ON POSSIBLE

NEUROPHYSIOLOGICAL BASIS
OF THE PROCESS OF SHORT
TIME INTERVAL ESTIMATION

N.P. Podvigin, T.V. Bagaeva, D.N. Podvigina,

Pavlov Institute of Physiology, Russian Academy of Science
E. Poeppel
Munich Institute of Medical Psychology
The paper is dealing with the experimental data
obtained during the neurophysiological
investigations of mechanisms of brain neuronal
networks which serve for the estimation of short
time intervals. The investigations are being
conducted at present.

The research tasks are:

a) to search high-frequency rhythmic processes, which
reveal the functioning of mechanisms of time
quantification of impulse currents, in visual system
neurons’ pulse activity.
b) to construct (on the basis of our own data and
literature evidence) the hypothetical scale of time
quanta’s values.
c) to describe possible structure-functional organization
of the first levels of neuronal system that serves for
estimation of short time intervals. We suppose that
the neuronal system of estimation of time intervals
operates on the basis of the scale of time quanta.
The processes of time quantification of impulse
currents in brain neuronal networks were
theoretically justified by John von Neumann, in
whose work the concepts of “temporary delays”
and “periodical impulsators” were first used for
description of neuronal networks.
The phenomenon of time quantification of signals has
also been revealed in physiological systems.
[Poeppel E., (1971); Poeppel E., Logothetis N.,
(1986); Ilmberger J., (1986) et al.] It’s believed that
the scale of subjective measurements of time
intervals operates on the basis of the minimal
temporary interval-quantum equal to 30-40 ms, that
is the period of gamma-oscillations. It’s also
believed that the scale of indication of current time,
as well as the scale of time quantification, are based
upon the system of neurons which have different
oscillatory frequencies. [Poeppel E., (1989), (1971),
(1978) et al.]
We’ve explored high-frequency range of rhythmic
activity of neurons (time intervals of from 20 to 2
ms and frequencies of from 0.05 to 0.5 kHz). That
enables us to determine if there are time quanta of
less than 30 ms in impulse signals and how the
high-frequency range of time quanta’s scale, which
takes part in brain neuronal networks’ operating,
may be organized.
Methods. Impulse reactions of neurons of cat
visual system were analyzed: the reactions of
retina ganglionic cells, neurons of lateral
geniculate nucleus (LGN) and cortex. Spectral
analysis of neuronal impulse reactions (within
the range of to 0.5 kHz), analysis of distributions
of inter-impulse intervals (DII) (measuring
precision – 0.1 ms) and autocorrelation analysis
were carried out.
In the Fig. 1 there is an example of the results
obtained using different types of analysis.
FIG.1 An example of the results obtained by using different types of analysis.
Registration information (the number of experiment, pool and record, the type of
reaction) is pointed out in the top graph.
The results. There are several suppositions we could take into
account beginning the research of neuronal system of time
intervals’ estimation. It’s possible, for instance,, that the
neuronal system of time intervals’ estimation is spatio-
temporally determined, and operations carried out with this
system are based on the assembly of neurons – oscillators
distributed within brain structures. It’s also possible that
the neuronal system in question is based on certain
temporal rules of continuous random processes – impulse
currents in brain neuronal networks. It’s not improbable
that the neuronal system under consideration combines
both principles mentioned above. We’ve proceeded from
the latter assumption.
While exploring the system of time quantification as a basis
for the system of time intervals’ estimation and proceeding
data analysis we’ve assumed that the most adequate (for
the mechanisms of brain neuronal structures) oscillatory
characteristic is not oscillatory frequency but the temporal
parameter of oscillations, that is their period.
Analysis of the range of high-frequency oscillations and small
values of inter-impulse intervals demonstrated that peaks in
spectral characteristics of neuronal impulse reactions could
be observed almost within the whole range analysed - up to
470 Hz, - and in DIIs there were modes on 0.9-1.2 ms.
(Fig. 2, 3)
FIG. 2 Spectral characteristics of impulse reactions of visual neurons with
