Naylor E
Symp Soc Exp Biol. 1985;39:63-93.
The best general hypothesis for the control of 'spontaneous' tidal and daily patterns of behaviour in coastal animals postulates an endogenous physiological pacemaker system which generates approximate periodicity, together with environmental adjustment of the clock(s) to local time. Free-running endogenous rhythms of circatidal, circadian, circasemilunar and circalunar periodicity have been demonstrated in a number of species in constant laboratory conditions, in some cases clarifying hitherto poorly understood aspects of the behavioural repertoire of animals in the sea. Entrainment of circatidal rhythmicity has been demonstrated using cycles of simulated tidal variables such as temperature, hydrostatic pressure, salinity and wave action. The crab Carcinus shows increased locomotor activity after changes of salinity (halokinesis); responses to 34% salinity entrain the endogenous clock, but responses to salinities above or below 34% are purely exogenous and do not persist in constant conditions after entrainment. Phase responsiveness of circatidal rhythms to pulses of tidal variables has been demonstrated in several species; phase response curves show marked differences from those of circadian rhythms. The endogenous basis of tidal and diel behaviour in marine molluscs and crustaceans involves matching spontaneous rhythms of neuroelectrical activity. Also, in decapod crustaceans a peptidic neurodepressing hormone (NDH) modulates neuroelectrical and behavioural rhythmicity. NDH is produced rhythmically in the eyestalk neurosecretory complex, perhaps partly under the control of other clock components elsewhere in the CNS. The physiological basis of circasemilunar, lunar (and annual) rhythms of behaviour has not been studied, but studies of synchronization of these rhythms have been undertaken. In some localities it has been shown experimentally that light intensities equivalent to moonlight are sufficient to entrain such rhythms. In other localities where moonlight is a less reliable cue the relative timing of tidal and daily variables has been shown to be important. So far there is no evidence that synchronization is achieved by absolute differences between tidal variables at neap and spring tides.
关于控制沿海动物“自发”潮汐和日常行为模式的最佳总体假说是，假定存在一个内源性生理起搏器系统，它能产生近似周期性，同时生物钟会根据当地时间进行环境调节。在恒定的实验室条件下，已在许多物种中证明了具有潮汐、昼夜、半月和月周期的自由运行内源性节律，在某些情况下，这也阐明了海洋动物行为模式中迄今了解甚少的方面。已通过模拟潮汐变量（如温度、静水压力、盐度和波浪作用）的周期来证明潮汐节律的夹带。螃蟹Carcinus在盐度变化后（盐度运动）表现出运动活动增加；对34%盐度的反应会夹带内源性生物钟，但对高于或低于34%盐度的反应纯粹是外源性的，夹带后在恒定条件下不会持续。已在几个物种中证明了潮汐节律对潮汐变量脉冲的相位响应性；相位响应曲线与昼夜节律的相位响应曲线有明显差异。海洋软体动物和甲壳类动物的潮汐和昼夜行为的内源性基础涉及神经电活动自发节律的匹配。此外，在十足目甲壳类动物中，一种肽类神经抑制激素（NDH）调节神经电和行为节律。NDH在眼柄神经分泌复合体中有节律地产生，可能部分受中枢神经系统其他生物钟成分的控制。行为的半月、月（和年）节律的生理基础尚未研究，但已对这些节律的同步进行了研究。在一些地方，实验表明相当于月光的光强度足以夹带此类节律。在月光不太可靠的其他地方，潮汐和日常变量的相对时间已被证明很重要。到目前为止，没有证据表明通过小潮和大潮时潮汐变量的绝对差异来实现同步。
