Manjarrez et al. (2003) Proc. of SPIE Vol.

5110, pp: xx-xx

Stochastic resonance in the spinal cord and somatosensory cortex of the

cat
E. Manjarrez*, G. Rojas-Piloni, H. Pérez, I. Méndez, Z. Hernández-Paxtián and A. Flores
Instituto de Fisiología,
Benemérita Universidad Autónoma de Puebla.
14 Sur 6301, Col. San Manuel. Apartado Postal 406. Puebla, Pue. CP 72570, México.

ABSTRACT

The aim of this study was to demonstrate the occurrence of stochastic resonance (SR) in spinal and cortical potentials
elicited by periodic tactile stimuli in the anaesthetised cat. The periodic tactile stimuli were applied on the central pad of the
hindpaw and the noisy tactile stimuli on the glabrous skin of the third hindpaw digit. This protocol allowed that the signal
and noise were mixed not in the skin but in the somatosensory regions of the central nervous system. The results show that
a particular level of tactile noise can increase the amplitude of the spinal and cortical potentials elicited by periodic tactile
stimuli. The topographical distribution of evoked potentials indicates that the effects of noise were spatially restricted. All
cats showed distinct SR behavior at the spinal and cortical stages of the sensory encoding. Such SR was abolished in the
cortical but not in the spinal recording after the sectioning of the ascending pathways. This suggests that the spinal
neurones may also contribute to the SR observed at the cortical level. The present study documents the first evidence that
the SR phenomenon occurs in the spinal and cortical somatosensory system itself and not only in the peripheral sensory
receptors.

Key words: Noise, tactile, information capacity, stochastic resonance, somatosensory, evoked potentials, spectrum map.

1. INTRODUCTION

Tactile information is transmitted from skin to the brain via relays within the central nervous system (CNS). Several lines of
evidence suggest that in cats (and to some extent in monkeys) the dorsal columns (DC) and the dorsolateral funiculus (DLF)
both contribute to tactile conditioned reflexes, two-point discrimination, tactile and proprioceptive placing, roughness
discrimination, and size discrimination 1. In cats and monkeys, tac-
Somatosensory
tile deficits are generally made much worse when combined le- Cortex
sions are made in the DC and the DLF than when a lesion is placed
in either alone 1.

Inputs to the DC system enter the spinal cord and ascend to the Thalamus
medulla, where they synapse at the gracile and cuneate nuclei. The
axons of postsynaptic cells form a fiber bundle that crosses in the Gracile and
Cuneate nuclei
brain stem to the opposite side and ascends to the thalamus. Out- Fig. 1. General view
puts of the thalamus are directed to postcentral cortical regions Lateral Cervical of two major
referred to as the somatosensory cortex.
nucleus somatic sensory
pathways in the cat
spinal cord: The
The spino-cervical tract is one of the major somatic sensory dorsal columns
pathways located in the DLF in the cat spinal cord 1. It transmits DLF
DC (DC) system and
information from cutaneous receptors to the cerebral cortex via Spino-Cervical
the spino-cervical
tract
relays in the lateral cervical nucleus and the contralateral thalamus. pathway (located
Figure 1 illustrates a highly schematic representation of the DC in the dorsolateral
and spino-cervical ascending pathways. funiculus, DLF).
Peripheral Sensory Receptors

* emanjar@siu.buap.mx; phone: +5222 22 295500 (ext 7326); fax: 5222 22 334511

Stochastic resonance (SR) is a counter-intuitive phenomenon of nonlinear systems that refers to the increase of the signal-
to-noise ratio (SNR) on the output, obtained through an increase of the noise level on the input. SR is commonly under-
stood to be the enhancement, by noise, of the response of a system to a weak input signal 2-4. Typically, the plot of SNR vs
input noise is an inverted U-like function characterised by maximal enhancement of SNR at a specific noise amplitude value.
SR type effects have been demonstrated in physical and biological systems 3,5-16.

The SR has been well described in the peripheral sensory system 17-24. However, it is not clear from these experiments
whether the electrical activity of somatosensory neurones in the CNS also exhibits the SR behavior, or whether the SR is
exclusively produced in the peripheral sensory receptors.

