Burden Neurological Institute, Bristol, England

THE CONVERGENCE AND INTERACTION OF VISUAL,
AUDITORY, AND TACTILE RESPONSES IN HUMAN

NONSPECIFIC CORTEX
W. Grey Walter
Burden Neurological Institute, Bristol, England
Introduction
The relation between the electrical responses in the human brain and
mental processes of integration and association is perhaps the most chal-
lenging of all the problems facing neurophysiologists today. Until quite
recently the prospect of unifying physiological and psychological concepts
by electro-encephalographic ( EEG ) techniques seemed to be receding
because of the baffling complexity of the intrinsic electrical rhythms, and
this difficulty has still not been overcome. The most intriguing, and at
the same time most elusive, of the properties of these rhythms is that,
although often remarkably constant in their variations with respect to
time, they fluctuate also, in a much less regular manner, within the three
dimensional space of the brain. The continuous analysis and display of
such a phenomenon presents serious difficulties, and no method has yet
given entirely satisfactory results.
The incentive to construct adequate equipment and to design and per-
form crucial experiments is undoubtedly limited by the uncertainty as
to whether the information likely to be obtained in this way is essential
to understanding brain mechanisms; few experimenters imagine that it
would be sufficient.
However difficult they may be to unravel, the time-and space-relations
of the intrinsic brain rhythms cannot be ignored, if only because so intri-
cate and orderly an arrangement seems unlikely to have survived the
struggle for existence with no functional identification. The awkward fact
that these rhythms are highly individual-even to the extent of total
absence in some people-must also be accepted as a fascinating part of the
enigma, where personality and disposition are essential features in the
architecture of human physiology.
The introduction of more versatile and sophisticated methods for ex-
tracting information about transient components of the EEG inevitably
diverted interest from the background activity because, by definition, the
methods of transient analysis tend to efface or at least diminish those
rhythmic features which are most prominent in a conventional record.
This unfortunate technical incompatibility has discouraged correlative
studies, but interaction between the multitude of intrinsic rhythms and
the responses evoked by sensory stimuli seems an indispensable part of
the cerebral information-flow system. These influences are complex and
often reciprocal; intrinsic rhythms of excitability-variation can modify the
amplitude and latency of evoked responses, and the arrival of signals in an
afferent channel can modulate the amplitude, phase, and frequency of
320
Walter: Auditory and Tactile Responses 321
intrinsic rhythms. These effects are typically elusive because of the “cere-
bral uncertainty principle”; any event which affects the brain ensures that
another similar event will have a different effect. In the present state of
knowledge a crude analogy may be drawn between these inter-relations
and the operation of a Vehicle Actuated Traffic Signal (VATS) system.
In such a system a regular time-sequence of “stop” and “go” signals can
be over-ridden by the approach of a vehicle toward an intersection. In
the absence of traffic the regular rhythm provides an equal chance of a
clear run in either direction, but the first vehicle to approach an inter-
section biasses the system in its favour, and the rhythm of the signals can
be “driven” by the vehicle proceeding from one crossing to another. When
traffic is very heavy however, the Vehicle Actuated mode is incompetent
and the time cycle supervenes, again providing every vehicle with an
equal chance of reaching its destination. Such problems are familiar to
traffic engineers, who are only too conscious of the enormous complexities
that develop in any but the simplest cases of regular orthogonal thorough-
fares with equal and constant traffic density. No driver in a big city would
underestimate the difficulty of providing an optimum flow rate for all at
all times. We must assume that in the human brain, with a population and
traffic density far greater than that of all terrestrial cities combined, a
solution to this problem has been elaborated, and, further, that some at
least of the failures in brain transactions-the various clinical states-may
arise from the innate inadequacy or breakdown under stress of some
traffic-control device.
The basic fact about any communication system is that all signals have
an origin and a destination. In the classical or “pre-reticular” physiology
of sensation the origin of sensory signals was assumed to be in the receptor
and their destination in the specific cortical sensory zone. We know now
that not all signals from the receptors reach the primary sensory regions
and, further, that most sensory signals can find their way, with surprisingly
little delay, to many regions far from their expected destination, where
they have a rendezvous with other signals of different provenance and
modality. These processes of sensory dispersion, convergence, and con-
tingent interaction in human subjects have been investigated in some
detail in the hope of establishing a working hypothesis of sensory inte-
gration and adaptive association. Some of the results have already been
reported elsewhere (Walter, 1963).
Methods
Material. Records have been taken from over 100 subjects, male and
female aged a few months to 50 years. Of these, fifty were normal vol-
unteers, including 30 school children. The clinical group included 30
autistic children as well as fifteen patients in whose brains 68 gold
electrodes had been implanted for therapeutic purposes (Crow, Cooper
and Phillips, 1961, FIGURE 1). In this latter group the majority of the
patients were suffering from chronic intractable psychoneurotic dis-
322 Annals New York Academy of Sciences
FIGURE1. Lateral view of electrode sheaves implanted in the brain of an epileptic

patient. The frontal sheaves indicate the plane and depth of the contacts in pre-
frontal nonspecific cortex. Each sheaf provides from five to seven contacts.
turbances but were free from organic brain disease; there were also three
cases of temporal lobe epilepsy with chronic implanted electrodes. Two
cases of “reading epilepsy” were examined and one patient with a severe
memory defect following influenza1 encephalitis.
In the normal adult group were three deep-trance hypnotic subjects,
two of them trained by Stephen Black for other physiological experiments.
The studies of normal and disturbed children were part of a long-term
research on brain maturation in relation to congenital or precocious
mental disorder.
Technique. Conventional EEG records were obtained with an Offner
Type T eight-channel instrument, fitted with a two-channel BNI-Faraday
automatic wave analyser. All channels were analysed with the eyes open
and shut and during mental activity. Two channels were then left for EEG
recording and analysis, the third was connected to a monitor circuit so as
to indicate stimuli and averaging epochs, the fourth to a bridge circuit to
indicate skin resistance and its changes (PGR), the fifth to electrodes on
the muscles of the left forearm to indicate operant movements by the
subject, the sixth to a mean-zero cardiotachometer, the seventh to a
pneumographic transducer around the chest of the subject, and the
eighth to a time marker.
In normal subjects and patients without intracerebral electrodes, the
two EEG channels were usually connected to electrodes on the vertex and
mid-occiput respectively with a common reference on the right mastoid
protuberance. Several other montages were tried but none is considered
entirely satisfactory, since the fields of the evoked potentials in nonspecific
areas are so widespread that no part of the head is quite unaffected by
them. The interpretation of scalp potentials was aided by simultaneous
records from intracerebral electrodes in fifteen cases. In these, with 68
electrodes in various parts of the frontal, temporal and sometimes occi-
pital lobes, records could be made with a trustworthy intracerebral average
reference ( AR) since it could be shown that only a minority of electrodes
were involved in the evoked responses and these could be eliminated from
the average circuit. In these cases, too, records from the scalp could be
obtained with an intra-cerebral AR, thus avoiding the ambiguities intro-
duced by field diffusion over the scalp.
The two EEG channels were connected to a two-channel Electronic
Average Response Indicator ( EAR1) using two barrier-grid storage tubes
in an arrangement already described (Cooper and Warren, 1961; Cooper
and Walter, 1962). A valuable feature of this device is that, by suitable
switching, the average values of responses in the two channels can be
written out immediately after a selected number of stimulus presentations
at a higher or lower paper speed and increased gain so as to obtain the
optimum clarity and resolution in both co-ordinates. It can be shown that
in the presence of random signals evenly distributed over the pass-band of
the transmission system (“white noise”), the “noise” pentration or signal-
noise gain of an averaging system for a time-coherent signal is proportional
to the square-root of the number of observations. The greatest relative
advantage is therefore obtained with a small number of observations and
the advantage rises very slowly with increasing numbers, while on the
other hand the likelihood of condition-constancy in the brain diminishes
rapidly with repetition. For these reasons most observations have been
made by averaging sets of twelve evoked responses, giving a minimum
signal-noise gain of about 3.5.In practice, the random components are
rarely “white” and many are isolated transients of waveform similar to
the evoked potentials themselves; in fact these are probably evoked re-
sponses, due to “spontaneous” or uncontrolled sensory and intracerebral
stimuli. The signal-noise gain therefore varied between 3.5 and the number
of samples, that is, twelve (FIGURE 2), which would be the figure if all
the interference were transients without coincidence. For most experi-
ments the averaging epoch was two seconds so as to provide time for
presentation of two or more stimuli, as well as a section before stimulation
as an indication of the intrinsic noise level. An automatic programmer was
used to ensure irregularity of intervals between samples but consistency
between one set and another; the interval varied from 5 to 10 seconds.
The stimulus generator was synchronised electrically with the time-base
of the storage-tube scanner, so that the stimuli always fell at the selected
instants in the average epoch (FIGURE 3 ) .
The first set of twelve samples was taken without stimuli and the
primary gain was adjusted so as to produce a noise-level in the average
of less than about one fifth full scale deflection peak-to-peak.
I
T 1 1 . I I .
0 1 5- 10 I2 10 25
--
FIGURE 2. The relation between number of samples and signal-noise gain oh-
tained by averaging. With “white noise” the gain rises only as the square-root of the
number of samples, while if the noise is due to random isolated transients the gain
can vary directly as the number. The operational conditions lie somewhere between
these limits. N = number of observations.
-w
I
I
I
-- Flash
.,
2secs,&
Clicks
’I
II
~ FUSNl 2 3 CLlClI 2 3
6
f bCr 2 Fbhl 1 3 (88 FbCLD
I .t V1 noun c
FIGURE 3. Schematic illustration of the stimulus program for a standard experi-

