M. Hulliger et Tk. Baumann, COMPONENT ANALYSIS OF THE RESPONSES OF SENSORY NEURONS TO COMBINED SINUSOIDAL AND TRIANGULAR STIMULATION, Journal of neuroscience methods, 53(2), 1994, pp. 173-188
A method for quantitative estimation of sensory neuron sensitivity to
small sinusoidal stimuli in the presence of sizable background drift (
in the stimulus or response) was developed. The performance of the met
hod was tested by analyzing the responses of 17 muscle spindle primary
(Ia) afferent neurons to concomitant sinusoidal and triangular stretc
hing of the soleus muscle. The efficacy and accuracy of several variat
ions of the method were examined. The variations included the use of p
robability density (PD) and average frequency (AF) histograms as the b
asis for calculations and two different algorithms for the decompositi
on of responses to combined sinusoidal and triangular stimulation. One
algorithm called the 'inherent-drift' method exploited the inherent h
alf-cycle repeat property of a sine wave to extract the drift componen
t. Another algorithm called the 'forced-drift' method first estimated
the drift by linear regression to a response to triangular stimulation
alone. The drift estimate (a slope value) was then subtracted from th
e response to combined sinusoidal and triangular stimulation of the sa
me triangular (background) velocity. A comparison of the performance o
f the drift correction method applied either to PD or AF histograms re
vealed no significant differences in the estimates of sinusoidal modul
ation. The limitations of the AF method were manifest primarily by pha
se lags at low mean levels of action potential discharge. Calculation
of the response parameters using the 'inherent-drift' correction proce
dure proved straightforward as long as there were at least two pairs o
f non-empty bins in the sine-cycle histograms on which to base the est
imate of drift. The method remained effective in determining sinusoida
l sensitivity in the face of distinct non-linearities (harmonic distor
tions) in the sine-cycle histograms. However, estimates of slope and t
he extraction of sinusoidal phase by the 'inherent' slope correction m
ethod became subject to large errors. Under such circumstances, more r
eliable estimates could be obtained by using the forced drift-correcti
on method instead. The importance of extracting the drift component pr
ior to estimating the sinusoidal response parameters was evaluated exp
erimentally and theoretically. In general, omission of a drift correct
ion introduced a large bias in the estimates of the phase of sinusoida
l response, whereas the estimate of sinusoidal modulation was rather i
nsensitive. Experimental findings were fully accounted for by theoreti
cal considerations. Analytically derived relationships identified low-
and high-risk regions more clearly for the estimate of sinusoidal mod
ulation than of phase. The relationship between biased modulation esti
mate and underlying drift showed minima characteristics with a low-ris
k region, where absolute errors and dependence on slope variations wer
e small. The precise shape of these characteristic curves depended on
the phase value of the sinusoidal response component. This suggests th
at in situations where the phase of sinusoidal responses can be estima
ted independently, experimental paradigms can be optimized by adjustin
g the phase of sinusoidal stimuli, so as to minimize the effects of co
mplete omission of drift correction or of inappropriate drift determin
ation on the estimates of modulation. In contrast, for phase estimatio
n low-risk regions are very narrow, indicating that inappropriate drif
t compensation almost invariably leads to significant errors in phase
estimates. In its current form, the method is directly applicable to s
ensory responses which consist of rate-modulated trains of action pote
ntials. Nevertheless, the same principles used to develop the method c
ould be applied to sensory responses in the form of receptor or genera
tor potentials. The method should therefore be applicable to the analy
sis of a wide variety of sensory systems.