The SR has been shown in psychophysical experiments of cutaneous tactile sensation in humans. These studies show that
the presence of a particular non-zero level of noise may significantly enhance the ability of an individual to detect sub-
threshold tactile stimuli 25-29. Furthermore, recent electrophysiological evidence, in humans, suggests that the cortical
neurones participating in the process also exhibits SR 14. However, it is not clear from these experiments whether the
electrical activity of the spinal neurones also exhibits SR behavior, or whether the SR is produced only in the peripheral
sensory receptors. In this context, the purpose of the present study was to substantiate whether SR occurs in the spinal and
cortical somatosensory system itself, in a preparation of the anaesthetised cat. Disclosure of this phenomenon in the spinal
and cortical stages of the sensory encoding could be important because a major goal of contemporary studies of sensory
processing is to understand the transformations of signals at each level within the central nervous system (CNS).

2. MATERIALS & METHODS

2.1 Preparation

Experiments were carried out in adult cats (2.0-3.5 kg) initially anaesthetised with pentobarbitone (35 mgkg-1 of weight, i.p.).
Most procedures were reported previously 15,30,31 and are briefly described here. Guidelines contained in NIH publication on
the principles of laboratory animal care 85-23, revised in 1985, were followed throughout.

2.2 Protocol of stimulation

The noisy tactile stimuli were applied with an indenter placed on the glabrous skin of the third hindpaw digit, whereas the
periodic tactile stimuli (signal) were applied on the glabrous skin of the central pad of the hindpaw (Fig. 2A). The output of
two independent function generators provided input to the stimulators. One of these (Tektronix CFG253 together with
Master-8 AMPI) generated the test stimulus signal while, the other (Wavetek 132), supplied the superimposed noise. This

A Noise Signal B
1.0
Power (au)

0.5 103 counts

2 mN

0.0
0 20 40 60 80 100
Frequency (Hz)

Fig. 2. A, Records of the input signal and input noise. Periodic tactile stimuli were applied on the glabrous skin of the central pad of the
hindpaw. Amplitude of periodic stimuli (test stimuli) was kept constant at 3 mN, whereas different levels of tactile noise were applied
on the glabrous skin of the third hindpaw digit. B, Power spectrum and amplitude distribution of the input noise.
protocol allowed that the signal and noise were mixed not in the skin but in the somatosensory regions of the CNS (see also
Mori and Kai, 2002 16). In order to avoid a possible peripheral mixing of the noisy stimuli and the periodic stimuli due to the
elasticity of the adjoining skin, the hindlimb and the third hindpaw digit, were held in a fixed position. Figure 4C shows that
the amplitudes of the input signal (measured with a transducer) were not affected by the different levels of input noise.

2.3 Test stimuli

Mecanical test stimuli (local skin displacements) were applied on the central pad of the hindpaw. Such stimuli consisted of
single pulses with a total duration of 10 ms, delivered at a constant frequency of 2.5 Hz. We have assumed that the SR might
be present at this frequency, because at this same frequency of tactile stimulation, we observed the SR in the human brain
waves 14. In our experiments, the amplitude of the test stimulus was kept constant at 3 mN (Fig. 4B-C), between 1.2 and 1.5
times the threshold level (xT) of the afferent volley
recorded on the surface of the spinal cord. We 25 µV
have assumed that the force of 3 mN would be
sufficient to activate slowly adapting and fast
adapting mechanoreceptive afferents innervating 2 mm 0 µV
the skin 32.

2.4 Input noise

80 µV
Noise with a power spectrum ranging from 0.1-60
Hz was applied on the third hindpaw digit (Fig.
2B). The range of the standard deviation of the
noisy stimuli (σn) was from 0.1-5 mN. The inset in
Fig. 2B shows the amplitude distribution (Gaussian)
of the input noise.

2.5 Electrophysiological recordings

Two glass micropipettes filled with NaCl 1.2 M (tip

diameter, 1.0-2.5 µm; 1.2-1.7 MΩ) were used to
record spinal and cortical evoked field potentials, 25 µV

within the dorsal horn at L6 (laminae III-VI), and

within the primary somatosensory cortex (S1) in
the right posterior sigmoid gyrus (layers III-V) (Fig.
40 µV
3). The micropipette entered the hindlimb repre-
sentation of the S1 somatosensory cortex located 100 ms
0 µV
in the posterior sigmoid gyrus 33. Surface evoked 5 mm
potentials were recorded digitally on a Synamps
EEG amplifier (NeuroScan, Inc. Sterling, VA) using
64 Ag-AgCl electrodes with 500 Hz and 10 KHz
sampling frequencies (0.15 Hz high-pass filter,
Noise
70 Hz low-pass filter) (Fig. 3). Bilateral spinal Signal

