Matthews Philip G D
Department of Zoology, The University of British Columbia, Vancouver, B.C., Canada.
J Comp Physiol B. 2018 Mar;188(2):195-210. doi: 10.1007/s00360-017-1121-6. Epub 2017 Aug 17.
This review examines the control of gas exchange in insects, specifically examining what mechanisms could explain the emergence of discontinuous gas exchange cycles (DGCs). DGCs are gas exchange patterns consisting of alternating breath-hold periods and bouts of gas exchange. While all insects are capable of displaying a continuous pattern of gas exchange, this episodic pattern is known to occur within only some groups of insects and then only sporadically or during certain phases of their life cycle. Investigations into DGCs have tended to emphasise the role of chemosensory thresholds in triggering spiracle opening as critical for producing these gas exchange patterns. However, a chemosensory basis for episodic breathing also requires an as-of-yet unidentified hysteresis between internal respiratory stimuli, chemoreceptors, and the spiracles. What has been less appreciated is the role that the insect's central nervous system (CNS) might play in generating episodic patterns of ventilation. The active ventilation displayed by many insects during DGCs suggests that this pattern could be the product of directed control by the CNS rather than arising passively as a result of self-sustaining oscillations in internal oxygen and carbon dioxide levels. This paper attempts to summarise what is currently known about insect gas exchange regulation, examining the location and control of ventilatory pattern generators in the CNS, the influence of chemoreceptor feedback in the form of O and CO/pH fluctuations in the haemolymph, and the role of state-dependent changes in CNS activity on ventilatory control. This information is placed in the context of what is currently known regarding the production of discontinuous gas exchange patterns.
本综述探讨了昆虫气体交换的调控,特别研究了哪些机制可以解释间断性气体交换循环(DGCs)的出现。DGCs是一种气体交换模式,由交替的屏息期和气体交换期组成。虽然所有昆虫都能够表现出连续的气体交换模式,但这种间歇性模式仅在某些昆虫群体中出现,而且只是偶尔出现或在其生命周期的特定阶段出现。对DGCs的研究往往强调化学感应阈值在触发气门打开方面的作用,认为这对产生这些气体交换模式至关重要。然而,间歇性呼吸的化学感应基础还需要内部呼吸刺激、化学感受器和气门之间一种尚未确定的滞后现象。而昆虫中枢神经系统(CNS)在产生间歇性通气模式中可能发挥的作用却较少受到关注。许多昆虫在DGCs期间表现出的主动通气表明,这种模式可能是CNS直接控制的产物,而不是由于内部氧气和二氧化碳水平的自我维持振荡而被动产生的。本文试图总结目前关于昆虫气体交换调节的已知信息,研究CNS中通气模式发生器的位置和控制、血淋巴中O₂和CO₂/pH波动形式的化学感受器反馈的影响,以及CNS活动中与状态相关的变化对通气控制的作用。这些信息将放在目前关于间断性气体交换模式产生的已知情况的背景下进行探讨。