Wipking W
Physiologische Ökologie, Zoologisches Institut der Universität Köln, Weyertal 119, D-50923, Köln, Germany.
Oecologia. 1995 May;102(2):202-210. doi: 10.1007/BF00333252.
The onset of larval diapause in the burnet moth Zygaena trifolii is clearly characterized by the larva molting into a specialized dormant morph. In a potentially bivoltine Mediterranean population (Marseille) two types of diapause can occur within 1 year: firstly, a facultative summer diapause of 3-10 weeks, and secondly, an obligate winter diapause, which can be lengthened by a period of thermal quiescence to several months in temperatures of ≤5°C. For the first time, three successive physiological periods have been experimentally distinguished within an insect dormancy (between onset of diapause and molting to the next non-diapause stage), using chilling periods of 30-180 days at 5°C, and varying conditions of photoperiod and temperature. These stages are: (1) a continuous Diapause-ending process (DEP); (2) thermal quiescence (Q); and finally, (3) a period of postdiapause development (PDD) before molting to the next larval instar. The result of transferring dormant larvae from chilling at 5°C to 20°C depended on the length of the chilling period. After chilling for 120-180 days, molting to the next instar occurred after 6-10 days, independent of daylength. This period corresponds with the duration of PDD. After shorter chilling periods (90, 60, 30 days and the control, 0 days) the period to eclosion increased exponentially, and included both the latter part of the previous diapause process and the 6-10 day period of PDD. However, photoperiod also influences the time to eclosion after chilling. Short daylength (8 h light / 16 h dark: LD 8/16) lengthened the diapause in comparison to long daylength (16 h light / 8 h dark: LD 16/8). Short daylength had a similar effect during chilling at 5°C, as measured by the longer time to eclosion after transfer. The shorter time to eclosion resulting from longer chilling periods (30-90 days) demonstrates that the state of diapause is continuously shortened at 5°C, and corresponds to the neuroendocrine controlled DEP. Presumably the DEP has already started after the onset of diapause. When chilling was continued after the end of the DEP, which ranged between 90 and 120 days, thermal quiescence (Q) followed (observed maximum 395 days). Different photoperiodic conditions during the pre-diapause inductive period modified diapause intensity (measured as the duration of diapause), in that a photoperiodic signal just below the critical photoperiod for diapause induction (LD 15/9) intensified diapause. Experiments simulating the summer diapause showed that PDD occurred in the range of 10-25°C. Higher temperatures (15 and 20°C) shortened the DEP at LD 16/8, so that at 20°C many individuals had already terminated diapause after 10-40 days and had molted after the 6-10 days of PDD. A temperature of 25°C unexpectedly lengthened the DEP to 110 days in several individuals. The ecological consequences and the adaptive significance of variation in the duration of the diapause are discussed in relation to the persistence of local populations predictably variable and rare climatic extremes throughout the year.
斑蛾Zygaena trifolii幼虫滞育的开始明显表现为幼虫蜕变为一种特殊的休眠形态。在具有潜在双季发生能力的地中海种群(马赛)中,1年内可能出现两种滞育类型:第一,为期3 - 10周的兼性夏季滞育;第二,专性冬季滞育,在≤5°C的温度下,通过一段热静止期可延长至数月。首次通过在5°C下进行30 - 180天的低温处理以及不同的光周期和温度条件,在昆虫休眠期(从滞育开始到蜕变为下一个非滞育阶段)内实验区分出三个连续的生理阶段。这些阶段分别是:(1) 一个连续的滞育结束过程(DEP);(2) 热静止(Q);最后,(3) 在蜕变为下一龄幼虫之前的滞育后发育阶段(PDD)。将处于休眠状态的幼虫从5°C低温转移到20°C后的结果取决于低温处理的时长。在低温处理120 - 180天后,6 - 10天内会蜕变为下一龄幼虫,与日长无关。这个时期与PDD的时长相对应。在较短的低温处理时长(90天、60天、30天以及对照0天)后,羽化所需时间呈指数增加,并且包括前一滞育过程的后期以及6 - 10天的PDD阶段。然而,光周期也会影响低温处理后的羽化时间。与长日长(16小时光照/8小时黑暗:LD 16/8)相比,短日长(8小时光照/16小时黑暗:LD 8/16)会延长滞育时间。在以转移后羽化所需更长时间衡量的5°C低温处理期间,短日长也有类似效果。较长低温处理时长(30 - 90天)导致的较短羽化时间表明,在5°C下滞育状态持续缩短,这与神经内分泌控制的DEP相对应。推测DEP在滞育开始后就已经启动。当在90至120天的DEP结束后继续进行低温处理时,随后会进入热静止(Q)阶段(观察到的最长时间为395天)。滞育前诱导期的不同光周期条件会改变滞育强度(以滞育持续时间衡量),即刚好低于滞育诱导临界光周期的光周期信号(LD 15/9)会增强滞育。模拟夏季滞育的实验表明,PDD发生在10 - 25°C范围内。较高温度(15°C和20°C)在LD 16/8条件下缩短了DEP,以至于在20°C时,许多个体在10 - 40天后就已经结束滞育,并在PDD的6 - 10天后蜕皮。25°C的温度意外地使几个个体的DEP延长至110天。结合全年当地种群可预测的可变且罕见的极端气候的持续情况,讨论了滞育持续时间变化的生态后果和适应性意义。