ment. Each presentation is during a two-second epoch over which the signals in two
EEG channels are stored in the averager tubes. Twelve such epochs, at irregular
intervals, make up a set of presentations lasting about two minutes; the average of
the twelve epochs is written out at once on the same record. The sequence of sets, as
indicated in the bottom diagram, composes a single experiment to ascertain the ex-
tent and nature of the effects described in the text.
The programmer was then set to provide visual stimulation by flicker
from a stroboscope at 15 flashes per second for about one second in the
middle of each of the twelve two-second samples.
This was repeated for two or three sets so as to ascertain the rate and
degree of “habituation” to visual stimuli. The duration of each set of
twelve presentations at irregular intervals was about two minutes.
The next few sets were taken with auditory stimuli in the form of repeti-
tive clicks in earphones, also at 15 per second. These clicks were esti-
mated at about 80 db intensity; subjectively they were loud enough to be
startling at first but not painful. In some cases lower intensities were used
and with deaf patients considerably higher ones. In most people the rate
and degree of habituation for auditory stimuli are lower than for visual
ones and few subjects showed complete habituation of the nonspecific
responses even after three or four sets, that is 35 to 50 stimuli. The se-
quence of sets of visual stimuli followed by auditory ones was adopted
because the presentation of auditory stimuli had been found to affect the
responses in other modalities but rarely vice-versa.
The next phase was to present a single auditory stimulus (click), or a
single flash, followed one second later by repetitive stimulation in the
other modality. The choice of sequence depended upon the behaviour of
the auditory and visual responses as indicated in the earlier trials. Previous
experiments had shown (Walter, 1963) that the commonest type of inter-
action was for the first (“Conditional”) response to be augmented and
the second (“Unconditional”) one to be attenuated progressively. There-
fore if, as is most common, the auditory responses were larger and more
persistent, the program was arranged to provide series of repetitive clicks
as unconditional stimuli preceded by single flashes as conditional ones.
This sequence was maintained for two or three sets. The subject was then
given a button to press with the left hand; usually this was not connected
to the stimulus-generator for one set of presentations in order to see the
effect of a purely instrumental response. The subject was asked to press
the button as soon as he heard the auditory stimulus (or whichever was
the second modality), With some children and clinical subjects, however,
in order to provide a modest incentive as soon as possible, the button
circuit was connected at once to the stimulus generator so as to terminate
the train auditory stimuli. This provided an effective operant response
whereby the subject could control the environmental situation and be
aware of this control by the abbreviation of the stimulus. The button-
circuit was so arranged that the unconditional stimulus could not be
avoided by anticipation. This was necessary to ensure that the full quota
of responses should be included in every average. After two or more sets
with the paired stimuli and operant response, the conditional stimulus was
withdrawn and a set was averaged with the unconditional stimulus alone.
The conditional stimulus was then restored, and in some cases this pro-
cedure was repeated several times.
The subject was then allowed a brief intermission and the whole
sequence was then gone through again but with repetitive tactile stimula-
tion delivered by a torque motor driving a blunt pin against the index
or middle finger of the right hand. The responses to these stimuli having
been averaged over enough sets to establish the habituation rate, the
tactile stimuli were presented as unconditional stimuli paired with which-
ever modality had previously been unconditional now in the conditional
clause. Again, the operant response (which could now control the tactile
stimuli ) was included, the conditional stimuli were withdrawn and
so forth.
A complete experiment including from 35 to 50 sets, each of twelve
presentations lasting about two minutes, lasted from one to two hours, but
for young children and ill or seriously disturbed patients the program was
shortened to about 30 minutes by omission of some of the associations
without operant control and reduction of the number of repetitive sets.
With combinations of three stimulus modalities and three types of sub-
ject response, 54 realistic situations can be provided, of which only a dozen
or so are included in this standard basic program. Other combinations
were used for special purposes, and these will be described in the appro-
priate context. In experiments intended to ascertain the cerebral criteria
of significance and contingency, supernumerary isolated stimuli in the
appropriate modality were introduced manually between programmed
presentations in order to dilute the conditional probability of association
by a known factor.
In experiments with hypnosis the standard program was presented sev-
eral times over before induction in order to obtain statistically valid
records of normal variations over an adequate time interval, since some
long-term changes can occur as well as shorter-term habituation and con-
tingent effects.
In patients with implanted electrodes an essential part of the thera-
peutic procedure is preliminary electrica1 stimulation of each electrode in
turn; electrodes with a relatively low threshold for a sustained local after-
discharge can be presumed to be in grey matter and are therefore excluded
from the therapeutic polarisation plan. The magnitude and extent of all-
or-none after-discharges at such electrodes can be compared directly with
the dimensions of the evoked responses, thus providing a measure of tissue
involvement and dispersion.
In order to facilitate comparison of the evoked responses during the
various phases of the experiments the average records from each channel
were traced together on a single chart. Similarly, the pulse and respira-
tion rates and PGR latencies were also charted for each subject. In this
way the principal observations from each experiment could be viewed in
a single field.
Renilts
Responses to visual and auditory stimuli can often be detected in con-
ventional EEG records, but their amplitudes are rarely much greater than
the background activity unless the stimuli are novel, startling, or in the
case of flicker, repetitive. The responses to visual stimuli have been inves-
tigated most completely because they are larger, are easily distinguished
from the receptor potentials (ERG) and have great clinical interest in
the precipitation of epileptic seizures (“photic activation”). It has been
assumed rather generally that these visual evoked responses arise in the
primary visual cortex, but comparison of records from the scalp with those
from electrodes implanted in the occipital lobes suggest that this is not
the case.
The responses to single flashes at irregular intervals in primary visual
cortex are stereotyped and persist unchanged over several thousand repeti-
tions while the scalp responses are variable and usually exhibit marked
habituation over a few dozen presentations. The conclusion is that the
responses as seen in scalp records from the occiput are compounded of
specific responses and nonspecific features from visual association areas,
the latter usually predominating.
Similar considerations apply to the interpretation of responses to audi-
tory stimuli, with the added complications that the much smaller primary
areas are nearer the receptor and are also surrounded by large tracts of
nonspecific parietal and temporal cortex. The proximity of the cochlea
to the temporal lobe introduces the possibility of interference by micro-
phonic potentials, and the contribution of the relatively minute strip of
primary auditory cortex is unlikely to be comparable with that of the adja-
cent masses of association and nonspecific tissue.
Specific tactile responses are also difficult to identify by location for the
same reasons; the narrow strip of somato-sensory cortex along the post-
central gyrus is very small compared with the adjacent nonspecific areas.
It was thought at first that the specific and nonspecific categories of
response might be distinguished by their waveforms and latencies but this
is rarely the case. The waveform of the nonspecific responses is extremely
variable, for reasons that will be described later, and the latency of these
responses even as recorded in the depths of orbital frontal cortex is rarely
longer than 25 msec. for auditory and tactile or 30 msec. for visual re-
sponses, figures not very different from those for the major components of
the corresponding responses in specific cortex. The most consistent differ-
ence between responses in specific and nonspecific cortex is in their con-
sistency, a distinction which is difficult to apply experimentally as well as
verbally. As already mentioned the truly specific responses in primary
receiving areas show no signs of habituation over long periods whereas
those in nonspecific regions are typically variable in this sense. Unfortu-
nately a response in nonspecific cortex which has undergone habituation
resembles a primary specific response even more closely than when it was
fresh, so that even this distinction is elusive and confusing. The most
satisfactory principle is to assume that, in experiments with human subjects,
all evoked responses are mainly due to nonspecific activation unless the
contrary can be proved.
An important consequence of this conclusion is that since, by definition,
responses in nonspecific cortex are diffusely projected over wide areas,
their electric fields as detected on the scalp are very extensive. This
means that no conventional electrode arrangement can be expected to
provide a “true” picture of these potential differences; bipolar montages,
whether longitudinal or transverse, inevitably produce the first derivative
of the potential differences with respect to space, no common-reference
system can be assumed free from activity at the reference electrode and
even the more promising Average Reference ( A R ) system is liable to
serious ambiguities if a large proportion of the electrodes are likely to be
involved in the disturbance.
These inferences have been confirmed by experiment. In patients with
intracerebral electrodes thirty or forty regions can be found which are
not involved in sensory responses (mostly in the frontal white matter).
When these are used as a combined AR the involvement of other regions
(including the scalp) can be explored. With this arrangement evoked
responses to auditory, visual, and tactile stimuli have been recorded from
the ears, chin, neck, and, in fact, all parts of the head. A further compli-
cation is that the time-relations of responses in the various regions of the
head are not static; measurable time differences have been found, some
consistent, some varying with the conditions and context of the stimuli.
In some cases the time-differences are such as to suggest a regular “sweep”
of afferent projection from front to back of the brain, rather as described
for the alpha rhythms by Cooper and Mundy Castle ( 1960). This suggests
that the “traffic-control” system outlined in the Introduction may in fact
direct and regulate the manifold diffuse sensory projections in an orderly
but flexible sequence of modulated waves. The systematic nature of this
process can be detected only when the signals are large and regular
enough to be identified clearly since the time-differences are only of the
order of 10 msec. and the averaging procedure, by definition, effaces
small irregularities.
Taking account of these complications it is perhaps surprising that any
useful records can be obtained from the scalp at all, and, indeed, were it
not for the opportunities presented by the intracerebral electrodes for
accurate local recording, the analysis of scalp potentiah would probably
have been regarded merely as a technical demonstration. However, the
peculiar-and still only partly explicable-distribution of electric fields
over the head must be accepted and interpreted as circumstances permit
in terms of intracerebral events, even if these seem surprisingly remote
from the superficial regions of apparently greatest activity.
Another factor to be considered is the incidence and source of artefacts.
These arise mainly from the eyes and scalp musculature, particularly in
association with novel or startling stimuli. Of these sources the eyes are
usually the most troublesome and electrode placements have to be ar-
ranged to minimise the potential differences from this source (which are
mainly antero-posterior in gradient ) as compared with the superficial
projection of the action-fields in nonspecific cortex.
This fieId structure has the effect of developing the highest potential
differences between an electrode at about the vertex and one near the
mastoid processes. With this transverse derivation potential changes due
to eye movements are extremely small. For this reason most records have
been taken with this simple montage, but there is no reason to suppose
that the vertical electrode is really above a particularly active cortical
zone. In fact the tissue immediately below the vertex is merely the sagittal
sinus-a pool of venous blood as remote from active tissue as any part of
the head. It was thought at first that the apparent activity of this region
might be due to a profusion of responses in the cingulate gyrus but in one
case where therapeutic electrodes were introduced into this very region,
remarkably W e response to sensory stimulation could be detected. The
conclusion is that the activity of the vertical region is due to the gradient
irregularities in the general field at this point and that the field itself is
generated by the compound activity of the medial and lateral frontal
cortex, where implanted electrodes do in fact indicate particularly large
and widespread responses.
Another component of the responses in nonspecific cortex is that from
the activation of the frontal orbital cortex. This is sometimes distinctive in
waveform and lability and presumably can contribute little to the fields on
the upper convexity of the scalp. It does, however, spread far into the
basal regions and is probably responsible for most of the activity detected
in such unlikely regions as the chin and neck. The mastoid projections
used-for lack of a better-as a common reference zone show activity from
both superior and inferior regions, but luckily these are often out of phase
so that some degree of cancellation and effective equipotentiality can be
expected. This is not always the case, and since the orbital and superior
components often respond differentially to habituation and conditioning
confusing results can be obtained unless intracerebral references are avail-
able. These effects have been allowed for as much as possible and the
observations described below have all been checked by intracerebral
analysis as well as scalp recording. The following are the principal effects
seen regularly in the population as a whole; the exceptions are in certain
clinical categories and young children.
Dispersive convergence. This apparently self-contradictory term is used
to describe the basic fact that signals in all modalities studied (sight,
hearing, and touch) converge toward frontal cortex but are widely dis-
persed therein (FIGURE 4).All cortical regions studied in these lobes have
shown responses, none has responded in one modality only. This general-
ization applies to the orbital cortex from the midline out to the lateral
border of the anterior fossa, the lateral frontal cortex from the frontal pole
back to the level of the tip of the temporal lobe and to medial cortex over
a similar extent. In many cases electrodes lay in sulci plunging deep into
the frontal lobe from the orbital, medial, or lateral surfaces and in these
too, responses to sensory stimuli were often large and distinctive. There
is no doubt therefore that, in a special but important sense, the prefrontal
cortex is essentially a part of the sensory system.
The size and latency of these responses vary considerably, but the aver-
age peak amplitude is of the order of 10 microvolts as recorded from a
,=
LEM. Average of I 2 Trials

