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Abstract. Oscillatory neural activity in the frequency The present study is an attempt to characterize the
range 7-12Hz is observed in the lateral geniculate thalamic oscillatory activity observed in the lateral
nucleus (LGN) of the lightly anesthetized cat. This geniculate nucleus (LGN) of the cat. The approach
paper describes a series of experiments in which the taken is based on engineering systems theory and
interactions between ongoing oscillatory potentials regards the LGN as a system with primary input from
and periodic photic and electrical stimuli are analyzed the retina and a measurable output signal in the form
using frequency domain techniques. The principal of a gross neural potential (see Fig. 1). This description
results of these experiments are consistent with a is clearly a simplification as recent physiological evi-
model of the neural system as an entrainable oscillator dence (Singer, 1977) has shown. However, it provides a
in which ongoing oscillations are suppressed by stimu- basis for expressing quantitative relationships describ-
lation at nearby frequencies, but coexist with stimulus ing retinal input and its influence on ongoing intrinsic
frequencies farther away. The physiological interpre- LGN activity. The intent of the work is not to model
tation of these results may be closely tied to the role of the physiological origins of LGN activity in detail, but
the LGN as a "gating" mechanism between retina and rather to derive a description of the interaction be-
cortex. tween the activity and retinal input. Systems theoreti-
cal models provide a useful description of this type of
interaction.
RECORDING
Introduction ELECTRODES
0340-1200/79/0033/0063/$02.60
64
In this paper we consider the effect of externally cell and interneuronal responses in the rat during
applied single-pulse and periodic stimuli upon the electric stimulation of the optic tract and visual cortex.
intrinsic, alpha-like, rhythmic activity of the cat's lateral They concluded that the LGN has recurrent inhibition
geniculate nucleus (LGN) that is centered at an expe- capable of oscillating at about 10 Hz. This result has
rimentally measured frequency, f,. This experimental also been discussed by Creutzfeldt (1966), Andersen
approach has led us to examine in detail a class of and Sears (1964), Andersen and Andersson (1968),
engineering models which seem to offer insight into Eccles (1969). More recent studies summarized by
interactions between intrinsic and extrinsic oscillatory Singer (1977) suggest that localized inhibition may be
mechanisms. One of these, the entrainable non-linear mediated by LGN interneurons while more diffuse
oscillator (Pavlidis, 1973), has been examined in the recurrent inhibition may be due to perigeniculate
context of other biological systems, for example, to neurons. These results have suggested that perigen-
model diurnal biorhythms which may be affected by iculate neurons may be associated with the widespread
imposed light-dark programs. We have experimentally oscillatory phenomena.
examined two effects in the LGN which would help to LGN single unit activity was categorized by Levick
differentiate between a linear system model and a non- and Williams (1964) as "stable" or "unstable". Bishop
linear entrainable oscillator model. These are: sup- et al. (1964) divided the unstable units into 2 groups:
pression of alpha-like activity by periodic stimuli, and "fast cyclic" and "rhythmic burst" types. The single
the dependence of the transfer characteristic upon unit stable mean rate and fast cyclic types of firing have
stimulus amplitude. This paper describes these effects their activity drastically changed when the retinal
and relates them to the entrainable oscillator input to the LGN is stopped, but rhythmic bursting
hypothesis. types of activity continue even after the LGN is
isolated from retinal input.
Although the LGN activity investigated in these
experiments is assumed to be intrinsic to the thalamus,
Background inputs from other parts of the brain may influence it.
Important inputs originate from the ascending re-
Previous physiological experiments suggest that there ticular activating system (ARAS) (Arden and
are at least two classes of spontaneous oscillations S6derberg, 1961; Ogawa, 1963; Satinsky, 1968;
which can be observed in the mammalian lateral Bowsher, 1970; Singer, 1973; Singer and Schmielau,
geniculate nucleus (LGN). One of these seems to 1976) and from the pons (Ponto-Geniculo-Occipital,
depend upon light-induced retinal activity (Kozak, PGO, Waves : Singer and Bedworth, 1974 ; Malcolm et
1971) whereas the other appears to be an intrinsic al., 1970; Bizzi and Brooks, 1963). Singer (1977) has
LGN oscillation due to the neural net of the LGN and recently reviewed corticofugal and reticular influences
its associated nuclei and does not seem to require on thalamic structures.
