A THEORY OF CEREBELLAR CORTEX
A THEORY OF CEREBELLAR CORTEX
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DOI：
10.1113/jphysiol.1969.sp008820
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发表时间：
1969-01-01
影响因子：
5.5
通讯作者：
MARR, D
中科院分区：
文献类型：
--
作者：
MARR, D
1.提出了小脑皮质的详细理论，其结果是小脑学习执行运动技能。描述了两种形式的输入-输出关系，都与皮质理论一致。一种适合学习动作（动作），另一种适合学习保持姿势和平衡（维持反射）。 2.已知下橄榄细胞和小脑浦肯野细胞在攀爬纤维输入诱导下具有特殊的一对一关系。对于学习动作，假设：（a）每个橄榄细胞对基本运动的大脑指令做出反应。任何动作在基本运动方面都有一个定义性的表征，并且这种表征具有神经表达，即下橄榄核中的一系列发射模式； (b) 在神经系统的正确状态下，浦肯野细胞可以启动其相应的橄榄细胞做出反应的基本运动。3.每当橄榄细胞放电时，它就会向其相应的浦肯野细胞发送脉冲（通过攀爬纤维输入）。该浦肯野细胞还（通过苔藓纤维输入）暴露于有关其橄榄细胞发射的环境的信息；研究还表明，在排练某个动作时，每个浦肯野细胞如何学会识别此类背景。之后，当学习了该动作时，仅上下文的发生就足以激发浦肯野细胞，从而引起下一个基本运动。因此，动作的进展就像排练时一样。4.结果表明，小脑皮层作为一种允许每个浦肯野细胞学习许多背景的结构的解释与各种类型细胞的分布以及它们已知的兴奋或抑制性质是一致的。结果表明，苔藓纤维颗粒细胞排列提供了所需的模式辨别能力。5.做出以下预测：(a)突触前和攀爬纤维（或突触后）活动的结合促进了从平行纤维到浦肯野细胞的突触。经英国牛津生理学会许可转载。(b)没有其他小脑突触是可修改的。(c)高尔基细胞由其上树突区和下树突区的较大输入驱动。6.为了学习维持反射，2(a)和2(b)被2’取代。每个橄榄细胞都受到一种或多种受体的刺激，这些受体的所有活性通常都会因刺激相应浦肯野细胞的结果而降低。 7.结果表明，如果满足(2’)，则受体→橄榄细胞→浦肯野细胞→效应器的回路可被视为由习得的苔藓纤维输入激活的稳定反射回路。这种类型的反射被称为习得的条件反射，它表明了如何……
1.A detailed theory of cerebellar cortex is proposed whose consequence is that the cerebellum learns to perform motor skills. Two forms of input-output relation are described, both consistent with the cortical theory. One is suitable for learning movements (actions), and the other for learning to maintain posture and balance (maintenance reflexes).2.It is known that the cells of the inferior olive and the cerebellar Purkinje cells have a special one-to-one relationship induced by the climbing fibre input. For learning actions, it is assumed that:(a)each olivary cell responds to a cerebral instruction for an elemental movement. Any action has a defining representation in terms of elemental movements, and this representation has a neural expression as a sequence of firing patterns in the inferior olive; and(b)in the correct state of the nervous system, a Purkinje cell can initiate the elemental movement to which its corresponding olivary cell responds.3.Whenever an olivary cell fires, it sends an impulse (via the climbing fibre input) to its corresponding Purkinje cell. This Purkinje cell is also exposed (via the mossy fibre input) to information about the context in which its olivary cell fired; and it is shown how, during rehearsal of an action, each Purkinje cell can learn to recognize such contexts. Later, when the action has been learnt, occurrence of the context alone is enough to fire the Purkinje cell, which then causes the next elemental movement. The action thus progresses as it did during rehearsal.4.It is shown that an interpretation of cerebellar cortex as a structure which allows each Purkinje cell to learn a number of contexts is consistent both with the distributions of the various types of cell, and with their known excitatory or inhibitory natures. It is demonstrated that the mossy fibre-granule cell arrangement provides the required pattern discrimination capability.5.The following predictions are made.(a)The synapses from parallel fibres to Purkinje cells are facilitated by the conjunction of presynaptic and climbing fibre (or post-synaptic) activity. Reprinted with permission of The Physiological Society, Oxford, England.(b)No other cerebellar synapses are modifiable.(c)Golgi cells are driven by the greater of the inputs from their upper and lower dendritic fields.6.For learning maintenance reflexes, 2(a) and 2 (b) are replaced by 2’. Each olivary cell is stimulated by one or more receptors, all of whose activities are usually reduced by the results of stimulating the corresponding Purkinje cell.7.It is shown that if (2’) is satisfied, the circuit receptor → olivary cell → Purkinje cell → effector may be regarded as a stabilizing reflex circuit which is activated by learned mossy fibre inputs. This type of reflex has been called a learned conditional reflex, and it is shown how such …