Ostriker G, Pellionisz A, Llinás R
Neuroscience. 1985 Feb;14(2):483-500. doi: 10.1016/0306-4522(85)90304-5.
The central nervous system expresses its function in natural frames of reference. A most conspicuous feature of such frames is their non-orthogonality. Gaze stabilization and, in particular, the sensorimotor transformations performed by the vestibulo-ocular reflex, are prime examples of such general coordinate transformations between and within multidimensional non-orthogonal frames. Since such operations can be described by tensor formalisms in an abstract manner, this methodology is applied here to develop a tensorial computer model of gaze stabilization. The representation of sensorimotor transformations by a reference-frame independent method obviates the necessity to simplify the intrinsic coordinate systems either by a reduction of the dimensionality or by a presumption of orthogonality. The frames of reference intrinsic to vestibulo-ocular reflex transformation (the vestibular semicircular canals and extraocular muscles) as well as the covariant character of the sensory input and the contravariant character of the motor output are physically obvious. A model built on these intrinsic systems of coordinates first serves to quantitate the degree of non-orthogonality in the extraocular muscle system, and thus to demonstrate both the necessity and the applicability of representing them by a formalism suitable for non-orthogonal systems, such as tensor network theory. The actual non-orthogonality of the gaze-stabilization system can be quantitated on the basis of the difference of covariant and contravariant expressions as follows. Tensor network theory describes sensorimotor transformations by employing a covariant embedding procedure. This, however, yields a covariant intention-type motor vector. If the central nervous system were to transmit these sensory-type components directly to the extraocular muscle motor mechanism, an error-angle would occur since covariants do not physically compose the intended movement. The error in every direction of gaze would be zero only if the extraocular muscle system would constitute an orthogonal set of rotation axes. Otherwise, the error, called refraction angle, is a measure of non-orthogonality. The complexity of the quantitation of non-orthogonality is compounded by the fact that these rotation axes change with the moving eye. Calculation of eye movements, executed both by covariant and contravariant vectors from primary and secondary eye positions, is based on the simplest assumption that the central nervous system establishes the covariant-contravariant transformation in the retinal tangent plane.(ABSTRACT TRUNCATED AT 400 WORDS)
中枢神经系统在自然参照系中发挥其功能。此类参照系最显著的特征是它们的非正交性。凝视稳定,尤其是前庭眼反射所执行的感觉运动转换,是多维非正交参照系之间以及内部这种一般坐标转换的主要示例。由于此类操作可以用张量形式主义以抽象方式描述,因此这里应用这种方法来开发一个凝视稳定的张量计算机模型。通过一种与参照系无关的方法来表示感觉运动转换,避免了通过降低维度或假定正交性来简化固有坐标系的必要性。前庭眼反射转换所固有的参照系(前庭半规管和眼外肌)以及感觉输入的协变特性和运动输出的逆变特性在物理上是明显的。基于这些固有坐标系构建的模型首先用于量化眼外肌系统中的非正交程度,从而证明用适合非正交系统的形式主义(如张量网络理论)来表示它们的必要性和适用性。凝视稳定系统实际的非正交性可以根据协变和逆变表达式的差异按如下方式进行量化。张量网络理论通过采用协变嵌入过程来描述感觉运动转换。然而,这会产生一个协变意图型运动向量。如果中枢神经系统将这些感觉型分量直接传递到眼外肌运动机制,由于协变量在物理上并不构成预期运动,就会出现误差角。只有当眼外肌系统构成一组正交的旋转轴时,每个凝视方向上的误差才会为零。否则,称为折射角的误差就是非正交性的一种度量。非正交性量化的复杂性因这些旋转轴随眼球运动而变化这一事实而加剧。根据初级和次级眼位的协变和逆变向量来计算眼球运动,是基于中枢神经系统在视网膜切平面中建立协变 - 逆变转换这一最简单的假设。(摘要截选至400字)