retinal input as a necessary condition for oscillatory Despite the influence of extrinsic factors, the work
behavior. The purpose of our experiments was to of Kato (1974) on the isolated, sliced LGN of the cat
determine what effect externally applied stimuli have provides evidence in support of an oscillatory process
upon the latter, intrinsic LGN oscillation, and to test that is intrinsic to the LGN and is due to the structure
the hypothesis that the LGN is an oscillating system of the neural net. Removal of visual cortices did not
that can be entrained by external periodic signals of stop the oscillation of neurons in the LGN in experi-
appropriate frequency and strength. ments of Satinsky (1968), and he also concluded that
Spontaneous neuronal activity is a feature of most the oscillatory activity may be due to reverberating
sensory systems (Thompson, 1967, p. 287). Oscillatory circuits within the LGN itself.
potentials in the visual system were first observed in Several previous determinations of the LGN tem-
the dark-adapted electroretinogram (ERG) and in the poral transfer characteristics have been made: Maffei
electric records from the optic tract (OT) by Granit and Rizzolatti (1966), using sine-modulated light in
(1933) and by Bartley and Bishop (1942). The response Nembutal-anesthetized cats; Lopes da Silva et al.
to single light flashes in human ERG's (Cobb and (1970a, b) using sine-modulated light in alert dogs
Morton, 1953) also had oscillatory components. having chronically implanted electrodes, and Rosen
Bishop and Davis (1960) showed that the LGN of the and Vastola (1973) who used sine modulation of rapid
cat generated slow (about 10 Hz) rhythms. electric shocks applied to the optic nerve to determine
The discovery of LGN principal cell repetitive the phase shift through the LGN. None of these
firing by Bishop et al. (1953) led them to postulate an experiments, however, were designed to investigate the
inhibitory interneuron associated with the LGN. interaction of the periodic stimulus with the intrinsic
Burke and Sefton (1966a, b, c) recorded single principal LGN oscillations.
65
stainless steel bipolar electrode. The electrode was variance in the noise spectral estimate, thus allowing
designed so that the exposed optic nerve could be the harmonic signal to stand out. Spectral averaging
pulled through it after enucleation. The electrode leads also permits one to determine the statistical signific-
were shielded, and the shielding was attached to a ance of spectral peaks. Our approach was to calculate
grounding electrode in the form of a stainless steel a mean and standard deviation of each spectral point
spiral wire wrapping the optic nerve between the point using period averaging of 10-20 repeated trials.
of stimulation and the brain. This minimized the shock
artifact as recorded in the brain.
Results
The overall electrical noise, recorded from coaxial
electrodes in Ringer's solution or from electrodes in Ambient LGN Activity
dead cats, was 5-10 gV peak-to-peak (filter band pass 1
Low frequency ambient activity in the LGN
to 300 Hz) for single sweeps and correspondingly less
(0.5-30 Hz) is characterized by a diffuse band of low
for averaged records (typically 2 laV for an average of
frequency noise in the 0.5 to 3 Hz region and by a
ten sweeps), Spectral analysis of the noise showed no
relatively sharper band of activity centered between 7
significant peaks at 60 Hz and its harmonics, or at any
and 12Hz, depending upon the individual cat. The
other frequencies.
latter activity will be referred to as LGN intrinsic or
The amplitude of the waveforms recorded during
natural oscillation. The amplitude, spectral width, and
the stimulation was from 100 to 600 gV, and therefore,
center frequency of the LGN natural oscillation vary
was well above the abient noise level. Locally made
with time in a single cat. There are periods when the
amplifiers with variable bandpass filters were used to
activity is quite regular. Quiet periods can occur under
amplify the signal and to filter out higher frequency
the effect of barbiturate anesthesia or when there has
signal components that could cause aliasing during
been no stimulus for one half to one hour. This effect
sampling. At this point in the signal processing, the
may be due to a change in the level of the cat's
stimulus signal plus the OT and LGN responses were
alertness or to a reduction of the on-going retinal
recorded using an Ampex model SP 300, AM-FM type
activity. During an active period the oscillation tends
recorder. On-line frequency analysis or multichannel
to occur in wave packets or spindles. The most re-
time averaging was done on a Digital Equipment
gularly occurring spindles are about one second in
Corp. PDP-8/L computer equipped with a multiplexed
duration and recur, on the average, every 2-4 s. Much
10 bit analog-to-digital converter, a 1 KHz clock, and a
of the time the spindles are less regular, both in their
digital-to-analog converter for display purposes. A
duration and in the interspindle intervals.