evoked potentials were recorded with an array of

32 (4x8) electrodes positioned on the surface of
the lumbar L5-L7 spinal cord. Bilateral cortical
evoked potentials were recorded with two arrays
of 16 (4x4) electrodes positioned on the surface of
the posterior sigmoid gyrus. Topographical spec-
trum maps were created using Scan 4.2 Software
from NeuroScan.
Fig. 3. Scheme of the experimental arrangement in a preparation of the
anaesthetised cat. The surface electrodes and the microelectrodes were
positioned in spinal and cortical somatosensory regions where the largest
evoked field potentials (EFPs) were detected. Typical topographical spec-
trum maps (corresponding to 0-10 Hz band) are illustrated.
2.6 Stimulation scheme

The stimulation scheme consisted of sequences that lasted 20 s, during which we applied either the periodic stimulus and
the noisy stimulus simultaneously or noise alone. We applied ten sequences, each with a different noise intensity level.
Figure 4B shows input signal and noise for four different levels of noise. The presentation order of the different noise levels
was varied randomly to remove possible serial effects.

2.7 Data analysis

Data acquisition of the input noise and of the spinal and cortical potentials was performed with a sampling rate of 500 Hz and
10 KHz. Spectral analysis of the spinal and cortical activity recorded during each of the 20 s stimulation epochs was
performed. The magnitude of the input noise was quantified by means of the standard deviation of the input force (σ n of
input noise). We estimated the effect of noise upon spinal and cortical evoked potentials from the output power spectra.
SNR (1) is defined as the ratio, at the input signal frequency (2.5 Hz), of the strength of the output power spectra peak (its
area) during pulse stimulation plus noise to the mean amplitude of the output power spectra occurring during input noise
alone. Both the mean amplitude and the area were calculated in the frequency interval of ±1 Hz around the input signal
frequency. The method to calculate SNR has been described in detail in a recent publication 14.

SNR = ∫ S ( f )df (1)

N(f )

S(f) corresponds to the power spectrum of the output neuronal activity (spinal or cortical) elicited by the periodic stimuli (2.5
Hz) plus noise. N(f) is the power spectrum of the output neuronal activity (spinal or cortical) elicited by noise alone. In
addition, the Shannon’s information capacity (2) at the spinal and cortical stages of the sensory encoding was calculated.
The information capacity (Rinfo) is the maximum achievable rate of information transmission through a communication
system in bits per second. For comparison, we performed an identical analysis of the signals before and after section of the
DC and the ipsilateral DLF.

 S(f ) 
Rinfo = ∫ log 2 1 + df (2)
 N( f )

2.8 Histology

At the end of the experiment, each animal was killed with a pentobarbitone overdose and perfused with 10 % formaline. The
spinal cord and the brain were removed while leaving the recording micropipettes in place. After complete fixation and
dehydration, both the spinal cord and the brain were placed in a solution of methyl salicylate for clearing, and subsequently
cut to have both sections containing the electrodes (Figs. 4D and 4G) and lesions in the T12 segment (Fig. 7F) of the spinal
cord.

3. RESULTS & DISCUSSION

We examined the effects of noise (applied on the glabrous skin of the third hindpaw digit) on the amplitude of the spinal and
cortical potentials produced by periodic tactile stimuli (applied on the glabrous skin of the central pad of the hindpaw) (Fig.
4A). The traces in Figs. 4E and 4H show averages of the spinal and cortical evoked potentials produced by the application
of the stimuli illustrated in Fig. 4B. The spinal evoked potentials were recorded at a depth of 1250 µm within the dorsal horn
(Fig. 4D), and the cortical evoked potentials were recorded at a depth of 1400 µm within the S1 somatosensory cortex (Fig.
4G). Figs. 4F and 4I illustrate plots of percentage changes of EFP-amplitude relative to control (taken as 100 %) versus σn for
one experiment. In these plots the control was considered as the response to the test stimulus alone (σn = 0). Each plot (Figs.
4H-I) exhibited a maximal enhancement of EFP-amplitude at a specific noise amplitude value. Note that spinal and cortical
evoked potentials were facilitated within a particular interval of noise amplitudes. An average of the spinal and cortical
facilitation peak was calculated for all animals (120.3±10.3 % (n=4) and 124.1±15.1 % (n=3), respectively; mean ± SD).

