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  • 2015-2019  (121,029)
  • 1975-1979  (24,223)
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  • Books  (79)
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  • 1
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    Biological cybernetics 17 (1975), S. 77-79 
    ISSN: 1432-0770
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Computer Science , Physics
    Notes: Abstract An approach is described by which certain deductions can be made concerning the natural underlying structure of visual space. The procedure is indirect and makes use of the notion of visual recognition defined with respect to an arbitrarily fixed structure. Consideration of the set of mappings associated with such recognition is shown to lead to a condition that must be satisfied by any proposed underlying structure.
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  • 2
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    Biological cybernetics 17 (1975), S. 129-135 
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract Linear frequency domain analysis of subthreshold current flow and cell to cell action potential timing studies have been used to examine the dynamic electrical behaviour of the electrotonic junction between Retzius cells in the leech. Both types of electrical transmission may be completely explained if the junction is modelled by a single resistive element. The possible role of axonal cable properties in the apparent junctional characteristics were considered, but the form of the predicted frequency response functions for such cables make it unlikely that they are involved in the observed electrical behaviour.
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  • 3
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    Biological cybernetics 17 (1975), S. 137-144 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract The behaviour of similar coupled non-linear oscillators of the type $$\dot x$$ =f(x, y, µ $$\dot y$$ =g(x, y, µ is to be investigated. The oscillators are assumed to be coupled by diffusion gradients. If some conditions on the magnitude of the diffusion coefficients are satisfied, it is proved that: 1) if the oscillators have the same period (identical value of the parameter μ) and different phases before coupling, after coupling they tend to synchronize the phases; 2) if the periods of the oscillators are not too different (in terms of the values of the parameter μ) before coupling, after coupling they tend to oscillate with the same period. It is suggested the possible role of diffusion as a synchronizing mechanism in some biological phenomena.
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  • 4
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    Biological cybernetics 17 (1975), S. 145-156 
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract We first show how to formalize environments in which situations can be changed only by action, and in which a situation should be brought about that is acceptable. This leads us to the concept of a problem constellation that consists of an action system, a reaction system, a judgement system, and an initial situation. We then present a cybernetical system which we call Robbimat. It is capable of testing problem constellations, of modifying them according to the results of the tests, and of judging the final constellation. We designed Robbimat in order to have favourable problem constellations automatically generated. As an example we discuss a simple game. It constitutes a win-loss-environment the structure of which can be deduced from the payoff function. We show how Robbimat analyses such an environment, and utilizing its structure classifies the initial situation.
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  • 5
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    Biological cybernetics 17 (1975), S. 183-193 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract This paper deals with information transfer from the environment and “self”-organization in open, nonlinear systems far from thermodynamic equilibrium — in the presence of either non-stationary phase jitter noise, or amplitude stationary noise. By “self”-organization we mean here the progressive formation within the system of sequential, ordered (coherent) relationships between appropriate dynamical variables-like for example, the phase differences between the oscillating components of the system. We take up (in Section II) the classical Laser as a specific example and examine in detail the influence of phase jitter noise in the mode (phase) locking process. We find—as expected—that phase fluctuations in the cavity cause degradation of the coherent behaviour (i.e. increase the entropy) of the system — which, however, levels off, or saturates with time. Further (in Section III) we examine systems where the number of self-sustained oscillating components may vary with time in such a way that the maximum entropy of the system increases faster than the overall instantaneous entropy. We put forth the hypothesis that in such cases — because of the increase of the redundancy — the system gets organized not just in spite of, but merely because of the presence of Noise. Possible applications in biological systems (especially concerning a model of cerebral organization) are briefly discussed. It is understood here, that the system has to display some preliminary dynamical structure before the organizing procedure takes over. What happens afterwards is the subject of this paper.
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  • 6
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    Biological cybernetics 18 (1975), S. 15-18 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract In 1969 Campbell et al. reported five spatial frequency response curves. Here a good curve fit is found to three of them based on a model of lateral inhibition from four off-centered geniculate cells whose receptive fields were at a common distance from that of an on-centered geniculate cell. The distance was 0.5° when the experimental curve was fitted by a difference of two Gaussians and 1.4° when fitted by a difference of two Bessel Functions. Of the remaining two response curves one was best fitted assuming no lateral inhibition and the other remains to be fitted.