Grass Model S 88 stimulator controlled timing of the
Figure 2 shows an example of irregular LGN
stimulus and of the start of computer data acquisition.
oscillatory activity. The time record, Fig. 2A, is similar
Analytical to a sine wave to which noise has been added. The
Frequency analysis was performed using the Fast obvious periodicity corresponds to approximately
Fourier Transform (FFT) algorithm (Cooley and 10 Hz, and the peak-to-peak (p-p) amplitude varies
Tukey, 1965) combined with other subprograms to from 20 to 120 ktV. The accompanying averaged ASD
digitize and display data, and then to average and plot plot, Fig. 2B, was obtained from ten segments of
amplitude spectra. Normally, 256 data points were sampled time data, each lasting 2.048 s. The interval
between consecutive sampling sections was ten se-
taken at a sampling rate of 125 Hz (8 ms/address). The
sampled data were multiplied by a raised cosine (han- conds, so that the total time during which the activity
ning) window. The windowed time series was the input was monitored amounted to slightly over two minutes.
For this time the bandwidth of the oscillation shown
data to the FFT algorithm. The result was a discrete
Fourier transform consisting of 128 complex points in on the plot was one Hz and the center frequency was at
the frequency domain. A discrete estimate of the 9.8 Hz.
Although the amplitude of the LGN oscillation is
Amplitude Spectral Density (ASD) was obtained by
taking the square root of the sum of the squares of the time-varying, the frequency of the peak amplitude
never varied more than _+1 Hz for any individual cat
real and imaginary points for each frequency incre-
with intact eyes. Larger changes were recorded when
ment of the FFT output. For convenience, these
the LGN was isolated from retinal activity after
discrete periodograms will be referred to as ASD data.
A specific number of successive ASD data files obtain- enucleation.
ed under the same experimental conditions were aver-
LGN Oscillations Following Bilateral Enucleation
aged together. The averaging procedure is desirable
when there is a harmonic signal superimposed upon a Because the principal afferent connections to the LGN
band-limited white noise, 'because it decreases the are from the retinal ganglion cells via the optic nerve,
67
A
there is a possibility that the 7-10Hz oscillation as
recorded in the LGN could actually originate in the
retina. Simultaneous gross electrode data from 14 cats
showed no evidence of this. There were, however, some
peaks in OT spectra falling in the 2-5 Hz range that
were mirrored in the LGN spectra.
In three cats, the eyes were removed bilaterally to
identify the role of the retinal input on the LGN gross
potential activity. The enucleation reduced the amount
B of low frequency (0.5-20 Hz) activity recorded from the
0. OT electrode, and also decreased the LGN amplitude
O. "~ ( Peokof oscillotion spectral density in the same frequency region, except
for the 7-9 Hz intrinsic oscillations which continued
~,~o. Bood .idth o, following bilateral enucleation. This procedure re-
o
duced other low frequency activity and made the
intrinsic oscillation more obvious. One cat showed a
large increase in the amplitude and regularity of the
LGN oscillations. This is illustrated in Fig. 3. The
"~ 0
typical reduction of the low frequency activity in both
I I
E 10 20 OT and LGN is apparent, along with the increase in
,ci
Frequency in Hz the amplitude of the LGN oscillations. The second
harmonic of the LGN oscillation becomes visible in
Fig. 2. A Intrinsic oscillatory activity in the cat's Lateral Geniculate
Nucleus (LGN). Darkness, eyes intact. A single sweep of ongoing the spectrum taken after enucleation. A slight dimi-
activity. Coaxial electrode. Amplifier band-pass 1-200Hz. B nution (from 8.8 to 7.3 Hz) of the center frequency also
Amplitude spectral density (ASD) of activity shown in A. Bandpass occurs.
1-30 Hz. Ten sweeps averaged in frequency domain. Sampling rate : Table 1 gives the frequency shifts for all three cats
125 Hz. Time series length: 2.048 s. The frequency peak of intrinsic
oscillation of the alpha-type is indicated by arrow after enucleation.
These shifts are equal to, or slightly greater than,
the frequency variability in the LGN oscillations of
cats which were not enucleated.
Table 1. LGN oscillation center frequency
2o] A B
,olFL LoN
,.,or.