In order to test the hypothesis that the noise has a large impact on the amplitude of the spino-cortical evoked responses, we
plotted topographical power spectrum maps (see method in Fig. 3) for ten different levels of noise. Figure 5 displays the
results obtained from one experiment, in which the noise and signal were applied with the same protocol of stimulation as
in Fig. 4A. Figures 5D,H, L and 5P,T,X illustrate cortical and spinal power spectrum maps for three different levels of noise
(σn = 0.07, σn = 0.60 and σn = 2.30). These maps were computed in the frequency band of 0-10 Hz. These maps show that the
amplitudes of the cortical and spinal evoked responses were increased when an appropriate level of noise (σn = 0.60) was

A NOISE + SIGNAL D G
3 mN Ans
500 ms Cru

SI

1 mm 1 mm
B E H
INPUT OUTPUT
Input Noise + Input Signal Spinal EFP Cortical EFP
σn = 0

σn = 0.75

σn = 2.01

σn = 4.70

4 mN
80 µV 40 µV
20 ms

Input Signal vs Input Noise Spinal EFP vs Input Noise Cortical EFP vs Input Noise
C F I
EFP amplitude (%)

150 150 150

of input signal (%)

EFP amplitude (%)

Amplitude

100 100 100

50 50 50
0 1 2 3 4 5 0 1 2 3 4 5 0 1 2 3 4 5

Input noise σn (mN) Input noise σ n (mN) Input noise σ n (mN)

Fig. 4. A, Scheme of the experimental arrangement. The periodic tactile stimuli were applied on the central pad of the hindpaw and the
noisy tactile stimuli on the glabrous skin of the third hindpaw digit. This protocol allowed that the signal and noise were mixed not in the
skin but in the somatosensory regions of the CNS. Drawings in D and G are histological reconstructions of the electrode tracks within the
dorsal horn at L6 and within the S1 somatosensory cortex. The electrodes were positioned in spinal and cortical somatosensory regions
where the largest evoked field potentials (evoked potentials) were detected. E and H, Averages (n=32) of spinal and cortical evoked
potentials for four different levels of noise (i.e., signal plus noise, σn= 0, σn =0.75, σn =2.01, σn=4.7; see Fig. 4B). C, Input signal versus
input noise. F, Facilitation of spinal evoked potentials (amplitude) versus σn for one experiment. Interrupted line represents the magni-
tude of a 95 % confidence interval. I, The same as F, but for the cortical evoked potentials.
 A Input E I
signal
σ = 0.07 σ = 0.6 σ = 2.3 3 mN
Input
noise

B Cortical F J
EFP
100 µV
C G K
1.0 1.0 1.0 500 ms
Power (au)

0.5 0.5 0.5

0.0 0.0 0.0

0 5 10 15 0 5 10 15 0 5 10 15
25 µV
Frequency (Hz)
D H L

0 µV

M Input Q U
signal
σ = 0.07 σ = 0.6 σ = 2.3 3 mN
Input
noise
N Spinal R V
EFP 30 µV
O S W
1.0 1.0 500 ms
1.0
Power (au)

0.5 0.5 0.5

0.0 0.0 0.0

0 5 10 15 0 5 10 15 0 5 10 15
Frequency (Hz)

P T X
25 µV

0 µV

Fig. 5. Effects of noise on cortical and spinal electrical activity recorded with the arrangement of 64 electrodes depicted in Figure 6A. The
effects of noise upon spinal and cortical evoked potentials were quantified from the output power spectra (Figs. 5C,G,K,O,S and W) and
the power spectrum maps (Figs. 5D,H,L,P,T and X). A-D and M-P, Results obtained for σn = 0.07. E-H and Q-T, Results obtained for
σn = 0.60. I-L and U-X, Results obtained for σn = 2.30. Power spectra were computed in the frequency band of 0 to 15 Hz and the
topographical power spectrum maps in the frequency band of 0 to 10 Hz. Arrows indicate the regions of maximal activity (>25µV).
 B Cortical SNR vs Input Noise

10 10 10 10 10 10
5 5 5 5 5 5
0 0 0 0 0 0

10 10 0.0 2.5 5.0 10 0.0 2.5 5.0 10 0.0 2.5 5.0

0.0 2.5 5.0 0.0 2.5 5.0 0.0 2.5 5.0
5 5 5 5

0 0 0 0
10 10
0.0 2.5 5.0 10 0.0 2.5 5.0 10 0.0 2.5 5.0 0.0 2.5 5.0
5 5
5 5
0 0
0 0
10 10 0.0 2.5 5.0 10 0.0 2.5 5.0 10
0.0 2.5 5.0 0.0 2.5 5.0 5 5
5 5
SNR

