Rat cutaneous RA afferents activated by two-dimensional skin stretch.

Skin develops biaxial stresses and strains when stretched. Rapidly adapting cutaneous mechanoreceptor neurons are known to be stretch sensitive, yet in the past, they have been studied using stretch stimuli applied along only a single direction. In this study, cutaneous rapidly adapting mechanoreceptors were studied in preparations of isolated skin in which the skin was stretched dynamically using biaxial stretch stimuli and in which loads and displacements were measured along two directions. Stretch stimuli followed a pseudo-Gaussian waveform and were applied along either one or two directions simultaneously. Associations between spikes and mechanical variables were determined using multiple logistic regression. When the skin was actuated along a single direction, holding the orthogonal axis fixed, spike responses were strongly associated with mechanical variables along the actuated direction. The variables were stress and its rate of change, the rate of change of strain, and the product of stress and its rate of change, which is proportional to strain energy density. When the skin was stretched along a single direction, spikes were very poorly associated with stress variables measured along the direction orthogonal to the stretch. Afferents showed weak directional selectivity: they were slightly more responsive to the variable stress along the circumferential direction of the hindlimb. When the skin was stretched biaxially (i.e., along both directions simultaneously) with identical pseudo-Gaussian noise stimuli, neuronal responses were associated with the same variables as above, but the associations were weaker.