SOUND
Lt.orb.fr. I8
R t orb.fr: k:f.(80 1
FLASH
TOUCH
I
I
1 T2
I
FIGURE4. Dispersive convergence and modality signature in pre-frontal non-
specific cortex. Both electrodes are in orbital cortex, 18 on the left and 80 on the
right. Both show responses in all three modalities, but the pattern and proportions
for each are distinctive. The auditory responses are the eighth average, after over 100
stimuli; the visual and tactile responses are the second averages after 28 stimuli.
single implanted electrode referred to the average of all the other 67. The
amplitude at the vertex is not much different from this and is actually
larger in some people, suggesting that the space-average provided by the
scalp electrode can summate activities in many regions. In some patients
the deep frontal evoked responses can be measured without automatic
averaging because of the low level of background intrinsic activity; these
measurements show considerable variations in size from stimulus to stimu-
lus and these are of course eliminated in the averaging process. Apart
from the habituation and other effects, however, the variations from aver-
age to average (that is when considered in sets of twelve) is much less.
The maximum peak amplitude of individual responses is never greater
than 30 microvolts. The significance of this figure as an indication of the
degree of cortical involvement in a particular modality can be seen when
it is compared with the maximum amplitude of discharge that the same
region of cortex can develop when stimulated electrically. As described
elsewhere (Walter, 1962), using a train of 400 psec. pulses at four to six
volts, cortical regions in the frontal lobes can be stimulated to produce
sustained local after-discharges. These involve only a few cubic millimetres
of the cortex immediately adjacent to the cathode electrode since other
electrodes only a few millimetres away may show no sign of the dis-
turbance. Such after-discharges are usually all-or-none and may last a
minute or more, to cease abruptly with no decrement. Their peak ampli-
tude is of the order of one millivolt and may reach three millivolts under
conditions identical to those during sensory response recording. If it be
assumed that these after-discharges represent the synchronised-and self-
excitatory-activity of the whole local population of neurons and their
processes, the inference is that the sensory evoked responses involve only
about one per cent of these structures.
As will be explained later, there is clear evidence that responses in any
one modality are strictly limited in the number of pathways accessible to
them, while responses in different modalities travel by entirely inde-
pendent routes to the surface of cortex where they mingle only at the
terminus of their line. The conclusion is that a signal from any receptor,
even if due to maximal sensory stimulation, can activate only about one
neuron in a hundred in frontal cortex, but will have this effect over a
vast surface containing several thousand million neurons. All significant
sensory stimuli therefore-with the provisions detailed below regarding
the context of their presentation-must be considered as activating some
tens of millions of neurons over a very extensive area, without necessarily
intruding on any other afferent signals or background activity.
The latency of the responses in these regions depends on modality, since
receptor and tract delays necessarily vary. Responses to auditory stimuli
such as clicks or the onset of tones have an average latency of 25 milli-
seconds (msec.) and those evoked by tactile stimuli to the hand show
about the same delay, Responses to visual stimuli (isolated flashes from
a stroboscope or the onset of a series of flashes-flicker) have latencies of
35 msecs. These figures are about the same as those for the latencies in
primary specific cortex and the inference is that, however they reach their
frontal destination, the pathway must be a fairly direct one. It is difficult
to exclude the possibility that the frontal responses are due to direct relay
from the specific regions by way of the long anterio-posterior commis-
sures. No patients examined so far have provided the test preparation of
total section of the optic radiations, and the delays in such a circuit would
be very short. The marked differences in behavior between the specific
and nonspecific responses indicates that, whatever the projection circuit,
there must be a highly selective element interposed between the provenance
and the destination. The most likely but still not entirely proven sources,
at least for the auditory and visual signals, are of course the nonspecific
relay centres of thalamus with their connections to the specific stations
in the geniculate bodies. The course of the tactile signals is less clear
since even in experimental animals “the sources of the sensory somatic
inflow which activate the various brain stem structures are not yet estab-
lished unequivocally” (Rose and Mountcastle, 1959).
An important feature of responses in nonspecific frontal cortex is their
extreme sensitivity to distraction, boredom, and drowsiness (FIGURE 5 ) .
R. 6.12.61. Averaqe of I 2 Trials
Post orb.med.It.- , b
/
[Flash
r”
~secr j-
51 1_
Flash
I L
DIST RAC TED by CONV E RS AT ION

FIGURE 5. The effect of accidental distraction by conversation on visual responses
in left orbital and right medial frontal nonspecific cortex. The various components are
affected differently by attention to a question about meal-times.
This property is discussed later in relation to more precisely specified con-
ditions but it is essential to appreciate that these phenomena are not direct
reflexions of specific responses, and that they are under the continuous and
vigilant control of some agency in close touch with the whole environ-
mental situation. It is possible, and even likely, that some regions at least
of the frontal nonspecific cortex receive signals both from the specific
primary zones through the long commissures and also from the thalamic
and brain stem diffuse systems, the latter providing general conditional
modulation of the detailed reports transmitted from the former.
ModaZity signature. Although nonspecific in the sense that it receives
signals from many, and possibly all, sensory sources, the pre-frontal cortex
does not respond identically to all modalities. As indicated above the
latencies of responses to visual stimulation are longer than in other
modalities but this is not a distinctive feature since the cortex could
scarcely "know" that the time differences were characteristic. The wave-
forms of responses, however, differ appreciably according to the sensory
modality and maintain some trace of their signature even through a
series of complex experiments in which their components may vary over
a wide range (FIGURE 6 ) . The largest responses are usually those to audi-
tory stimuli; they contain several components or phases, of which the first
surface-positive wave is the most persistent, and the superimposed surface-
negative one the most prominent in novel or significant conditions. The
later components are small and evanescent in auditory responses. Visual
responses are altogether more labile than auditory ones and are usually
more complex with two or more negative components superimposed on
the positive wave, sometimes followed by a very long slow negative surge
peaking at about 200 msecs. The most peculiar features of visual responses
are the coherent after-rhythms seen in some subjects particularly when
the stimuli are significant and conditional (FIGURE 12).These after-rhythms
were observed by Barlow (1960) and Brazier (1960, 1962) using auto-
correlation as well as averaging techniques. They can be seen also in
superimposition records (Walter, 1962) and are remarkable for the con-
stancy of their phase and frequency, the latter being usually within the
alpha range for the subject. In some cases however, the frequency of an
after-rhythm is quite different from any intrinsic component, suggesting
that this effect is not necessarily related to the alpha mechanisms. On the
other hand, subjects who display no alpha rhythms with eyes shut never
seem to develop a coherent after-rhythm even following highly significant
visual stimuli, so there must be some connection between the two effects.
The anatomical distribution of the after-rhythms also indicates a relation
to visual mechanisms, Records from electrodes in primary visual cortex
show the effect in restricted regions, often those less involved in specific
evoked responses, and in some subjects the after-rhythms are more distinct
in records from parietal or vertical scalp electrodes than in any single
intracerebral derivation. This indicates that the effect is due to precise
synchronisation of activity over a wide area rather than to intense rhyth-
micity in any particular zone. The observation that the tendency to pro-
duce coherent after-rhythms can be enhanced by inspiration of CO, and
13.12.61. Averoqe of I 2 Trials
T SOUND
SCALP’
\
_____-
Vertex- Lmastoid
t.occip.-Lmastoid
- ------
-b--
J ,
G?__
frecr a j
F LASH
T OUC H
f 4
r- 1
-
FIGURE 6. Modality signature in scalp records of responses in a normal subject
during early stages of standard experiment. The vertical derivation shows responses
in all three modalities but each has its distinctive pattern.
reduced by voluntary hyperpnoea (Walter, 1962) is further evidence of

