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形状的神经编码:皮肤机械感受器对划过猴子指腹的波浪形表面的反应。

Neural encoding of shape: responses of cutaneous mechanoreceptors to a wavy surface stroked across the monkey fingerpad.

作者信息

LaMotte R H, Srinivasan M A

机构信息

Department of Anesthesiology, Yale University School of Medicine, New Haven, Connecticut 06510, USA.

出版信息

J Neurophysiol. 1996 Dec;76(6):3787-97. doi: 10.1152/jn.1996.76.6.3787.

DOI:10.1152/jn.1996.76.6.3787
PMID:8985876
Abstract
  1. The role of cutaneous mechanoreceptors in the tactile perception of shape was investigated. Objects whose surfaces were shaped as a pattern of smooth, alternating convex and concave cylindrical surfaces of differing radii of curvature were constructed such that there were no discontinuities in the slope of the surface. These "wavy surfaces" were stroked across the fingerpad of the anesthetized monkey and electrophysiological responses of slowly adapting type I mechanoreceptive afferents (SAs) and rapidly adapting type I mechanoreceptive afferents (RAs) were recorded. 2. For both SAs and RAs, each convexity indenting the skin evoked a burst of impulses and each concavity of the same curvature that followed elicited a pause in response. "Spatial event plots" (SEPs) of the occurrence of action potentials as a function of the location of the object on the receptive field were obtained and interpreted as the responses of a spatially distributed population of fibers. With increasing magnitude of curvature (equivalently, decreasing radius of curvature) of convexity, the mean width of the burst in the SEPs for each fiber type (representing the width of a region of skin containing active fibers) decreased and the mean discharge rate during the burst increased. Over a range of velocities of stroking from 1 to 40 mm/s, the number of RAs activated increased with velocity, whereas SAs were active at all velocities. For both SAs and RAs, the burst rates increased with velocity, whereas the widths of the bursts and pauses remained approximately invariant. Thus the spatial measures of burst or pause width provide a robust representation of the size of a feature on the object surface. 3. For a given velocity of stroking, the spatially distributed pattern of averaged discharge rates (spatial rate profile, SRP) provided a representation of the shape of the wavy surface. The distance between neighboring peaks in the SRP for individual RAs and SAs was approximately the same as the distance between the peaks of the wavy surface. The averaged SRP for a population of SAs provided a better representation of shape than that for RAs. Whereas active regions in the SEP can be isomorphic to the two dimensional form of the stimulus "footprint" in contact with the skin surface, the SRP, which in addition encodes the features of the stimulus in the third dimension normal to the skin surface, is not isomorphic to the stimulus shape. 4. When the sizes as well as the shapes of objects are varied, it is hypothesized that a central processing mechanism extracts the invariant property of shape from the slopes of the rising and falling phases of an SRP that has been normalized for overall differences in discharge rates. These differences would be expected to occur with variations in the parameters of stimulation such as compressional force, stroke trajectory, and stroke velocity. It was shown that a common feature of the mean SRP for SAs evoked by each wavy surface convexity, regardless of its radius, was the constancy of the slope from the base to the peak and from the peak to the base. Thus a possible code for the constant curvature of a cylinder is the constancy of the slopes along the rising and declining phases of the triangular-shaped spatial response profile evoked in the SA population by the cylindrical convexity.
摘要
  1. 研究了皮肤机械感受器在形状触觉感知中的作用。构建了表面形状为具有不同曲率半径的光滑、交替的凸圆柱面和凹圆柱面图案的物体,使表面斜率没有间断。将这些“波浪形表面”划过麻醉猴子的指腹,并记录慢适应I型机械感受传入纤维(SA)和快适应I型机械感受传入纤维(RA)的电生理反应。2. 对于SA和RA,每个使皮肤凹陷的凸面都会引发一阵冲动,随后相同曲率的每个凹面会引发反应暂停。获得了动作电位发生的“空间事件图”(SEP),其作为物体在感受野上位置的函数,并被解释为空间分布的纤维群体的反应。随着凸面曲率大小增加(等效于曲率半径减小),每种纤维类型的SEP中爆发的平均宽度(代表包含活动纤维的皮肤区域的宽度)减小,爆发期间的平均放电率增加。在1至40毫米/秒的一系列抚摸速度范围内,被激活的RA数量随速度增加,而SA在所有速度下均有活动。对于SA和RA,爆发率均随速度增加,而爆发和暂停的宽度大致保持不变。因此,爆发或暂停宽度的空间测量提供了物体表面特征大小的可靠表示。3. 对于给定抚摸速度,平均放电率的空间分布模式(空间速率分布图,SRP)提供了波浪形表面形状的表示。单个RA和SA的SRP中相邻峰值之间的距离与波浪形表面峰值之间的距离大致相同。SA群体的平均SRP比RA群体的平均SRP能更好地表示形状。虽然SEP中的活动区域可以与与皮肤表面接触的刺激“足迹”的二维形式同构,但SRP除了对垂直于皮肤表面的第三维中的刺激特征进行编码外,与刺激形状不同构。4. 当物体的大小和形状都变化时,假设一种中枢处理机制从已针对放电率总体差异进行归一化的SRP的上升和下降阶段的斜率中提取形状的不变属性。预计这些差异会随着刺激参数的变化而出现,如压缩力、抚摸轨迹和抚摸速度。结果表明,每个波浪形表面凸面引发的SA的平均SRP的一个共同特征是,无论其半径如何,从底部到峰值以及从峰值到底部的斜率是恒定的。因此,圆柱体恒定曲率的一种可能编码是由圆柱形凸面在SA群体中引发的三角形空间反应轮廓的上升和下降阶段的斜率的恒定。

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