Fig. 3A-D. Simultaneous OT and LGN am-
plitude spectral densities (ASD's) before and
o
0
o lo 0 I0 after bilateral enucleation. Different prepara-
0 tion than in Fig. 2. Ten sweeps averaged for
each plot. Sampling frequency 125 Hz. Sweep
2O D
1~ OT Before OT After
lengths 2.048s, as in Fig. 2B. A and C.
Before enucleation, darkness. The alpha-like
! LGN intrinsic oscillation is indicated by the
arrow. Oscillatory activity at this frequency is
IO L___LL,~NoiseLevel absent from the OT record. B and D. Same
~o o
,
10
ot
0 10
,
preparation, after bilateral enucleation. The
oscillatory LGN activity is more coherent,
has a higher amplitude and its second har-
Frequency in Hz Frequency in Hz monic is visible
68
,]i'
p < .01 differed only slightly from f~. r: ratio of the
p= .02 difference between the means at neighboring
points to the standard error of the difference
'tJ C 20 C22 of the means, p: t-test significance level, S.L. -
steady light. Coaxial electrodes. Amplifiers'
I I band-pass : 1-30 Hz. Sampling rate : 125 Hz ;
IO 2'0 30 Dark 10 2O 3O 256 points sampled
70
c 20
Oj ,
A and p are shown between the last point that we
interpret as "suppressed" and the first "recovered" one.
All five cats analyzed in this way showed a significant
jump that differentiated the "suppressed" response
-8 from the "recovered" one. This "jump" occurred be-
= I0
tween 15 and 25Hz. We do not have quantitative
0 . . . . information on the influence of the stimulus strength
upon the suppression effect.
fFi B
L G N Frequency Response
The averaged amplitude spectra of simultaneously
~
<
zo
O , , ~
J
,
recorded L G N and O T responses to periodic stimulus
trains of various frequencies were compared. The
spectra were averaged over 10 to 20 successive trials.
,o],CTRANSF
9 R
The stimulus frequency runs were presented in random
fn C order. In some cats, the stimulus strength was varied
by using neutral density filters for photic stimulus and
by varying the stimulus voltage for electric stimulus.
The raw d a t a for each stimulus frequency, f~, consisted
:~0.5 of two averaged amplitude spectra, one for the L G N
and one for the OT. The mean values and the standard
<
deviations of the spectra of f~ were used for the
o frequency response determinations.
0 5 I'0 1'5 io In order to derive a frequency response estimate for
F r e q u e n c y in Hz
the L G N , we divided the L G N frequency response
Fig. 7. A and B LGN and OT tuning curves with electric stimulation function by the corresponding O T frequency response
for one experimental run. Amplitude of spectral peak correspond- function, for each case separately. Figure 7 shows an
ing to the stimulus frequency is plotted versus stimulus frequency. example of the L G N frequency response, the O T
Brackets show _+1 S.E. C Transfer characteristic is derived from OT frequency respons e and the calculated transfer charac-
and LGN. Arrows mark the frequency of the natural oscillation,
f,, which was measured prior to periodic stimulus tests
teristic, for one experiment using electric shock stimuli
to OT. The standard error of the mean of each data
point is marked by brackets and was determined from
the L G N and OT frequency responses. Standard
the L G N natural oscillation recovers when the sti- errors are found using an approximation for the ex-
mulus frequency is sufficiently high. pected value of the ratio of two random variables.
Figure 6 shows that this recovery takes place above
15 Hz. Since all the points on the plot have standard
errors, the significance of the increase in the amplitude
of the L G N natural oscillation above flicker frequen- where :
cies of 15 Hz, can be tested by taking the ratio, r, of the E x = standard error of the transfer characteristic
difference of the mean values divided by the standard E 1 = standard error of the L G N response
error of the difference of the means: E o = standard error of the O T response
A t = L G N response mean value at fs
x-y
r = (sX2 +sp2)i/2 A 0 = the O T response mean value at f~.
This error expression is an upper bound, since it
where: neglects a co-variance term which would reduce the
2 = mean value of the first recovered point error .if the input and output were correlated.
= mean value of the last suppressed point A total of 17 frequency response runs were made in
s2 = standard error of 2 nine cats. Runs for three stimulus intensities were
sp = standard error of y. made in four of these cats. D a t a from six of the nine
F r o m r the probability, p, that this difference would be cats show an absolute or relative m a x i m u m in the
exceeded by chance can then be determined from L G N frequency response, when the stimulus fre-
tables of the t distribution. On each part of Fig. 6, the r quency, fs, is at or near the L G N natural frequency, f,.