0 0
0 0 10
10 10 10
0.0 2.5 5.0 0.0 2.5 5.0
0.0 2.5 5.0 0.0 2.5 5.0 5
5 5 5
0
0 0 0
10 0.0 2.5 5.0 10
10 10 0.0 2.5 5.0
0.0 2.5 5.0 0.0 2.5 5.0 5 5
5 5
0 0
0 0
10 10 10 10 0.0 2.5 5.0 10 0.0 2.5 5.0 10
0.0 2.5 5.0 0.0 2.5 5.0
5 5 5 5 5 5
0 0 0 0 0 0

0.0 2.5 5.0 0.0 2.5 5.0 0.0 2.5 5.0 0.0 2.5 5.0 0.0 2.5 5.0 0.0 2.5 5.0

Input noise σn (mN) Input noise σn (mN)

A C Spinal SNR vs Input Noise

14
7
0

14
7
0

14
7
0

14
7
0
SNR

14
7
0

14
7
0

14
7
0
14
7
0
0 2 4 0 2 4 0 2 4 0 2 4

Signal Noise Input noise σn (mN)

Fig. 6. SNR versus input noise for cortical and spinal electrical activity recorded with the array of electrodes depicted in Figure 6A. B,
Cortical output SNR versus σn . C, Spinal SNR versus σn. These results were obtained from the same experiment illustrated in Figure 5.
SNR values were calculated with eq. (1). Further explanations in text.
used. The power spectra illustrated in Figs. 5C,G,K and Figs. 5O,S,W were calculated from the recordings illustrated in Figs.
5B,F,J and Figs. 5N,R,V, respectively. The recordings illustrated in Figs. 5B,F,J and Figs. 5N,R,V were obtained from the
electrodes indicated by arrows in Fig. 5H and Fig. 5T. We characterized SR by examining the cortical and spinal output SNR
obtained for each input noise level (σn). Figures 6B-C show the cortical and spinal output SNR (versus σn). SNR values were
calculated with the power spectra of the signals recorded from the 64 electrodes illustrated in Fig. 6A. The plots in Figs. 6B,C
were placed in the same arrangement as the electrodes in Fig. 6A. In most cases (Fig. 6B, Right; and Fig. 6, Left), the SNR
versus input noise plots were typical of SR phenomena. Similar plots were obtained in two other experiments. These results
suggest that the effects of noise were spatially restricted. This modulation of the SNR suggests, quite strongly, that the
information transmitted through the somatosensory system (i.e., Rinfo) is also modulated by the input noise.

Furthermore, we studied the effects of noise on the spinal and cortical potentials produced by periodic tactile stimuli, before
and after sectioning of the DC and the ipsilateral DLF. We used the same protocol of stimulation illustrated in Figs. 4A-B. All
animals we examined exhibited SR-type behavior elicited by tactile stimulation upon spinal and cortical activity recorded
with micropipettes. Figure 7A illustrates typical recordings and power spectra obtained from the spinal and cortical evoked
potentials. Figure 7 was obtained from three cats (indicated by different symbols). Figures 7B-C (and 7D-E), illustrate the

Before section

Cortical S1
A B D
8 400
Po wer (au)

6
SNR

4 200

Rinfo
80 µV
500 ms
0
2 3 4 0 0
Hz
0 1 2 3 4 5 0 1 2 3 4 5
Input noise σn (mN) Input noise σn (mN)

Spinal L6
C E
P ow er (au)

9
14 600
SNR

40 µV
0 7 400
Rinfo

2 3 4
Hz
0 200
0 1 2 3 4 5 0 1 2 3 4 5
Signal Noise Input noise σn (mN) Input noise σn (mN)

Fig. 7. A, Scheme of the experimental arrangement. Local field potentials were recorded within the primary somatosensory cortex (S1)
and within the dorsal horn in the spinal lumbar segment L6. Plots in Fig. 7A show typical power spectra obtained from the field potentials
elicited by tactile stimuli at 2.5 Hz. Cortical (B) and spinal (C) output SNR versus σ n, for three different cats (indicated by different
symbols) in control conditions (before section). D-E, The same as B-C, but for Shannon’s information capacity (R info). SNR was
calculated with eq. (1). R info was calculated with eq. (2). Further explanations in text.
output SNR (and Rinfo) at the two stages of sensory encoding: cortical (Fig. 7B, Fig. 7D; filled symbols) and spinal (Fig. 7C,
Fig. 7E; open symbols) versus σn in control conditions. Similarity in the inverted U-like feature of the graphs calculated from
each of the cats is such that a qualitative general description of these curves may apply to all. As the noise amplitude
increases, SNR values become larger. Hence, a positive slope and an upsurge of the function can be observed as the curve
rises steeply and turns convex. A maximum value of SNR is reached and the slope becomes zero within a particular interval
of noise amplitudes. Beyond such peak, with higher noise amplitudes, the slope becomes negative as the curve subsides
gradually.