dominant peaks on different frequency values within frequency range under
consideration (20-500 Hz). Data from ganglionic cells (GC), neurons of lateral
geniculate nucleus (LGN) and visual cortex of the cat. Digits above plots indicate
the number of the experiment, pool (p) or neuron (n) number, the number of the
record, neuron type, brain structure, the frequency of the dominant peak (Hz).
If the data were averaged the numbers of averaged records are pointed out in
brackets. Abscissa axis: two scales – frequencies (Hz) and values of oscillatory
periods corresponding to the frequencies.
FIG. 3 Distributions of inter-impulse intervals (DII) in reactions of visual
neurons which have maximum modes within the range of short inter-impulse
intervals (0.1 – 30 ms). DII of the reactions of cat canglionic cells (GC), LGN
neurons and visual cortex neurons (VC) to visual stimulus. The arrows under the
top histogram indicate the range analyzed.
Multiple values of modes in DIIs and
spectral characteristics could be
observed. (Fig 4, 5)
FIG. 4 The examples of DIIs with multiple mode values in impulse reactions
of neurons from three levels of visual system.
FIG. 5 The distribution of frequency and periods of gamma-oscillations of three
types. On the left there are the histograms of distributions of dominant peaks’
number (ordinate axis) in spectral characteristics of responses of three types
of neurons. Abscissa axis: frequency range analyzed. On the right there are the
distributions of oscillatory periods from the histograms shown on the left.
Multiple (or near multiple) values of
DII modes were revealed within
different subranges of the frequency
range analyzed. (Fig. 6)
FIG. 6 DIIs of neuronal impulse reactions (LGN and GC neurons) which peaks
have multiple values within different parts of analyzed range of inter-impulse
intervals (0.1 – 30 ms). For details see registration information for each DII.
One interesting moment should be
mentioned: (Fig. 7) in DIIs there
were high-frequency components
(with a period of about 1 ms)
modulating the function of inter-
impulse interval distribution.
FIG. 7 On the top there is an example of DII in background activity of retinal
ganglionic off-neuron while the retina was illuminated (420 Lx). The registration
was 3 minute long. 1667 impulses were recorded. Under the DII there are the
distributions of high-frequency spikes within each peak (I, II, III) in the top
graph.
According to the literature evidence [Poeppel E., (1989),
(1995), (1996) et al.] and our own data we suggest (as
a working hypothesis) the temporal scale within
which the system of multiple time quantification of
signals and the system of subjective estimation of time
intervals operate. The structure of this scale (Fig. 8) is
based upon following theses:
a The system of time quantification of impulse currents
has logarithmic scale of values of temporal quanta
which are multiple to minimal (“fundamental”)
temporal quantum.
b The value of fundamental temporal quantum is
determined in the range of 1-2 ms based on time
resolution of impulse sequence in neuronal chains.
c For the system of time quantification informatively
significant parameters of oscillatory processes are not
their frequencies but their time characteristic –
periods.
d A neurophysiological basis for time quantification
scale is the network of oscillating neurons distributed
in neuronal structures and also quasi-periodic pulse
bursts in neuronal activity revealed in DIIs of impulse
signals.
e Such network of oscillating neurons is a system with
distributed parameters. Elements of the system –
temporal scale supporters – are the operators without
strictly determined spatio-temporal coordinates.
FIG. 8 The hypothetical scale of temporal quantification of impulse currents. On
the left there is the scale of logarithmic values of temporal quanta (lg T), of their
absolute values and the scale of the frequency values which periods correspond to
the values of temporal quanta. Minimal temporal quantum is equal to 1.5 ms.
Suggested temporal scale of multiple
values of temporal quanta may serve
as a neurophysiological basis for the
system of time interval estimation,
i.e. the time perception system, and
also may temporally determine
neuronal processes of functioning
brain.
Here are the main aspects of structural organisation
of hypothetical neuronal system for subjective
estimation of time intervals:

a) The principle of reception of external signal by

time perception system seems to differ greatly
from the ones of sensory systems. We suppose
that sensitive (“receptive”) level of time
perception system is based upon the structure of
neurons – oscillators, which is a number of
quantified standard values of time intervals. The
values of periods of oscillatory processes with
different frequencies are supporters for these
standard time intervals.
It should be mentioned that some particular features
of the system in question (for example, these
ones are: the system of neurons – oscillators is
not spatially determined; the current of
oscillations generated by neurons has a discrete
nature; information on temporal characteristics of
events of outer world is used by all brain systems
at the same time, and others) allow us to suggest
that the system of perception and estimation of
time intervals has some elements of stochastic
system. This aspect isn’t discussed in the paper.
b) Such system of etalons – indicators of input
temporal characteristics is a specific field of “time
receptors”.
We think that within the system estimating time
intervals there are (as well as for other sensory
modality) operating diapasons – some parts of the
common temporal scale which are determined by
the values of time intervals estimated.
c) With the change of the values of time intervals
measured position of operating diapason (or
dynamic characteristic of the measurements) may
move (as well as for other sensory modalities)
within common temporal scale (or the system of
sub-scales within one common temporal scale may
be used).
d) Signals generated by the field of “time receptors”
seem to be primarily processed with the operations
which are algorithmically similar to the ones that
take part in processing of other modality signals.
For example it’s been shown by now that there are
threshold mechanisms of time perception which
determine absolute and differential thresholds;
mechanisms of temporal and spatio-temporal
summation; [Podvigin N., (1979), (1970) et al.]
mechanisms of taking the logarithm of the values
of time intervals measured [Eisler H., (1975)] etc.
Thus we have attempted to set out the hypothesis of
structure – functional organization of the first
levels of neuronal system for measuring and
estimating of time intervals in time perception
processes.

The concept of “fundamental” temporal quantum

which determines the measuring temporal scale,
multiple to this quantum, has been theoretically
justified for time quantification systems in
micro-world phenomena. [Markov M., (1958);
Blochintzev D., (1970) et al.] Usage of this
concept enables to determine the order of
accuracy of calculation of short time intervals.
 References
[1] Poeppel E. (1971) Oscillations as a possible basis for time perception.
Studium Generale, Vol. 24, pp. 85-107
[2] Poeppel E., Logothetis N. (1986) Neuronal oscillations in the brain.
Discontinuous initiations of pursuit eye movements indicate a 30-Hz
temporal framework for visual information processing.
Naturwissenschaften, Vol. 73, pp. 267-268
[3] Ilmberger J. (1986) Auditory excitability cycles in choice reaction time and
order threshold. Naturwissenschaften, Vol. 73, pp. 743-744
[4] Poeppel E. (1989) The measurement of music and the cerebral clock: a new
theory. Leonardo, Vol. 22, No. 1, pp. 83-89
[5] Poeppel E. (1974) Oscillations as possible basis for time perception.
Studium Generale, Vol. 24, pp. 85-107
[6] Poeppel E. (1978) Time perception. In: Handbook of sensory physiology.
Vol. VIII, chapter 23 Perception, pp. 714-729
[7] Poeppel E. (1989) Taxonomy of the subjective: an evolutionary
perspective. In: Jason W. Brown, M.D. (ed), Neuropsychology of visual
perception, pp. 219-232
[8] Poeppel E. (1995) Time perception: problems of representation and
processing. In: Micael A. Arbib (ed), The handhook of Brain Theory and
Neural Networks, pp. 987-990
[9] Poeppel E. (1996) Reconstruction of subjective time on the basis of
hierarchically organized processing system. In: M.A. Pastor, J. Artieda
(eds), Time, Internal Clocks and Movement, pp. 165-185
[10] Markov M. (1958) Hyperons and kaons, Phismath State Publishing
House, Moscow
[11] Blochintzev D. (1970) Space and time in micro-world, Nauka, Moscow
[12] Podvigin N. (1979) Dynamic properties of neuronal structures of visual
system.
[13] Podvigin N (1970) On the non-linear transformations of signal in retina.
In: The research of principles of information processing in visual system,
pp. 28-39
[14] Eisler H. (1975) Subjective duration and psychophysics. Psychological
review, Vol. 82, No. 6, pp. 429-450