71
1
0.2
(1966) and Singer et al. (1972). The spontaneous ac-
0. I
tivity, Fig. 2A also agrees in frequency with the results
0 /
-0.1
s ,5 io of Bishop and Davis (1960). Spectral analysis of the
0.2 activity, Fig. 2B, is like that found by Lopes da Silva et
0.3 al., (1970a, b) in the dog LGN, except that the fre-
quency of the ongoing oscillation in the dog is about
+0.5
12Hz instead of 7-10Hz in the cat.
"-'Z~ 9 40 Volt Stimulus Most of the experimental frequency response find-
+0.25 \ 0 t 0 Volt Stimulus B ings are similar to those of Maffei and Rizzolatti
\ V 7 Volt Stimulus (1966), who determined the L G N transfer characteris-
0 9 i tics using single unit electrodes. Their gain vs fre-
5 ',~" ' ' OH quency curves agree roughly with ours in that the
response tends to fall off above 10 Hz. Our findings
-0.2 5 in x
show a peak which occurs at the natural oscillation
frequency. The region in which this peak occurs is
-0.5 rather narrow and, unless the test frequency incre-
Fig. 9A and B. Phase transfer points taken from Fourier transform of
ments are equal to 1 Hz or less, it is not likely that the
the time-averaged stimulus, LGN response and OT response. 2.048 s peak would be found. Maffei and Rizzolatti's test
samples taken from the middle of a 3 s long train of square wave frequencies in this region were at 2, 5, and 10 Hz. Their
photic flicker data sampled at 250Hz. Analog filters set 0-30Hz. frequency increments were too widely spaced to have
Stimulus frequency range is from 5-20Hz. A Phase shift (L-O)
resolved the peak response which our data show.
through the LGN taken from a photically stimulated cat. B LGN
phase shift for an electrically stimulated cat. Three different stimulus The phase of our transfer characteristics changes in
voltages are used: 40, 10, ? V the same amount over approximately the same fre-
quency interval as the phase response which Mallei
and Rizzolatti (1966) show. Both the phase changes
amount to approximately - 180 degrees. However, our
7 V, tend to keep the phase shift nearly constant over a data show a phase lead below the natural frequency
3Hz range. Increasing the stimulus strength to 10V and a phase lag when the stimulus is above the natural
decreases this range of "flatness" to 2 Hz. There is no frequency. The data of Maffei and Rizzolatti (1966)
evidence of "flatness" around f, in data from the 40 V show a phase which is always less than or equal to zero
stimulus run. The rate of phase change of a linear fit to degrees, i.e., there is no phase lead in their data. It is
the data in the 3 to 10Hz range gives a slope of possible that the dynamic population response obtain-
approximately -0.15 radians per Hz. If the shift were ed with a gross electrode may differ from the responses
caused by a time delay, the delay would amount the of a single unit.
approximately 75 ms. Rosen and Vastola (1973) also investigated the
To summarize, L G N data from both electrically transfer properties of the lateral geniculate nucleus
and photically stimulated cats show a phase decrease using sinusoidally modulated pulse trains of electric
with increased stimulus frequency. There is a relative shocks applied as a stimulus to the optic tract. It is not
lead below f, and a relative lag above f~. The LGN possible to compare their results with ours directly
phase passes through zero in the region where fs =f,. because they show the relationship between the modu-
For photic stimulation, zero phase occurs at a fre- lation frequencies of the stimuli and the average firing
quency 1-2 Hz higher than f~. The phase shift cannot rates of single cells in the LGN. This averaging
be adequately characterized as a simple time delay. technique measures only the shift in mean firing rate
due to sinusoidal modulation. Such a response is quite
different from the mean transfer functions which are
Discussion discussed here.
The power spectrum of LGN activity recorded by
Comparison of Results with Previous Findings Lopes da Silva et al. (1970a) shows a peak at l l . 7 H z
The pulse response in Fig. 4 with the oscillatory after- which corresponds to the alpha rhythm of the dog
effect is similar to the slow rhythms found in the cat (Lopes da Silva et al., 1973a, b). This spectrum looks
L G N by Bishop and Davis (1960). There is a single quite similar to the spectrum of the spontaneous
spike followed by several cycles of an 8-10 Hz damped activity which we show for cat, Fig. 2B, except that the
oscillation. The period of these oscillations corres- frequency peak for cat is, on the average, somewhat
ponds in frequency to the 100ms postsynaptic IPSP's below 10 Hz.