some sort of pacemaker, since C0,-like states of attention and alertness-
can be assumed to activate the diffuse brain stem reticular formation.
Both the biochemical and psychological methods of “activation” typically
suppress rather than augment the intrinsic alpha rhythms and their op-
posite action on the coherent after-effects of visual stimulation should
be an important clue to the significance of this effect.
Responses to tactile stimulation are the most variable and elusive of
the three modalities. The variation between individuals is particularly
striking; in some normal subjects the tactile responses are as large as in
the other modalities, in others they are scarcely visible. The largest
responses of all in this modality have been found in some congenitally
blind children and there was some correlation between the extent of these
responses and proficiency at Braille. In several normal subjects also, the
presence of unusually large, extensive and persistent tactile responses
seemed to be associated with a high sensitivity to texture (Walter, Cohen,
Cooper, and Winter, 1963).
The waveform of tactile responses contains the same components as
the other modalities but the late negative swing is often more pronounced
and seems to spread more widely over the scalp than the earlier features.
In general, the delayed negative surges are much less prominent in
intracerebral recordings than in those from scalp electrodes, suggesting
that these components, rather as in the case of the coherent after-rhythms,
are signs of widespread superficial cortical change at a low level. Simi-
larly also they tend to be augmented by novelty and significance and to
disappear completely when the stimulus is an unconditional one preceded
by a conditional signal in another modality.
Zdiodromic projection. This term is used to describe the fact that signals
in the various modalities evoke responses that travel by “private lines”
to the surface of the nonspecific cortex. The proof of this is simple. If two
stimuli are presented in the same modality at decreasing intervals, an
interval is reached at which the second response dwindles and finally
disappears. This is usually at a stimulus interval of 100 to 200 msec. This
may be described as intra-modality occlusion, and indicates that the struc-
tures involved in the first response are refractory or incapable of respond-
ing to the second stimulus for a considerable period, and that no “spare
lines” are available. The first response, of course, remains unchanged.
Now, if the second stimulus is in another modality this effect is not seen;
instead, as the interval between the stimuli is diminished the responses
superimpose algebraically, producing a different waveform derived exactly
from the sums and differences of the two responses (FIGURES 7 and 8 ) .
It is possible to arrange an interval between say a flash and click such that
the responses cancel almost entirely, as though the second response were
influencing its predecessor, which is absurd. The effect can be obtained
both with intracerebral and with scalp recordings and with any combina-
tions of modalities. It indicates that the structures involved in each
modality are separate and independent right up to the terminus of the
network at the surface of the nonspecific cortex. This effect illustrates a
very important technical point; absence or suppression of electrical change
in a recording does not necessarily indicate electrical inactivity. Instru-
mental synthesis is undoubtedly the origin of many “typical” EEG wave-
forms-instrumental cancellation may similarly be responsible for effects
described as “suppression,” “activation patterns” and the like.
Idiodromic projection to nonspecific cortex has an important bearing on
the physiological interpretation of the interactive effects described below.
The dependence of afferent pathways in the several modalities suggests
E L 1 3 - A R ?H Q‘lif62
A
FLNH
n
1 DO
-sot
FIGURE 7. Idiodromic projection demonstrated by presentation of a flash stimulus
followed by a click at various intervals. Derivation from an electrode in left orbital
cortex referred to intracerebral AR. Trace 1 shows response to click alone, Trace 2 to
flash alone, Trace 3 the response to a flash followed by a click 100 msec. later. The
two responses cancel except for the first component of the flash response and the last
of the click response. Traces 4 and 5 with stimuli at 80 and 30 msec. separation,
show progressive instrumental summation of the two responses as the components
coincide.
that the interaction between their effects is mainly at the superficial corti-
cal level even if the preliminary selection processes are pre- or sub-cortical.
A corollary of this supposition seems to be that the selection (on the
basis of conditional significance) must be mediated by cortico-fugal con-
trol of the sub-cortical relays. This hypothesis leads to the concept of
retroactive cortico-basal circuits in which the maintenance of stability
would be a major problem. Failure to maintain stability could result in
cumulative oscillations; the notorious susceptibility of many epileptics
and some normal subjects to “activation” by rhythmic sensory stimulation
may well be evidence of this inevitable weakness. The waveforms of the
normal evoked responses may be described as micro-wave-and-spike and
the sustained after-discharges evoked by local electrical stimulation of
normal cortex have a similar appearance at a higher amplitude; the
difference between these and the enormous potentials of the petit mal
wave-and-spike may be only a matter of degree, and dependent on the
extent of synchronisation and cortical permeation. The contribution of
metabolic factors should not be overlooked in developing these notions:
hypersynchrony can often be potentiated by mild alkalosis or hypocapnia
(as in voluntary hyperpnoea) and this has a number of known effects on
5
c
9 --
10
11
FIGURE 8. Idiodromic projection in scalp recording from vertex to intracerebral

AR. Same subject as in FIGURE 7 . Trace 1: click response alone; Trace 2: flash alone;
Trace 3: with 270 msec. between stimuli both responses appear complete, including
the secondary negative surge of the click response. Traces 4, 5, and 6: at intervals
between 130 and 70 msec. the primary components cancel almost completely but the
secondary negative surge remains. Traces 7 and 8: at shorter intervals the responses
summate. Traces 9, 10, 11: with two stimuli in the same modality. There is no can-
cellation since both responses occupy the same pathway.
oxygen availability in the brain and on the brain stem reticular system.
This is really another problem, but it can intrude on experiments of the
type described here since some subjects overbreathe automatically during
stress or excitement and thus modify both the intrinsic background and
interactive stability of the non-specific responses.
Diferentiation. Most of the earlier reports on sensory evoked responses
in man dealt with the effects of sustained rhythmic visual stimuli, for the
excellent reasons that these are the only effects that ca.n be detected in
conventional records, and they also have valuable clinical applications. In
the experiments described here both single, sustained, and repetitive
stimuli were used. In general, and with a few notable exceptions, the
responses evoked by single, sustained or repetitive stimuli in the same
modality are indistinguishable in non specific cortex. The responding
system behaves as if it were coupled to the receptor through a differentiat-
ing network, much as the stages of a conventional amplifier are coupled
by differentiating capacitors and resistors, with a short time-constant. In
physiological terms, the nonspecific responses are “on-effects.” The effect
of this is, of course, that the output of the system is informed about the
onset of the stimulus at the input but not about its absolute level or con-
tinuance. In a linear differentiating system there should be an equal and
opposite off-effect, and this has been observed in some subjects for visual
and auditory stimuli but not for tactile ones. There is, of course, no reason
to suppose that the cerebral systems are linear, and furthermore the re-
ceptors themselves contain a wide variety of elements with “on,” “off and
“on-off properties.
The theoretical importance of differentiation is considerable, however
it be achieved. Many years before experiments of this sort were possible
it was suggested (Walter, 1953) that an essential preliminary operation in
the establishment of conditional associations in the brain was the differ-
entiation of all signals before projection to nonspecific regions “for
information only.” This was considered necessary because for legitimate
and economical association, only the onset or initiation of an uncondi-
tional stimulus is important. In the working model of this hypothesis
(“CORA”) differentiation is in fact an indispensable part of the prelimi-
nary processing; if information about the steady or absolute level of all
possible stimuli were transferred to the later stages of the system, these
would soon be jammed with irrelevant and useless stores of obsolete data.
Recognition of the theoretical need for a process does not of course supply
an explanation of its mechanism and many experiments remain to be done.
One of the most promising is reciprocal or negative stimulation, using
interruption of steady stimuli such as “dark flashes” or “clicks of silence.”
The results of these experiments are still under review-they are com-
plicated by the unavoidable quality-inversion; a disagreeable sound which
a subject quickly learns to turn off cannot be used as a continuous stimulus
with intervals which the subject must learn to shorten. The interruption
must be made disagreeable and the stimulus correspondingly attractive.
In a few subjects the differentiation effect is much less marked in the
early phases when isolated stimuli are presented. The two in whom a
repetitive nonspecific response was seen to the visual flicker were both
very intelligent young students and the rhythmic responses in the non-
specific regions dwindled after three or four dozen presentations. In one
of these subjects (SMP) there was no on-effect, the flicker evoked a
following response in both vertex and occiput and there was a large off-
response only in the vertex ( FIGURE 9 ) . The time-relations of the following
responses in the two regions showed a steady drift during the 1.3 seconds
of exposure; at first the waves appeared out of phase between vertex and
occiput (with a common reference on the mastoid) while at the end of
the exposure they were precisely in phase. These relations were main-
tained through four sets of 12 presentations each, indicating a very high
degree of stability and coherence. The most likely interpretation of this
finding is that during the first half-second of the exposure the anterior
nonspecific responses were leading the posterior ones by about 30 msec.;
this time difference changed during the last few hundred msec. until for
the last four or five flashes the responses were synchronised in all regions.
Rare though such an appearance is, it demonstrates that nonspecific
regions can receive detailed information about the course of a stimulus
as well as about its onset, and that such broadcast details are subject to
systematic time- and space-manipulation. Apparently correlated with these
effects were the subjective reports by these two subjects; both said that
at first they thought that the flicker and the visual illusions it produced
might be important since they had read accounts of them. Later they
regarded the visual stimuli as signals only, to be controlled, or as warnings
for other actions. In accord with this change in attitude the responses
during conditioning lost their rhythmic character and developed instead a
typical on-effect. The degree of differentiation must therefore be under
the control of some mechanism related to “attitude,” rather as the time-
constant of an amplifier can be regulated according to the phenomenon
to be recorded. As in the case of the other labile components, such a
control system, operated in effect by statistical appraisal of associated
events, implies a cortico-fugal channel with the potential weaknesses
already referred to.
Habituation. This term has been adopted in neurophysiological usage
to describe the progressive attenuation of certain cerebral responses to
monotonous regular stimuli. It is not the happiest word since in English
a habit is something a person cannot help doing: it is better to think of
the French “s’habituer”-to get used to something. In many early reports
on habituation in allegedly specific areas the effect was attributed to
central regulation of receptor or afferent-pathway transmission. This
explanation now seems less plausible since in many cases habituation
seems to have been due to indirect action on the pupil of the eye or the
intrinsic ear muscles, A further possibility, particularly in human studies,
is that the responses considered as specific contained components from
nonspecific regions. When recordings have been made from electrodes
implanted in human visual cortex no sign of habituation has been de-
tected; the complex local responses are identical over several thousand
sets of stimuli. In the same subjects, however, scalp records from the
577 jsc )Lv : * a 3 -