SR after sectioning of DC and IDLF

The SR behavior was abolished in the cortical (Fig. 8C, Fig. 8E) but not in the spinal (Fig. 8D, Fig. 8F) recordings after
sectioning of DC and ipsilateral DLF (Fig. 8A-B). Averages of the cortical SNR and Rinfo peak values before section (in the
band from 1 to 2mN; Figs. 7B and 7D) were calculated for three animals (4.8±1.4 and 303.5±68.7 bits/s, respectively). These
averages were different (P<0.001; Students t-test) from the averages of the cortical SNR and Rinfo values (0.5± 0.5 and
85.4±21.3 bits/s, respectively; in the band from 1 to 2mN; Figs. 8C and 8E) obtained after the sections illustrated in Fig. 8B.

After section

A Cortical S1
C E
8 400
T12

Rinfo
B 4 200
SNR

0 0
0 1 2 3 4 5 0 1 2 3 4 5
T12 Input noise σn (mN) Input noise σn (mN)

1 mm
Spinal L6
D F
14 600
7
Rinfo

400
SNR

0 200
Signal Noise 0 1 2 3 4 5 0 1 2 3 4 5
Input noise σn (mN) Input noise σn (mN)

Fig. 8. A, Scheme of the experimental arrangement. Cortical (C) and spinal (D) output SNR versus σ n, for the same experiments
illustrated in Figure 7, but after section (at T12) of the DC and the ipsilateral DLF. B, Drawings from the histological sections obtained
from three cats (indicated by different symbols). E-F, The same as C-D, but for Rinfo. Note that SR disappears in the cortical (filled
symbols), but not in the spinal (open symbols) stages after section. Further explanations in text.
SR occurs in the spino-cortical somatosensory system itself and not only in the peripheral sensory receptors

Our results show that certain range of noise can increase spinal and cortical evoked field potentials (Figs. 4F and 4I). This
result provides a possible explanation of the SR observed in SNR and Rinfo of the spinal and cortical activity evoked by
mechanical tactile stimuli (Figs. 7B-E). In particular, we demonstrated that the SR embodies electrical processes of the
spino-cortical somatosensory system itself (see also Manjarrez et al. 2002 15). We used a protocol, as shown in Fig. 4A, that
allowed that the signal and noise were mixed not in the skin but in the somatosensory regions of the CNS. In this context,
our results show that the SR occurs in the spinal and cortical somatosensory system itself and not only in the peripheral
sensory receptors.

To our knowledge, the present investigation documents the first explicit explanation of the occurrence of SR phenomena
concerning the electrical activity of the spinal and cortical stages of sensory encoding in an in vivo preparation. Our results
agree well with the psychophysical findings of Collins et al. 25-27, who described that the ability of an individual to detect a
sub-threshold tactile stimulus can be enhanced by introducing a particular level of noise. Furthermore, our results are
consistent with recent evidence that the SNR of cortical activity elicited by mechanical tactile stimuli in humans is optimized
by the presence of noise 14.

Our experiments show that the range of noise intensities necessary for the enhancement of SNR was within physiological
limits (1 to 4mN), in the same range in which noise can improve tactile sensation in humans 14,25. Furthermore, our experi-
ments show that the SR was evident at 2.5 Hz, at the same frequency of tactile stimulation where the SR in the human brain
was observed 14. These evidences suggest that noise could play a major physiological role in tactile sensation by soma-
tosensory neurones, both in cats and humans.

Several causes may explain the different profiles observed in the SNR and Rinfo graphs obtained from different experiments
(Figs. 7B-E). The diversity of these profiles between animals may be attributed to their different sensitivity to stimuli,
dissimilarities in skin elasticity, receptor density, and irregularity of the background activity at the spinal, brainstem,
thalamic, and cortical levels.