73
Systems Models shift depends to some extent upon the kind of stimulus.
The strength of the electric shock train stimulus in
While it is difficult to model a complex neural network
some experiments does affect the phase response,
in sufficient detail to interpret the observed behavior in
particularly when the stimulus frequency is close to the
terms of specific physiological phenomena, it is often
L G N natural frequency.
fruitful to explore analogous behavior in order to gai n
The experimental results concerning all the above
insight into a physiological system. Among physical
three points may be related to certain classes of
systems, a particular class of nonlinear system, the
models. The gain results are like those of a second-
entrainable oscillator, would seem to offer such an
order damped linear system, but the results could also
analogy for the present study.
be consistent with the response of a non-linear system
Engineering systems theory treats frequency en-
with small gain dependence upon the stimulus ampli-
trainment in the context of a well-defined self-
tude. The phase results are similar to those of a non-
oscillating system with a fundamental frequency, f,.
linear system because the phase is dependent upon the
Such a system can be mathematically described by a
stimulus strength. In some non-linear systems the
nonlinear differential equation. The mathematical be-
change in phase with stimulus strength is a more
havior of the model is examined using a periodic
sensitive parameter than is the change in gain.
forcing function. Frequency entrainment is said to
A variety of models can be invoked to explain
occur when the driving signal "takes over control of
experimental results. As complexity increases, more
the oscillation frequency" (Cunningham, 1958, p. 213)
results are explained. A single non-linear entrainable
i.e., the component at f, is suppressed or abolished
oscillator model explains our results more completely
during periodic driving by signals of appropriate fre-
than a single linear system model. As a further step, an
quency and sufficient strength. This effect depends
ensemble of non-linear entrainable oscillators might be
upon the strength and frequency of the injected signal
considered.
and upon the characteristic of the nonlinear system
itself.
The following principal characteristics of the ex-
perimental system with periodic stimuli may be com- Conclusions
pared to the entrainment model: The intrinsic oscillatory activity of the L G N suggests
1) Background activity. There is an ongoing neural that large numbers of neurons may "spontaneously"
activity in the L G N and which can be characterized as synchronize. The origin of this synchrony is not clear.
having a frequency component of fairly narrow band- It is reasonable from physiological considerations
width (Fig. 2). The "ringing" response to single light (Singer, 1977) to suppose that such oscillatory activity
pulses (Fig. 4D) strongly suggests that there is an originates within the thalamic structures but may be
interaction between an external visual stimulus and the strongly influenced by other inputs. Recurrent in-
background activity as recorded in the LGN. hibitory feedback exists between the perigeniculate
2) Suppression by Periodic Stimulation. As discussed nucleus and the LGN principal cells, and the mapping
above, frequency entrainment is an effect which occurs of this inhibitory influence in the L G N appears to be
in a system that can be described by a non-linear widespread rather than local. Reticular formation
differential equation that allows a periodic system inputs, in turn, appear to inhibit perigeniculate neu-
"output" to appear even when there is no system rons (Singer, 1977). In addition, recurrent inhibitory
"input". This intrinsic output can be suppressed or interneurons occur within the LGN, though their effect
completely eliminated by certain periodic input sig- seems to be more local. Thus, from anatomical con-
nals. Periodic electrical stimulation to the OT sup- siderations, there appears to be a network of neural
presses the intrinsic, alpha-like, oscillation which oc- interactions which could produce oscillatory activity
curs in the LGN. due to feedback inhibition with a delay resulting from
3) Gain and phase response during periodic stimu- synaptic summation.
lation. The L G N transfer characteristics show a maxi- The control of inhibitory feedback by reticular
mum or relative maximum when the stimulus fre- formation inputs suggests a mechanism which could
quency is at the L G N intrinsic oscillatory frequency. induce or suppress such oscillatory actNity. In this
There is no significant change in the transfer character- case, oscillations would tend to occur when the re-
istic with change in stimulus strength. The phase shift ticular formation is least active or in states of low
of the fundamental frequency through the L G N de- alertness. Behavioral manifestation of the occurrence
creases with increasing frequency, and has a shift of of alpha activity in EEG also suggests that such
zero when the stimulus frequency is at or near the oscillatory activity may be associated with a "resting"
L G N natural frequency, s The nature of the phase rather than "active" state of the neural system.
74
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