Tisdale Ryan K, Vyssotski Alexei L, Lesku John A, Rattenborg Niels C
Avian Sleep Group, Max Planck Institute for Ornithology, Seewiesen, Germany.
Brain Behav Evol. 2017;89(4):249-261. doi: 10.1159/000475590. Epub 2017 Jul 7.
The functions of slow wave sleep (SWS) and rapid eye movement (REM) sleep, distinct sleep substates present in both mammals and birds, remain unresolved. One approach to gaining insight into their function is to trace the evolution of these states through examining sleep in as many taxonomic groups as possible. The mammalian and avian clades are each composed of two extant groups, i.e., the monotremes (echidna and platypus) and therian (marsupial and eutherian [or placental]) mammals, and Palaeognaths (cassowaries, emus, kiwi, ostriches, rheas, and tinamous) and Neognaths (all other birds) among birds. Previous electrophysiological studies of monotremes and ostriches have identified a unique "mixed" sleep state combining features of SWS and REM sleep unlike the well-delineated sleep states observed in all therian mammals and Neognath birds. In the platypus this state is characterized by periods of REM sleep-related myoclonic twitching, relaxed skeletal musculature, and rapid eye movements, occurring in conjunction with SWS-related slow waves in the forebrain electroencephalogram (EEG). A similar mixed state was also observed in ostriches; although in addition to occurring during periods with EEG slow waves, reduced muscle tone and rapid eye movements also occurred in conjunction with EEG activation, a pattern typical of REM sleep in Neognath birds. Collectively, these studies suggested that REM sleep occurring exclusively as an integrated state with forebrain activation might have evolved independently in the therian and Neognath lineages. To test this hypothesis, we examined sleep in the elegant crested tinamou (Eudromia elegans), a small Palaeognath bird that more closely resembles Neognath birds in size and their ability to fly. A 24-h period was scored for sleep state based on electrophysiology and behavior. Unlike ostriches, but like all of the Neognath birds examined, all indicators of REM sleep usually occurred in conjunction with forebrain activation in tinamous. The absence of a mixed REM sleep state in tinamous calls into question the idea that this state is primitive among Palaeognath birds and therefore birds in general.
慢波睡眠(SWS)和快速眼动(REM)睡眠是哺乳动物和鸟类都存在的不同睡眠亚状态,其功能仍未明确。深入了解它们功能的一种方法是通过尽可能多地研究不同分类群的睡眠情况来追溯这些睡眠状态的演化。哺乳动物和鸟类分支各自都由两个现存类群组成,即单孔目动物(针鼹和鸭嘴兽)和兽亚纲(有袋类和真兽亚纲[或胎盘类])哺乳动物,以及鸟类中的古颚类(食火鸡、鸸鹋、几维鸟、鸵鸟、美洲鸵和鹬鸵)和新颚类(所有其他鸟类)。先前对单孔目动物和鸵鸟的电生理研究发现了一种独特的“混合”睡眠状态,它结合了慢波睡眠和快速眼动睡眠的特征,这与在所有兽亚纲哺乳动物和新颚类鸟类中观察到的界限分明的睡眠状态不同。在鸭嘴兽中,这种状态的特征是与快速眼动睡眠相关的肌阵挛抽搐、松弛的骨骼肌以及快速眼动,同时在前脑脑电图(EEG)中出现与慢波睡眠相关的慢波。在鸵鸟中也观察到了类似的混合状态;尽管除了在脑电图出现慢波的时期出现外,肌张力降低和快速眼动还与脑电图激活同时出现,这是新颚类鸟类快速眼动睡眠的典型模式。总体而言,这些研究表明,完全作为一种与前脑激活相关的整合状态出现的快速眼动睡眠可能在兽亚纲和新颚类谱系中独立演化。为了验证这一假设,我们研究了凤头鹬鸵(Eudromia elegans)的睡眠情况,凤头鹬鸵是一种小型古颚类鸟类,在体型和飞行能力方面更类似于新颚类鸟类。根据电生理和行为对24小时内的睡眠状态进行评分。与鸵鸟不同,但与所有已研究的新颚类鸟类一样,凤头鹬鸵中快速眼动睡眠的所有指标通常都与前脑激活同时出现。凤头鹬鸵不存在混合快速眼动睡眠状态,这对这种状态在古颚类鸟类乃至整个鸟类中是原始状态的观点提出了质疑。