其脂质较低的不饱和指数相吻合。