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[十足目小龙虾的化学感受系统]

[Systems of chemoperception in decapod crayfish].

作者信息

Fedotov V P

出版信息

Zh Evol Biokhim Fiziol. 2009 Jan-Feb;45(1):3-24.

Abstract

The review presents data on some peripheral and central structures in the system of perception of chemical stimuli in crayfish and other Decapoda. The hair receptors on chelipeds, antennas, antennules are innervated by mechano-and chemoreceptor neurons. Antennules are the specialized crayfish chemoreceptor organs, on the surface of which groups of exteroceptors are located. There is an ordered disposition of exteroceptor receptive fields in the form of receptor hair bushes on claws of ambulatory feet (AF), antennas, antennules, and other movable appendages. Behavioral experiments have shown sensitivity of crayfish to odor of individuals of their gender, sex partners as well as the presence in crayfish of pheromones providing connection of female with offspring at the initial stages of the life cycle. Occasional chemosensory cells innervating hair bushes on the crayfish AF respond to amino acids, amines, nucleotides, and sugars. Minimal thresholds of reaction of the studied Decapoda chemoreceptors in response to some chemical compounds correspond to concentrations of 0.1-1 microM. For some chemoreceptors, dose-dependent effects have been shown. Alongside with monomodal chemoreceptors, the crayfish have bimodal receptor perceiving mechanical and chemical stimuli. The efficient response of crayfish chemoreceptors can be obtained to the substance that includes amino group with the hydrogen bridge with carboxyl group, contains no more than 3 carbon atoms in the chain, and is characterized by a certain stereoform. Among chemoreceptors there are fast and slow adapting cells. Efficiency of response of individual chemoreceptors depends on temperature of medium. Chemoreceptors reacting to ecdysterons have been revealed in crayfish. Ecdysterons play a great role in intra- and interspecies communications in Crustacea. Based on the study of efferent responses of interneurons of the first and higher orders in the first thoracic crayfish ganglion to stimulation of the own receptive fields, a concept is put forward of the structural-functional organization of afferent projections at the segmental level. Peculiarities of afferent projections from antennule chemoreceptors are considered. The data are presented on connections of these chemoreceptors with antennular, olfactory, and additional lobules, various cell groups, interneurons of the first, second, and third orders located in various brain parts. An attention is drawn to connections of serotonin neurons in glomeruli with endings of chemoreceptor neurons and projection of interneurons of the higher orders,. which are located in the internal medulla of the crayfish eye stalks. Several principles of integration of the chemoreceptor information in central parts of the crayfish nervous system are discussed. The giant serotonin neurons revealed in crayfish glomeruli most likely participate in formation of memory to certain chemical actions. Polymodal receptor signals in the central chain of the perception system activate autonomic centers, and the changes of the animal functional state can be evaluated from the heart responses. The crayfish heart responses recorded by novel noninvasive methods allow detection of the initial and other phases of the stress state at changes of the chemical quality of medium. Progress of the current biochemical and electrophysiological methods of study of chemoreceptors allow hoping for learning of fine chemoperception mechanisms in invertebrate and vertebrate animals.

摘要

这篇综述介绍了小龙虾和其他十足目动物化学刺激感知系统中一些外周和中枢结构的数据。螯足、触角、小触角上的毛感受器由机械感受器和化学感受器神经元支配。小触角是小龙虾特有的化学感受器器官,其表面分布着几组外感受器。在步行足(AF)的爪、触角、小触角和其他可移动附肢上,外感受器感受野以感受器毛丛的形式有序排列。行为实验表明,小龙虾对同种性别个体、性伴侣的气味以及在生命周期初始阶段使雌性与后代建立联系的信息素的存在具有敏感性。支配小龙虾AF上毛丛的偶尔化学感觉细胞对氨基酸、胺、核苷酸和糖有反应。所研究的十足目动物化学感受器对某些化合物反应的最小阈值对应于0.1 - 1微摩尔的浓度。对于一些化学感受器,已显示出剂量依赖性效应。除了单峰化学感受器外,小龙虾还有感知机械和化学刺激的双峰感受器。小龙虾化学感受器对含有与羧基形成氢键的氨基、链中碳原子不超过3个且具有特定立体构型的物质能产生有效反应。化学感受器中有快速适应和缓慢适应的细胞。单个化学感受器的反应效率取决于介质温度。在小龙虾中已发现对蜕皮甾酮有反应的化学感受器。蜕皮甾酮在甲壳纲动物的种内和种间通讯中起重要作用。基于对小龙虾第一胸神经节中一阶和高阶中间神经元对自身感受野刺激的传出反应的研究,提出了节段水平传入投射的结构 - 功能组织概念。考虑了来自小触角化学感受器的传入投射的特点。给出了这些化学感受器与小触角、嗅觉和附加小叶、位于不同脑区的各种细胞群、一阶、二阶和三阶中间神经元之间连接的数据。还关注了肾小球中5 - 羟色胺神经元与化学感受器神经元末梢的连接以及高阶中间神经元的投射,这些高阶中间神经元位于小龙虾眼柄的内髓质中。讨论了小龙虾神经系统中枢部分化学感受器信息整合的几个原则。在小龙虾肾小球中发现的巨大5 - 羟色胺神经元很可能参与对特定化学作用的记忆形成。感知系统中枢链中的多峰感受器信号激活自主神经中枢,动物功能状态的变化可从心脏反应中评估。通过新型非侵入性方法记录的小龙虾心脏反应能够在介质化学性质改变时检测到应激状态的初始阶段和其他阶段。当前研究化学感受器的生化和电生理方法的进展使得有望了解无脊椎动物和脊椎动物的精细化学感知机制。

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