Participation of dorsal horn spinal neurones in the mechanism of generation of SR at the cortical level

Previous work from our laboratory 30,31 has shown that the somatosensory cortical neurones can be driven by dorsal horn
spinal neurones with spontaneous activity through the DC and the spino-cervical tract. The cell bodies of origin of the
spino-cervical tract are located in the lumbo-sacral dorsal horn (laminae IV-VI) 34, in the same region where the neurones
producing spontaneous negative cord dorsum potentials are located. These observations show that the first synapse for
most tactile afferents is also located within the dorsal horn. In this context, the present study suggests that such dorsal
horn spinal neurones of the spino-cervical tract may also contribute to the mechanism of generation of SR at the cortical
level. Therefore, the present results (Fig. 8) are consistent with this possibility, because the SR disappears in the cortical
but not in the spinal recording after section of the DC and the ipsilateral DLF 30.

4. CONCLUSION

We conclude that the SR may occur in the spino-cortical somatosensory system itself, in a preparation of the anaesthetised
cat.

ACKNOWLEDGEMENTS

This work was partly supported by the following grants: CONACyT J36062-N (E.M), Fondo-Ricardo J. Zevada (A.F),
FOMES-BUAP and FIUPEA P/PIFI 2001-22-FI-17, México.
 REFERENCES

