Shunkina Ksenia V, Zaytseva Olga V, Starunov Viktor V, Ostrovsky Andrew N
Laboratory of Evolutionary Morphology, Zoological Institute, Russian Academy of Sciences, Universitetskaja nab. 1, 199034 Saint Petersburg, Russia.
Laboratory of Evolutionary Morphology, Zoological Institute, Russian Academy of Sciences, Universitetskaja nab. 1, 199034 Saint Petersburg, Russia ; Department of Invertebrate Zoology, Faculty of Biology, Saint Petersburg State University, Universitetskaja nab. 7/9, 199034 Saint Petersburg, Russia.
Front Zool. 2015 Oct 12;12:28. doi: 10.1186/s12983-015-0112-2. eCollection 2015.
Though some elements of the bryozoan nervous system were discovered 180 years ago, few studies of their neuromorphology have been undertaken since that time. As a result the general picture of the bryozoan nervous system structure is incomplete in respect of details and fragmentary in respect of taxonomic coverage.
The nervous system of three common European freshwater bryozoans - Cristatella mucedo, Plumatella repens (both with a horseshoe-shaped lophophore) and Fredericella sultana (with a circular lophophore) had numerous differences in the details of the structure but the general neuroarchitecture is similar. The nervous system of the zooid consists of the cerebral ganglion, a circumpharyngeal ring and lophophoral nerve tracts (horns), both sending numerous nerves to the tentacles, and the nerve plexuses of the body wall and of the gut. A number of the important details (distal branching of the additional radial nerve, pattern of distribution of nerve cells and neurites in the ganglion, etc.) were described for the first time. The number and position of the tentacle nerves in Cristatella mucedo was ascertained and suggestions about their function were made. The revealed distribution of various neuromediators in the nervous system allowed us to suggest functional affinities of some major nerves.
Despite the basic similarity, both the ganglion and the lophophore nervous system in Phylactolaemata have a more complex structure than in marine bryozoans (classes Gymnolaemata and Stenolaemata). First of all, their neuronal network has a denser and more complex branching pattern: most phylactolaemates have two large nerve tracts associated with lophophore arms, they have more nerves in the tentacles, additional and basal branches emitting from the main radial nerves, etc. This, in part, can be explained by the horseshoe shape of the lophophore and a larger size of the polypide in freshwater species. The structure of the nervous system in Fredericella sultana suggests that it underwent a secondary simplification following the reduction of the lophophore arms. Colony locomotion in Cristatella mucedo is based on co-ordinated activity of two perpendicular muscle layers of the sole and the plexus of motor neurons sandwiched between them. The trigger of this activity and the co-ordination mechanism remain enigmatic.
尽管苔藓虫神经系统的一些元素在180年前就已被发现,但自那时以来,对其神经形态学的研究却很少。因此,苔藓虫神经系统结构的总体情况在细节方面并不完整,在分类学覆盖范围方面也很零碎。
三种常见的欧洲淡水苔藓虫——穆氏冠苔藓虫、羽苔虫(两者都有马蹄形触手冠)和苏氏弗雷德里克苔藓虫(有圆形触手冠)的神经系统在结构细节上有许多差异,但总体神经结构相似。个虫的神经系统由脑神经节、围咽环和触手冠神经束(角)组成,两者都向触手发送许多神经,还有体壁和肠道的神经丛。首次描述了一些重要细节(额外放射状神经的远端分支、神经节中神经细胞和神经突的分布模式等)。确定了穆氏冠苔藓虫触手神经的数量和位置,并对其功能提出了建议。在神经系统中发现的各种神经介质的分布使我们能够提出一些主要神经的功能关联。
尽管基本相似,但苔藓虫纲的神经节和触手冠神经系统的结构比海洋苔藓虫(裸唇纲和窄唇纲)更复杂。首先,它们的神经元网络具有更密集和更复杂的分支模式:大多数苔藓虫纲动物有两条与触手冠臂相关的大神经束,它们的触手有更多神经,主要放射状神经发出额外的和基部的分支等。这在一定程度上可以通过触手冠的马蹄形和淡水物种中多虫体较大的尺寸来解释。苏氏弗雷德里克苔藓虫的神经系统结构表明,随着触手冠臂的减少,它经历了二次简化。穆氏冠苔藓虫的群体运动基于足底两个垂直肌肉层和夹在它们之间的运动神经元丛的协调活动。这种活动的触发因素和协调机制仍然是个谜。