0
I sc6
FIGURE 9. Anomalous differentiation of visual responses in normal subject A.

Vertex to mastoid. B . Occiput to mastoid derivations. Trace 1: first presentation of
flicker at usual recording speed showing desponse at 15/sec. Traces 2, 3, and 4:
responses to flicker recorded at higher speed to show consistency of following re-
sponse, lack of .on-effect and marked off-effect only at vertex. During these presenta-
tions the subject was concentrating on the visual illusions evoked by the flicker.
Trace 5: repetitive clicks at 15 per sec. evoke a typical on-effect, just visible follow-
ing in vertical derivation and no off-effect. Traces 6 to 16: Association of conditional
click with flicker and operant response emphasises the off-effect and finally evokes an
on-effect in the occipital derivation.
571 367 I src

FIGURE
9. (Continued)
occipital region as near to the visual cortex as possible, show considerable

variation particularly of the surface-negative components ( SNC ) , After
a few dozen stimuli the only feature left is often the single surface-
positive wave. The so-called visual responses in man are therefore
mixtures of specific and nonspecific components and the same applies to
responses in other modalities.
Habituation is a prominent characteristic of nearly all responses in non-
specific cortex (FIGURE 10). The rate and extent vary over a very wide
range, however, even in normal subjects, and if clinical states are included
!
5
10
II
12
FIGURE 10. Habituation of resuonses to auditory stimuli in orbital cortex (same
subject as FIGURE 4 ) . A. 12 primary records to show degree of masking of responses
by intrinsic activity. B. Average of these first twelve presentations showing clear re-
sponse, followed by second, fourth, seventh and ninth averages, each of twelve pres-
entations. Habituation is clearly demonstrated after about 50 stimuli at the intervals
shown in FIGURE 3. The first effect is attenuation of the SNC.
Walter: Auditory and Tactile Responses
AVERAGE
I - 12
13-24
3 7-48
73-84
97-108
3--- 2 5ec.-
FIGURE
10. (Continued)
the diversity is even greater. In one patient in a state of manic excitement

the effect was actually reversed; responses to monotonous stimuli increased
progressively over the first few dozen presentations. This inversion of the
usual order is so rare and peculiar that we have dubbed it “Noitautibah
for lack of a more explanatory term. Whatever its relation to clinical
states, the fact that habituation can be reversed at all indicates that it is
not an invariable rule and, like so many other cerebral effects, is contin-
gent on the context of the stimuli and the attitude of the subject.
The most important neuro-electric aspect of habituation is that the
various components of the responses are affected differentially. The main
components, as described earlier, are a surface positive wave and super-
imposed surface negative components; it is these latter that are first
affected by habituation. In some subjects the positive wave persists over
a hundred or more presentations and in pathological cases with cortical
degeneration this may be the only detectable component. The interpreta-
tion of these differences depends on the assumptions made about the sig-
nificance of electric sign in relation to cortical elements. The most plausible
explanation is that the surface positive components indicate activation
of neuronic somata in the deeper cortical levels, while the negative ones
are signs of graded depolarisation in the apical dendritic feltwork. This
interpretation is corroborated by study of the phase relations of responses
within the nonspecific cortex; when one electrode is on the scalp or cortical
surface and another just below its border, the main components of the
responses are invariably out of phase (with both electrodes referred to
AR) (FIGURE 11).This can only mean that the effective generator is a
polarised sheet of tissue with laminar poles about three mm. apart. On
these assumptions, habituation would seem to influence mainly the diffuse
----A *i----- -
1
CLICU --- ----I sac -+-
1
FlAsH
FIGURE 11. Comparison of phase relations of responses to flicker and clicks in
cortical, subcortical and scalp derivations. A . Responses in two adjacent electrodes
23 and 24 placed as in diagram their contact surfaces three mm. apart, both referred t o
AR. The phase inversion is almost perfect. B . With a shorter interval between the
start of flicker and the click, algebraic summation is seen in both regions, the sec-
ondary response in cortex remains uncancelled. C . Simultaneous averaging of scalp
( vertex) and intracerebral derivations, both referred to intracerebral AR shows
phase inversion, a larger primary SNC and secondary negative surge in scalp deriva-
tions.
radiation of activity across the surface of cortex. Further consideration

of these mechanisms raises again the question of the provenance of the
responses in nonspecific prefrontal cortex. The identification of two com-
ponents, differentially susceptible to habituation ( and as will be described
later, to conditioning) presents the possibility that there are two sources
and these could well be the diffuse sub-cortical projections on the one
hand and the long commissural pathways from the specific primary zones
on the other. This explanation would be helpful also in accounting for the
typical differentiation or on-effect with the occasional appearance of sus-
tained responses; the diffuse projections from the sub-cortical relays would
act as an alerting or warning signal: “something has happened,” and the
secondary projections from specific areas as detailed messages : “This has
happened.” As the novelty and importance of detail wanes with monot-
onous repetition the latter components would dwindle, leaving only the
general nonspecific alerting which in turn would fade as the absence of
significant implication was established.
The possibility that nonspecific habituation might include effects due
to receptor shuttering cannot always be ignored, but it is extremely
remote. In the first place specific responses have shown no such effects in
these conditions and an even more striking proof is that very slight
changes in the quality, context or intensity of the stimulus can abolish
habituation instantly. The contingent restoration of habituated responses
by significant association will be considered later under that heading:
the easy reversibility of the process is an argument against adoption of
the Pavlovian expression “Extinction of the Orienting Response,” for
these responses are by no means extinct-they are liable to erupt on the
slightest provocation.
The time-course of habituation in any particular person is a complex
function of real time, stimulus abundance, stimulus interval, and regu-
larity. Trite though they may be, the distinctions between this effect and
the superficially similar ones of “adaptation” as applied to sense organs
and “accommodation” as applied to nerve fibres should never be forgotten.
In both these latter the decline of activity in response to a sustained
stimulus is a fair approximation to an exponential curve and indeed the
simple algebraic expressions used for physical systems fit the results of
physiological experiments quite well. Habituation, on the other hand,
rarely follows a smooth course and the most common form in normal sub-
jects is for a sustained response level to fall rather abruptly to a low level
after 20 to 50 presentatians at irregular intervals over about 10 minutes.
The impression is rather that the cerebral attention is maintained until
some threshold of triviality is reached. In the clinical subjects who dis-
played little or long-delayed habituation, their mental state could be
summed up by the plaint: “nothing is too trivial to worry about!”
Habituation has sometimes been compared with fatigue, but the differ-
ence between the two is fundamental; the larger the disturbance the
more rapid and complete the development of fatigue, whereas habitua-
tion is faster to smaller stimuli.
From the technical standpoint habituation is a nuisance since it illus-
trates all too clearly the uncertainty principle of cerebral action and
makes the prolonged or repeated study of isolated responses impossible,
except in those special cases, usually with psychiatric anomalies, in which
the effect is minimal. The temptation to use such subjects for experi-
mental studies is hard to resist, since they can provide invariant records
for several hours at a time, involving several hundred stimuli, and offer
incomparable opportunities for investigating the action upon these re-
sponses of other variables. Information obtained in this way has been
included here, with the reservation that the psycho-physiological con-
ditions were certainly abnormal.
Contingent amplification. This term introduces the basic concept that
the extent of responses in nonspecific cortex varies according to the con-
tingency in which they are presented. The phenomenon of habituation
is really a special case of this; a novel stimulus presented repeatedly
without association may be considered as possessing at first an indefinitely
high potential significance, which diminishes steadily with every repeti-
tion until it reaches a negligible level. In these experiments all the stimuli
were novel, for none of the subjects had previously lain on a bed while
occasional bright flashes, clicks in earphones, or mechanical touches on
their fingers were administered apparently without order or reason. In
most normal subjects between 20 and 100 repetitions were enough to
establish that there was no significant relation between the stimuli and
any other events, and the responses were then “habituated.”
At this stage in the experiments, stimuli in two modalities were pre-
sented, the first a single flash, click or touch, the second a series of clicks,
touches, or flashes one second later. Even when the responses to the
stimuli in the first modality had habituated completely, this simple
association provided a high enough significance to restore them at least
to their original level (FIGURE 12). In some subjects, particularly the
younger ones, the SNC of the conditional responses were amplified con-
siderably beyond their size during the first presentation of unassociated
stimuli, and this effect was usually maintained over 25 to 50 trials.
The next stage was to allow the subjects to press a button at the presen-
tation of the second, “Unconditional,” stimulus. This usually amplified
still further the responses to the first Conditional stimuli. When, again,
the button circuit was connected so as to terminate the Unconditional
stimulus, this addition of an operant defensive element enhanced and
consolidated yet again the responses to the Conditional warning stimulus.
Elaborate and tenuous though this interaction may seem, it is one of
the most consistent and predictable of all the effects observed in these
experiments. The delicacy of the relation between conditional responses
on the one hand and unconditional stimuli and operant responses on the
other is vividly illustrated in experiments with hypnotic suggestion. As
is well known and generally agreed, hypnosis is not associated with any
particular changes in the EEG; when subjects in deep hypnotic trance
are told that real stimuli cannot be perceived, evoked responses still
appear as usual. However if they are told that the Unconditional stimuli
are going to be particularly powerful and abundant the nonspecific re-
sponses to the Conditional stimuli are often further amplified, even when
in fact the number of unconditional stimuli is precisely the same as
before. The imaginary stimuli can be as effective as the real ones. The
reverse effect in hypnosis, to be described later, is even more consistent
and convincing, but the evidence of true psychophysiological interaction
in these conditions is ineluctable.
t
S.T.W.
A
, Y-n.
J;T;LI I
FIGURE 12. Development of contingent effects in normal subject. A. First average