1. W.D. Willis and R.E. Coggeshall, “Sensory Mechanisms of the Spinal Cord”, Plenum Press, New York, 1991.
2. K. Wiesenfeld and F. Moss, “Stochastic resonance and the benefits of noise: from ice ages to crayfish and SQUIDs”,
Nature 373, pp. 33-36, 1995.
3. L. Gammaitoni, P. Hanggi, P. Jung and F. Marchesoni, “Stochastic resonance”, Rev Mod Phys 70, pp. 223-287, 1998.
4. V.S. Anishchenko, A.B. Neiman, F. Moss and L. Schimansky-Geier, “Stochastic resonance: noise-enhanced order”,
Physics-Uspekhi 42, pp. 7-36, 1999.
5. J.P. Segundo, J.F. Vibert, K. Pakdaman, M. Stiber and O. Diez-Martínez, “Noise and the neurosciences: a long history,
a recent revival and some theory”. In: Pribram, K.H. (ed) Origins: Brain and Self Organization, Laurence Elbaum
associates Publishers, pp. 300-331. Hillsdale, N.J., 1994.
6. D.F. Russell, L.A. Wilkens and F. Moss, “Use of behavioural stochastic resonance by paddle fish for feeding”, Nature
402, pp. 291-294, 1999.
7. G. Winterer, M. Ziller, H. Dorn, K. Frick, C. Mulert, N. Dahhan, W.M. Herrmann and R. Coppola, “Cortical activation,
signal-to-noise ratio and stochastic resonance during information processing in man”, Clin Neurophysiol 110, pp.
1193-1203, 1999.
8. W.C. Stacey and D.M. Durand, “Stochastic resonance improves signal detection in hippocampal CA1 neurons”, J
Neurophysiol 83, pp. 1394-1402, 2000.
9. I. Hidaka, D. Nozaki and Y. Yamamoto, “Functional stochastic resonance in the human brain: noise induced sensitiza-
tion of baroreflex system”, Phys Rev Lett 85, pp. 3740-3743, 2000.
10. I. Hidaka, S. Ando, H. Shigematsu, K. Sakai, S. Setoguchi, T. Seto, Y. Hirooka, A. Takeshita and Y. Yamamoto, “Noise-
enhanced heart rate and sympathetic nerve responses to oscillatory lower body negative pressure in humans”, J
Neurophysiol 86, pp. 559-564, 2001.
11. Y. Yamamoto, I. Hidaka, N. Iso-o, A. Komai, R. Soma and S. Kwak, “Noise-induced compensation for postural hypoten-
sion in primary autonomic failure”, Brain Res 945, pp. 71-78, 2002.
12. P. Hanggi, “Stochastic resonance in biology. How noise can enhance detection of weak signals and help improve
biological information processing”, Chemphyschem 3, pp. 285-290, 2002.
13. L.M. Ward, A. Neiman and F.Moss, “Stochastic resonance in psychophysics and in animal behavior”, Biol Cybern 87,
pp. 91-101, 2002.
14. E. Manjarrez, O. Diez-Martínez, I. Méndez and A. Flores, “Stochastic resonance in human electroencephalographic
activity elicited by mechanical tactile stimuli”, Neurosci Lett 324, pp. 213-216, 2002.
15. E. Manjarrez, G. Rojas-Piloni, I. Méndez, L. Martínez, D. Vélez, D. Vázquez, and A. Flores, “Internal stochastic reso-
nance in the coherence between spinal and cortical neuronal ensembles in the cat”, Neurosci Lett 326, pp. 93-96, 2002.
16. T. Mori and S. Kai, “Noise-induced entrainment and stochastic resonance in human brain waves”, Phys Rev Lett 88,
pp. 1-4, 2002.
17. J.K. Douglass, L. Wilkens, E. Pantazelou and F. Moss, “Noise enhancement of information transfer in crayfish mecha-
noreceptors by stochastic resonance”, Nature 365, pp. 337-340, 1993.
18. J.E. Levin and J.P. Miller, “Broadband neural encoding in the cricket cercal sensory system enhanced by stochastic
resonance”, Nature 380, pp. 165-168, 1996.
19. P. Cordo, J.T. Inglis, S. Verschueren, J.J. Collins, D.M. Merfeld, S. Rosenblum, S. Buckley and F. Moss, “Noise in human
muscle spindles”, Nature 383, pp. 769-770, 1996.
20. J.J. Collins, T.T. Imhoff and P. Grigg, “Noise–enhanced information transmission in rat SA1 cutaneous mechanorecep-
tors via aperiodic stochastic resonance”, J Neurophysiol 76, pp. 642-645, 1996.
21. M. Juusola and A.S. French, “The efficiency of sensory information coding by mechanoreceptor neurons”. Neuron
18, pp. 959-968, 1997.
22. C. Ivey, A.V. Apkarian and D.R. Chialvo, “Noise-induced tuning curve changes in mechanoreceptors”, J Neurophysiol
79, pp. 1879-1890, 1998.
23. F. Jaramillo and K. Wiesenfeld, “Mechanoelectrical transduction assisted by Brownian motion: a role for noise in the
auditory system”, Nature Neurosci 1, pp. 384-388, 1998.
24. S. Bahar, A. Neiman, L:A. Wilkens and F. Moss, “Phase synchronization and stochastic resonance effects in the
crayfish caudal photoreceptor”, Phys Rev E 65, pp. 050901, 2002.
25. J.J. Collins, T.T. Imhoff and P. Grigg, “Noise-enhanced tactile sensation”, Nature 383, pp. 770, 1996.
26. J.J. Collins, T.T. Imhoff and P. Grigg, “Noise-mediated enhancements and decrements in human tactile sensation”, Phys
Rev E 56, pp. 923-926, 1997.
27. K.A. Richardson, T.T. Imhoff, P. Grigg and J.J. Collins, “Using electrical noise to enhance the ability of humans to detect
subthreshold mechanical cutaneous stimuli” Chaos 8, pp. 599-603, 1998
28. W. Liu, L.A. Lipsitz, M. Montero-Odasso, J. Bean, D.C. Kerrigan and J.J. Collins, “Noise-enhanced vibrotactile sensi-
tivity in older adults, patients with stroke, and patients with diabetic neuropathy”, Arch Phys Med Rehabil 83, pp. 171-
176, 2002.
29. N.T. Dhruv, J.B. Niemi, J.D.Harry, L.A. Lipsitz and J.J. Collins, “Enhancing tactile sensation in older adults with electri-
cal noise stimulation”, NeuroReport 13, pp. 597-600, 2002.
30. E. Manjarrez, G. Rojas-Piloni, D. Vázquez and A. Flores, “Cortical neuronal ensembles driven by dorsal horn spinal
neurones with spontaneous activity in the cat”, Neurosci Lett 318, pp. 145-148, 2002.
31. E. Manjarrez, G. Rojas-Piloni, L. Martínez, D. Vázquez, D. Vélez, I. Méndez and A. Flores, “Amplitude of somatosensory
cortical evoked potentials is correlated with spontaneous activity of spinal neurones in the cat”, Neurosci Lett 323, pp.
187-190, 2002.
32. M. Trulsson, “Mechanoreceptive afferents in the human sural nerve”, Exp Brain Res 137, pp. 111-116, 2001.
33. D.J. Felleman, J.T. Wall, C.G. Cusick and J.H. Kaas, “The representation of the body surface in SI of the cats”. J
Neurosci 3, pp. 1648-1669, 1983.
34. R.N. Bryan, D.L. Trevino, J.D. Coulter and W.D. Willis, “Location and somatotopic organization of the cells of origin of
the spino-cervical tract”, Exp Brain Res 17, pp. 177-189, 1973.