Sokolov E N, Nezlina N I
Institute of Higher Nervous Activity and Neurophysiology, M. V. Lomonosov Moscow State University, Russian Academy of Sciences, Moscow.
Neurosci Behav Physiol. 2008 Jan;38(1):1-14. doi: 10.1007/s11055-008-0001-7.
The conditioned reflex is characterized by plasticity supporting bilateral selective connections between its input and output. In simple nervous systems, input stimuli are represented by selective detectors connected to command neurons by plastic synapses whose activity increases on learning and decreases on extinction. The process of associative learning occurs when excitation of the detector and the command neuron coincide. Short-term memory in a plastic synapse is associated with phosphorylation of postsynaptic receptor molecules and does not require protein synthesis. Long-term memory is associated with early gene expression, structural genes, and protein synthesis. The simple "detector-command neuron" association has increased in complexity during evolution. At the input, pre-detector interneurons activating a specific detector converge on the command neuron: the command neuron determines the selectivity of the mechanisms of conditioned reflexes for complex stimuli. The output mechanism has also become more complex: command neurons have become more specialized and premotor interneurons have appeared between them and motor neurons, excitation of premotor neurons being passed to groups of motor neurons responsible for the configuration of the behavioral act. Conditioned reflexes combining more complex signals at the input with more flexible results at the output allow a diversity of behavioral acts.
条件反射的特点是具有可塑性,支持其输入和输出之间的双侧选择性连接。在简单的神经系统中,输入刺激由通过可塑性突触与指令神经元相连的选择性检测器来表征,这些突触的活动在学习时增加,在消退时减少。当检测器和指令神经元同时兴奋时,就会发生联合学习过程。可塑性突触中的短期记忆与突触后受体分子的磷酸化有关,不需要蛋白质合成。长期记忆与早期基因表达、结构基因和蛋白质合成有关。在进化过程中,简单的“检测器-指令神经元”关联的复杂性增加了。在输入端,激活特定检测器的前检测器中间神经元汇聚到指令神经元上:指令神经元决定了条件反射机制对复杂刺激的选择性。输出机制也变得更加复杂:指令神经元变得更加专业化,并且在它们和运动神经元之间出现了运动前中间神经元,运动前神经元的兴奋传递给负责行为动作构型的运动神经元群。在输入端组合更复杂信号、在输出端产生更灵活结果的条件反射允许产生多种多样的行为动作。