of responses to flicker alone showing differentiation and flicker following at 15/sec.
in vertex (compare with FIGURE 9). B. First average of responses to clicks at 15 per
sec. C. First average of responses to touch also at 15 per sec. All responses show
typical waveform including secondary negative surges but no significant coherent
after-rhythms. D. Second average of association between flash and repetitive touch
with operant response ( T R . . . C ) . The visual response, which had previously
habituated, is returning, there is a marked coherent after-rhythm in the occipital
channel and the reaction time is about 150 msecs. E. Withdrawal of the conditional
visual stimulus restores the tactile response to its original size and waveform (com-
pare with C ) and there is no coherent rhythm. The reaction time is 200 msecs. F.
Restoration of the visual stimulus evokes an amplified response, with a small second-
ary negative wave, attenuates the tactile response and the coherent after-rhythm re-
appears in the occipital channel. The tactile response is not attenuated as much as
in D because the contingency is lower after the withdrawal of the visual stimulus
in the previous set ( E ) . The reaction time is again reduced to 150 msec.
Contingent attenuation. As a corollary of the contingent amplification
of the Conditional responses to an associated pair of stimuli, the Uncon-
ditional responses are steadily and regularIy attenuated as the significance
of the implication accumulates (FIGURE 12). Here again, the first effects
are on the SNC of these responses but in manv subjects the whole uncon-
ditional response disappears completely after 30 to 50 presentations. This
I ~
F
- - V
T R ...L
r - - -
I %/+---
2 F k, src*.7 c -
FIGURE
12. (Continued)
effect makes the investigation of response interaction details very diffi-

cult since the effects of the second stimulus may have disappeared before
the desired results have been obtained. This was particularly tiresome in
the experiments on idiodromic projections, since the paired responses had
to be presented together many times at various separation intervals in
order to acquire convincing records. The relatively slow development
of contingent attenuation in some clinical states, notably chronic anxiety,
was exploited for the first studies, which were later checked by fewer
presentations at critical intervals in normal subjects.
In general these reciprocal relations between conditional and uncondi-
tional effects are signs of prompt, effective, and economical adaptive be-
haviour. In mentally disturbed patients, both children and adult, the
contingent adjustments are slow and may fail to develop even after
several hundred presentations. The clinical implications and applications
are another problem, but there seems to be a connection between states
of tension-anxiety, chronic depression, cyclic variations in mentality, per-
haps other less defined conditions, and the contingent interaction of
responses in this simple situation.
A very important extension of these studies would be to vary the time-
separation of the two categories of stimulus. Over the range 0.5 to 2
seconds there are few substantial differences but measurement of the
maximum time-interval over which these effects can be sustained would
be of great interest in relation to personal disposition and mood vari-
ations. In the school of Pavlov, lengthening the time-lapse between con-
ditional and unconditional stimuli proved particularly effective in type-
identification, experimental neurotogenesis and irradiation of inhibition.
The interest of this procedure in clinical application could be very great
since many conditions seem to involve disturbance of time-sense, par-
ticularly when this involves a stoical attitude to delay, isolation or neglect.
In relation to the effect of conditional or unconditional stimuli it should
be mentioned here that in spite of special search and detailed record
scrutiny, no trace has been seen of a “conditioned brain response, that
is a consistent waveform at the moment when an unconditional response
would have appeared when the unconditional stimulus has actually been
withdrawn. On the other hand Conditioned motor and autonomic re-
sponses are commonplace in these conditions, so the classical association
was undoubtedly present. This failure to observe an effect even when it
was particularly sought for (as most of the others were not) does not
exclude the possibility by any means. The standard averaging technique
demands presentation of at least six pairs of stimuli and when the condi-
tional stimulus is unreinforced it may well be that only the first few trials
would show signs of cerebral conditioning in the above sense, and on
the average these would be effaced by the absence of the effect in the
later ones. Averages taken of first presentations only in these conditions
have shown no sign of conditioned responses in the brain. Other phe-
nomena such as contingent alpha suppression, variance attenuation and
so forth are often seen and would have to be allowed for in appraising
records in such conditions.
Unconditional restoration. When the responses to an unconditional
stimulus have undergone complete contingent attenuation, withdrawal
of the conditional stimulus invariably restores them, at least to their
original amplitude (FIGURE 12). The effect on the SNC is particularly
striking since these are the first to fade; even when the negative com-
ponents were quite small in isolated responses and entirely suppressed in
unconditional ones they may become the most prominent feature after
withdrawal of the preceding conditional stimulus.
When records of contingent attenuation and unconditional restoration
are examined without reference to other phases of the experiment the
impression given is of a simple occlusion by the first, conditional, re-
sponse. That the effect is far from simple is shown by the comparison of
earlier phases when, during the first associations both conditional and
unconditional responses appeared together. An even more dramatic proof
of the subtlety of the effect is given by diluting the significance of the
association. This can be explained in several ways; the simplest is to start
by considering the system as a computer of conditional probability. In
the experiments as described so far the subject is first provided with in-
formation suggesting that neither visual nor auditory nor tactile stimuli
are of any importance. The probability of their significance in relation
to other events, indefinite at the outset, sinks steadily toward zero as
more and more signals are received with no presage or sequel. The situ-
ation is then radically altered; two signals are presented in sequence with
an invariable association or syntax-“IF flash, THEN clicks, so, press
button!” Again, as more and more such events are experienced the prob-
ability of the association being significant grows till after a score or SO of
presentations the odds against association being accidental are well over
a million to one. Concurrently with this sequential analysis the evoked
responses go through the evolutions already described.
Now, when the conditional stimulus is suddenly withdrawn the condi-
tional clause in the proposition above vanishes, leaving only the indica-
tive statement and the imperative. From here on the probability can never
reach quite the proximity to certitude that it attained before. For example
there may first have been 24 presentations in which all, 24/24, were
unequivocal associations. When 12 unconditional stimuli are then given
without conditional warnings, only 24 out of 36 experiences have been
associated, and even if the next set of twelve are again all associations
there are still only 36/48 linked experiences, giving an overall probability
of 0.75 instead of 1.0. In confirmation of this, the contingent attenuation
is much less marked in unconditional responses during the set of associ-
ations immediately following the one to test withdrawal of the conditional
stimuli.
This observation suggested that the criteria of significance adopted b y
the brain could be assessed by providing a suitable number of super-
numerary conditional stimuli, without unconditional reinforcement, be-
tween associations. In this way the significance of the association can be
diluted by any desired factor; for example if 24 clicks are presented, of
which only 12 are followed by flashes to be extinguished by pressing the
button, the objective probability of the clicks implying flashes is just 0.5.
This inference, however, is only true if there have never been any clicks
or flashes alone or together before, that is if the 24 clicks are the whole
population of clicks, not just an arbitrary sample of a larger population.
This is not the case of course and it is still uncertain to what extent the
subjects extract a trailing or running average, and there is certainly great
variation between individuals. The basic effect, however, is quite clear
and invariable; the more unreinforced stimuli in the conditional mode,
the larger are the SNC of the unconditional responses (FIGURE 13). In
5TH l $ l b 3 W
o-n
- d *+
c
I
c
r
FIGURE13. The effects of significance dilution or equivocation on conditional
responses. Same subject as FIGURE 12. A. Standard first association of flash with
clicks and operant response, showing contingent amplification, attenuation, secondary
negative surge and coherent after-rhythm. B. The same after 12 further presentations.
The association is firmly established, the negative surge is increased, coherent rhythm
amplified, auditory response attenuated and reaction time reduced to 50 msec. C .
Withdrawal of visual stimulus restores auditory response, abolishes coherent rhythm
and lengthens reaction time to 160 msec. D. Restoration of visual stimulus re-estab-
lishes situation as in B, but with less after-rhythm and longer reaction time. E . Pres-
entation of 24 flashes of which only 12 were followed by clicks dilutes probability of
association within set to 0.5; primary visual responses remain but secondary wave
and coherent rhythm are attenuated while contingent attenuation of auditory response
is abolished, as though the conditional stimuli had been withdrawn as in C. Reaction
time is over 200 msecs. F. Unequivocal association of 12 flashes and clicks in next
set increases secondary negative surge and contingent attenuation, but coherent
rhythm is absent and reaction time is still long; the significance can never be quite
the same again. G. After 46 unequivocal associations the coherent rhythm begins to
re-appear and reaction time is shorter. H . Repetition of significance dilution to 12
associations in 43 flashes again restores auditory response, abolishes secondary nega-
tive surges and coherent rhythm and lengthens reaction time. I . Presentation of flash
only at end of experiment without auditory stimulus shows lapse of contingent am-
plification and total lack of coherent rhythm in absence of association.
I ul11bJoa
1
cr-.. 2d>&
- 1 f-
F c
FIGURE
13. (Continued)
the normal subjects studied so far the effect of significance dilution or

equivocation is complete at a probability level of 0.5, as in the example
above, but the interpolation of a "run" of unequivocal associations can
strengthen the subjective significance rather more than is objectively
justifiable, at least in the more sanguine subjects (FIGURE 14). Perhaps
the most surprising-and neurophysiologically most significant-effect of
probability dilution, is that the responses to the conditional stimuli, even
when highly diluted by abundant identical unassociated ones, persist
unchanged; they may even seem to be slightly enhanced or at least clari-
fied. This last effect is sometimes due to a reduction of intrinsic activity
by the profusion of stimuli and by the subjective excitement expressed
by several subjects when they realise that they are involved in a guessing
or gambling game.
Another very striking effect of probability dilution or contingency
attenuation is the suppression of coherent after-rhythms. As already men-
--Wr “%b
C
H
-4 v-0 ,
-,.------!L1I-
4
- 1
F12/$) t
r
I
- o - m i i
- 1
1 -r
F
FIGURE
13. (Continued)
tioned, in subjects who display this phenomenon, the amplitude and

persistence of the coherent rhythms following conditional visual stimuli
grow steadily as the significance of association mounts toward certainty.
When the conditional probability of the visual stimuli is diluted by pres-
entation of supernumerary flashes, even though the visual responses
proper persist, the after-rhythms disappear. After a series of such experi-
ences these rhythms take a long time to recover and may not reach their
highest level for several days after the equivocal experience. These
rhythms are seen mainly in the parieto-occipital regions, but the slow
secondary negative waves which occupy the same sort of fields as the
primary components in nonspecific cortex follow a very similar course
during variations of contingency: they are prominent in phases of novelty
or near-certainty and diminish when the association is attenuated. The
slow secondary waves seem to return more promptly than the after-
! A
Av. No
c II)
I
i
1 2
FIGURE 14. Comparison of equivocal association and withdrawal of conditional

stimulus in another subject. The tracings in A are from eight sets of twelve presen-
tations of a click followed by flashes with operant response during various phases of
the experiment. The numbers on the left of each tracing indicate the order in se-
quence. In B are tracings from six sets with decreasing degrees of significance dilu-
tion, No. 7 with 12 clicks out of 23 associated. This was followed by 8 in A with
unequivocal association then by 9 with 12/18 associations. The effect on the visual
response is less dramatic here and in 16 than in 7 because the subject felt reassured
by the interposed unequivocal presentations. Tracings 10, 11 & 17 with single prob-
abilities below 0.5 show visual responses as large and complex as those in C , below,
when no conditional stimuli were presented. The conditional responses persist even
when their significance is diluted.
rhythms when the association is re-established and thus resemble more,

though inversely, the behaviour of the' unconditional responses.
The spatial congruence and inverse relation of the conditional slow
secondary negative waves and nonspecific unconditional responses sug-
gest that the contingent attenuation of the latter may be physically re-
lated to prolonged widespread depolarisation of the superficial dendritic
feltwork, of which the negative surges are the outward sign. This mech-
anism of interaction has been invoked to account for other similar phe-
nomena and may be another example of the technical limitations of
electro-biological methods. The apparent attenuation of unconditional
responses following significant conditional ones, if due to previous slow
depolarisation of the active structures, could mean simply that their re-
sponses are progressively potentiated rather than suppressed, and this
would seem a logically attractive explanation. A further advantage of
this interpretation is that it includes a mechanism for the acceleration
and even anticipation of the motor responses evoked by the unconditional
Walter : Auditory and Tactile Responses 355
A I
FLASHES
I FUSHES
14. (Continued)
FIGURE
stimulus. Activity of the operant muscles was recorded in the experiments,

as well as the effect of the defensive action; although this is really another
subject, the shortening of the reaction time with the conditional warning
usually ran parallel with the contingent changes in the nonspecific re-
sponses. In some subjects, the preparatory rise in muscle tone, following
the conditional but preceding the unconditional stimulus and operant
response, developed and subsided with the secondary negative wave.
The reaction time varied in the same way but very short reaction times
(that is responses in effect to the time-lapse after the conditional stimulus
rather than to the unconditional stimulus itself) were often more closely
associated with the appearance of coherent after-rhythms, suggesting
that these may have a timing as well as a storage function.
Hypnotic restomtion. This effect is really a special case of uncondi-
tional restoration, but it is particularly interesting since there is otherwise
so little reflection in the neurophysiological variables of the often spec-
tacular changes in behavior and memory induced by hypnotic suggestion.
Most of these changes may be explained away as purely histrionic, but
certain features, particularly the spontaneous amnesia, are hard to €3
into this category.
In these experiments several attempts were made to induce changes
in responsiveness by suggestions of regression, sleep, and negative hal-
lucination. In one subject the last named, suggestion of deafness to con-
ditional clicks, had a marginal effect on the degree of contingent
attenuation. The most consistent and repeatable results were obtained by
suggestion of profuse stimuli in the conditional mode without reinforce-
ment. For example when the subject had gone through the usual routine
presentations in a normal state, had been hypnotised, and again com-
pleted the standard series, with clicks as conditional and flicker as
unconditional stimuli, she was told that she would hear frequent clicks
all the time and an example was given with real clicks. The normal set
of twelve associations was then presented, with no real extra clicks, and
this was repeated five times. The effect on the unconditional responses
was as if the conditional stimuli had been withdrawn or diluted by real
unreinforced stimuli; the SNC of the visual responses reappeared (FIGURE
15). The operant responses of the subject, as indicated both by the elec-
tromyogram and the reaction time to the visual stimuli, also changed as
f-
~~
sa$ 1qlJK 6 L (D)
FIGURE15. Dilution of conditional probability by suggesting under hypnosis.

After establishment of standard association of clicks with flashes and operant response
under hypnosis (1-5) the SNC of the visual responses was attenuated. The subject
was told she would hear frequent random clicks. There was progressive re-appear-
ance of unconditional visual responses and diminution of the auditory responses.
(6-10). This effect was abolished immediately by cancelling the suggestion. ( 12-13).
if the conditional stimuli had disappeared or lost their significance. The
psycho-galvanic responses (PGR) in this subject also indicated a changed
attitude to the situation. She had participated in so many experiments
before that in the usual conditions all autonomic effects had long since
habituated, but when the suggestion of supernumerary unreinforced clicks
was made, frequent large PGRs appeared again, some between presenta-
tions and some following the “real” conditional stimuli. Reappearance
of autonomic reactivity was seen also in other subjects presented with
equivocal stimuli after a long run of associations without exceptions, As
in this latter situation of real equivocation, the nonspecific responses to
the real conditional stimuli persisted, even when their significance had
been so diluted by imaginary random stimuli that they no longer affected
the unconditional responses.
During the phase of direct suggestion of random clicks under hypnosis,
a set of visual stimuli was presented with no real clicks, that is, without
conditional stimuli. Unexpectedly the responses to the visual stimuli did
not show either the usual unconditional restoration or hypnotic restora-
tion. A tempting inference is that the imaginary auditory stimuli were
more significant or distracting, though presumably random, when experi-
enced alone than when mixed with real clicks. Distraction is probably
the better term and it is easy to understand that an imaginary or illusory
experience could be more compelling and engaging without sensory
stimulation. A rather similar state is reported by some subjects under the
influence of hallucinogens, who resent external influences and prefer to
contemplate their inner experiences. The hypnotic subjects cannot com-
ment on this since they display the typical amnesia for events during
deep trance.
Discussion
The properties of human cerebral responses to sensory stimuli as de-
scribed above seem so complex and varied as to defy coherent explana-
tion. Even if the variations between individuals could be ignored the
diversity of interactive patterns would be difficult to classify, but personal
differences must be expected and accepted as a part of the problem.
Actually it is the personal characters that provide an essential clue to
resolution of these difficulties since they provide apparent exceptions that
literally prove the rules in the sense of testing their validity.
One source of confusion must be considered before discussing the
mechanisms underlying these effects. Reports from different centres vary
considerably both in their nomenclature and in their content; there is at
present neither an agreed terminology nor a conventional or standardised
experimental situation. The range of possible and reasonable situations
is bewilderingly wide. Given three stimulus modalities ( vision, hearing
and touch) and three modes of subject response (none, instrumental,
operant) there would seem to be 54 distinct situations, any or all of
which would be reasonable for human subjects and these can be com-
bined and permuted in an indefinitely large number of patterns. The
“simplest” situation, where a single stimulus is presented repeatedly with-
out possibility of an effector response, is the commonest but is really far
from simple, since it is presenting the brain with a peculiar challenge,
a novel experience of unknown significance which must be appraised
statistically merely on the basis of its repetition, regularity, and perhaps
autocorrelation. This elementary, “classical” situation, in fact, gives the
most inconsistent and confusing results, which can be interpreted only by
assuming that in such a predicament the brain employs some a priori
criterion of “regularity” or “randomness.” There seems little doubt that
such a criterion is incorporated in the cerebral mechanisms, but it is cer-
tainly not a universal one, and as would be expected, personal variations
are found to be greatest when the situation is apparently most simple.
The rate and course of habituation to a set of single, irregular stimuli is
far more variable from person to person than is the pattern of interaction
to associated stimuli of unequivocal significance.
The discrepancies in the reports from various centres are largely due
to this factor; provided it is recognised there should be little serious dis-
agreement. It is not surprising that no two groups of workers have
selected the same experimental arrangements since there are so very
many to choose from, but some agreement on standard conditions might
be useful if co-ordinated research, mutual aid and complementary facil-
ities are envisaged for the future. The application of these methods of
studying human brain mechanisms to clinical problems will be particu-
larly difficult and confusing as long as every investigator is studying a
different problem with different methods.
The eight or so properties of evoked responses in nonspecific cortex
described above provide what may be called an operational specification
for the input circuits of a learning machine. They may be considered in
two groups. The first includes the four features which relate to the pre-
liminary operations on all afferent signals irrespective of their context:
Dispersive Convergence, Modality Signature, Idiodromic Projection, and
Differentiation. Since these mechanisms are concerned with the extrac-
tion of meaning from sensory stimuli, these preliminary processes may
be thought of in linguistic terms as specifying the orthography of the
cerebral code, the established procedure and accepted abbreviations for
signals from all sources. The second group of properties, Habituation,
Contingent Amplification, Contingent Attenuation, Unconditional Resto-
ration, specify the conjugation and syntax of signals according to their
context and destination.
Dispersive Convergence, Modality Signature, and Idiodromic Projec-
tion indicate between them the nature of the networks leading from the
first afferent relays to the cortical surface. They ensure that all nonspecific
areas receive information about all signals of uncertain or well-defined
significance, without overload or inter-modality modulation. The signals
thus relayed are distorted but not irreversibly corrupted. The process of
Differentiation is similar to the practice of writing an important term
with a Capital Initial whereby, after regular usage, it can be readily
identified, as in EEG. It is interesting that in some subjects this conven-
tion is observed to the letter; the responses to the first few dozen presen-
tations of Flicker have capital initials but are spelled out, while later
ones, in which only the onset of the unconditional stimulus matters, are
represented by the initial alone. The theoretical need for differentiation
was predicted and defined many years ago (Walter, 1953) in relation to
cerebral mechanisms of learning; it was demonstrated that this operation
-as a cerebral rather than receptor property-was essential for all signals
that might be classified as unconditional since knowledge of the start of
such events is all that is necessary to establish the significance of their
association. This prediction has been amply confirmed and helps to ex-
plain the graphic complexity of the evoked responses to novel or unclas-
sified stimuli. Any signal may be classified as novel, neutral, conditional
or unconditional: as a novel signal is repeated, its class must ultimately
be changed into one of the other three but until this transition has been
achieved the cerebral selective processes must in effect maintain a “null
hypothesis” and assume as little as possible about the ultimate category.
This means that the responses to the first few presentations of a stimulus
are likely to have features characteristic of both unconditional and condi-
tional responses; the projections to nonspecific cortex will be a blend of
both derivatives, one clipped or differentiated as if for unconditional
classification, the other protracted or extended in case the signal turns
out to be of conditional significance. If the stimulus remains neutral, that
is, it is neither preceded nor succeeded by any other event, then both
the differentiated and protracted components must wane, but they will
not necessarily wane at the same rate or in the same sequence. A stimulus
with established unconditiona1 significance-for example, a loud noise-
would be more likely to retain the differentiated appearance, while one
with a conditional quality would remain protracted. The difference be-
tween these pre-classified types might only be of degree but would
depend on experience and context.
In these experiments, the primary components of responses to single
auditory stimuli were in fact usually more persistent in normal subjects
than those to visual or tactile ones, while the secondary negative waves
of the visual and tactile responses were generally more prominent. The
component relations could usually be reversed by suitable association.
In one patient suffering from a chronic intractable anxiety with marked
fear of sudden noises, the nonspecific responses to auditory stimuli per-
sisted unchanged over several hundred presentations, retaining through-
out both their primary differential and secondary protracted components.
This effect suggested that, for this patient, the auditory stimuli, however
monotonous and objectively insignificant, were treated by the brain as
if they were perpetually both conditional and unconditional; in his own
words, he “could never get used to sudden noises.”
The relatively prolonged after-effects of conditional stimuli-explicitly
demonstrated in adaptive effector action-are reflected in two features of
the nonspecific responses, the coherent after-rhythms and the secondary
negative surges. The former, arising mainly in the parieto-occipital asso-
ciation areas, are related primarily to visual signals; they may be con-
sidered as conveying orderly information in the visual modality to other
zones. The secondary negative surges, their fields spatially congruent
with those of the primary negative waves, seem to have a more general
function. Since their amplitude and duration vary inversely with the SNC
of unconditional responses which they overlap or precede, it is tempting
to regard them as a primary dendritic gate, opened by significant condi-
tional signals and promoting the initiation of relevant effector action on
the conditional signal. In such conditions the unconditional signal has
become merely a timing pulse to indicate when the action planned in
response to the conditional signal should be taken. In accord with this
the muscular activity of the subjects often showed a preparatory incre-
ment in response to the conditional signal though the operant movement
was deferred until the onset of the unconditional one.
Some subjects learned to time their actions so as to control the uncon-
ditional signal after only a single flash, click or touch, that is with an
apparent reaction time of only about 50 msec. This was obviously due
to the establishment of a “time” association since the interval between
conditional and unconditional stimuli was always exactly one second.
When this was achieved the secondary negative surges were often par-
ticularly prominent and the degree of contingent attenuation of the
unconditional responses correspondingly great. The absence of this com-
ponent in the intracerebral recordings, its relative prominence in vertical
derivations and the correlation with relevant effector action suggest that
it may indicate diffusion of dendritic depolarisation into the motor regions
so as to form a functional link between the nonspecific sensory systems
in the frontal lobes and the motor areas in the Rolandic region. This
conjecture may throw some light on the controversy regarding the loca-
tion and nature of “closure” in conditional reflex formation. This may be
seen as dependent on both selective processing by nonspecific mech-
anisms as suggested by Western experimenters and also on cortico-
cortical interaction as assumed in the classical Pavlovian hypotheses. It
is not unusual in such disputes for the assertions on both sides to be
corroborated and their denials refuted.
Acknowledgment
These investigations are being made with V. Aldridge, R. Cooper, and
A. Winter. Most of the equipment is built by W. J. Warren, and the
patients are under the care of H. J. Crow and V. 0. G. Smyth. Part of the
expenses was defrayed by generous grants from the Mental Health Re-
search Foundation of London, and the Parapsychology Foundation of
New York. To all of these and to the very patient subjects I am deeply
indebted for the privilege of their collaboration.
References
BARLOW,J. S. 1960. Rhythmic responses to flash and alpha activity. EEG Clin.
Neurophysiol. 12: 317-326.
BRAZIER, M. A. B. 1960. Long persisting electrical traces in the brain of man. MOS-
COW Colloquium on EEG of Higher Nervous Activity. EEG Clin. Neurophysiol.
Suppl. No. 13.
BRAZIER, M, A. B. 1962. Normal and Abnormal Oscillatory phenomena in the electri-
cal activity of the brain. In Neural Physiopathology. Grenell, Ed. Hoeber, New
York, N. Y.
COOPER,R. & A. C. MUNDYCASTLE.1960. Spatial and temporal characteristics of
the alpha rhythm: a toposcopic analysis. EEG Clin. Neurophysiol. 12: 153-165.
COOPER,R. & W. GREYWALTER.1962. The conjugation and syntax of evoked re-
sponses in human non-specific cortex. EEG Clin. Neurophysiol. 14: 786.
COOPER,R. & W. J. WARREN.1961. The use of Barrier Grid storage tubes 9511A for
extraction of average evoked responses from the EEG. J. Physiol. 157: 38P.
CROW,H. J., R. COOPER& D. G. PHILLIPS.1961. Controlled multifocal frontal leucot-
omy for psychiatric illness. J. Neurol. Neurosurg. Psychiat. 24: 353-360.
ROSE, J. E. & V. B. MOUNTCASTLE. 1959. Touch and kinesthesis. I n Handbook of
Physiology. Williams & Wilkins. Baltimore, Md.
WALTER,W. GREY.1953. The Living Brain. Duckworth. London, Eng. Norton, New
York, N. Y.
WALTER,W. GREY. 1962. Oscillatory activity in the Nervous System. In Neural
Physiopathology. Grenell, Ed. Hoeber. New York, N. Y.
WALTER,W. GREY. 1963. Specific and nonspecific cerebral responses and autonomic
mechanisms in human subjects during Conditioning. I n Colloquium on Specific
and Unspecific Mechanisms of Sensori-motor Integration. Baldacci Foundation
and IBRO. In press.
WALTER,W. GREY,J. COHEN,R. COOPER & A. L. WINTER.1963. Analysis of Intrinsic
brain rhythms and responses evoked by visual, auditory and tactile stimuli in a
group of congenitally blind children. Conference on Technology & Blindness.
American Foundation for the Blind. In press.

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THE CONVERGENCE AND INTERACTION OF VISUAL,

AUDITORY, AND TACTILE RESPONSES IN HUMAN

FIGURE1. Lateral view of electrode sheaves implanted in the brain of an epileptic

FIGURE 3. Schematic illustration of the stimulus program for a standard experi-

LEM. Average of I 2 Trials

R. 6.12.61. Averaqe of I 2 Trials

DIST RAC TED by CONV E RS AT ION

reduced by voluntary hyperpnoea (Walter, 1962) is further evidence of

FIGURE 8. Idiodromic projection in scalp recording from vertex to intracerebral

577 jsc )Lv : * a 3 -

FIGURE 9. Anomalous differentiation of visual responses in normal subject A.

571 367 I src

occipital region as near to the visual cortex as possible, show considerable

the diversity is even greater. In one patient in a state of manic excitement

radiation of activity across the surface of cortex. Further consideration

FIGURE 12. Development of contingent effects in normal subject. A. First average

effect makes the investigation of response interaction details very diffi-

the normal subjects studied so far the effect of significance dilution or

tioned, in subjects who display this phenomenon, the amplitude and

FIGURE 14. Comparison of equivocal association and withdrawal of conditional

rhythms when the association is re-established and thus resemble more,

stimulus. Activity of the operant muscles was recorded in the experiments,

sa$ 1qlJK 6 L (D)

FIGURE15. Dilution of conditional probability by suggesting under hypnosis.

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