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  • 7
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    Biological cybernetics 18 (1975), S. 19-29 
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract A model of neural network extracting binocular parallax is proposed. It is a multilayered network composed of analog threshold elements. Three types of binocular neurons are included in this model. They are binocular simple neurons, binocular gate neurons and binocular depth neurons. The final layers of this model consist of elements which correspond to the binocular depth neurons. The performance of the model has been simulated on a digital computer. The results of the computer simulation show that every element of this model acts like neurons found in cat's and monkey's visual system and this model extracts binocular parallax caused by simple line components satisfactorily.
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  • 8
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    Biological cybernetics 18 (1975), S. 49-60 
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract Current clamp data of the squid axon indicate that there is a qualitative change in the adaptive response as the magnitude of the current step is increased. Large stimulus currents have a strong inhibitory effect on spike generation and on active responses in general. Such currents always lead to only one action-potential and to the elimination of post-spike subthreshold oscillation. In view of a direct connection between stimulus current and potassium current I K, the potassium channel of the Hodgkin-Huxley model is reinterpreted in a natural way such that the K+ conductance is directly dependent on I K in addition to a voltage dependence. The I-Kdependence seems to dominate whenever the stimulus current is greater than approximately 35 μA/cm2. For current ramps, and large current steps, such a current formulation leads to good agreement with the data.
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  • 9
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    Biological cybernetics 18 (1975), S. 69-80 
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract In a preceding paper, Poggio and Reichardt (1973a), a phenomenological theory describing the visual orientation behaviour of fixed flying flies (Musca domestica) towards elementary patterns was presented. Some of the problems raised in this first paper are treated here in more detail. The mapping between the position dependent torque distribution — D(ψ) characteristics — associated with a given pattern and the stationary orientation distribution p(ψ), is studied taking into account that the fluctuation process (generated by the fly) is coloured gaussian noise. Under certain critical conditions this may lead to an “early symmetry breaking” in the mean values of the p(ψ) distribution. The validity of the “superposition principle” has also been examined. Although shift and superposition give the main qualitative features of the “attractiveness profile” D(ψ), associated with a 2-stripe pattern, superposition does not hold quantitatively for stripe separations up to about 80°. Evidence is presented suggesting that such an effect is due to inhibitory interactions between input channels of the fly's eye. Implications of this finding with respect to the problem of spontaneous pattern preference are also discussed.
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  • 10
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    Biological cybernetics 18 (1975), S. 137-153 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Zusammenfassung Ausgehend von detaillierten Vorstellungen zur Kinetik der myofibrillären ATPase und der Ankopplung der Enzymwirkung an die Mechanik der Myosinbrückenzyklen wird ein quantitatives chemo-mechanisches Modell für den fibrillären Insektenflugmuskel vorgeschlagen. Mit der Wahl eines einheitlichen Parametersatzes gelingt es praktisch alle bisher in der Literatur beschriebenen Resultate zu simulieren. Dazu gehören der verzögerte Spannungsaufbau bei rechteckförmigen und sinusoidalen Längenänderungen, charakteristische Nichtlinearitäten in den oszillatorischen Längen-Spannungsschleifen, die Aktivierung der ATPase Aktivität durch statische und dynamische Dehnung, der proportionale Anstieg des Wirkungsgrades der chemo-mechanischen Kopplung mit der Effektivgeschwindigkeit bei der Oszillation, sowie die Existenz von Autooszillationen. In Übereinstimmung mit strukturellen und biochemischen Befunden erlaubt das Modell eine molekulare Interpretation des Aktivierungsmechanismus über ein Zusammenwirken von Ca2+-Konzentration und Rückkopplung der Muskelkraft sowohl bei der Aktivierung des Aktinfilaments (über Verschiebungen in den Tropomyosinmolekülen) wie auch am Ca2+-abhängigen Regulationssystem des Myosins im fibrillären Flugmuskel. Damit gelingt es die Ca2+-Abhängigkeiten der ATPase Aktivität, der Frequenzverscheibung auf der mechanischen Ortskurve, sowie der Zeitkonstanten bei sprungförmigen Längen-und Spannungsänderungen vorauszusagen. Die in Röntgenanalysen gefundene Abnahme der Zahl der am Aktinfilament angepackten Myosinbrücken nach höheren Oszillationsgeschwindigkeiten hin wird durch die Annahme einer geschwindigkeitsabhängigen Ratenkonstante für das Loslassen der Myosinbrücken im Modell realisiert. Zusätzlich wird es möglich, für die bisherige Modellbidung am Wirbeltiermuskel wichtige Experimente, wie das Verhalten des Systems bei kleinen, sehr schnellen Wegänderungen (Huxley und Simmons, 1971) oder bei sprungförmiger Entlastung des Muskels (Civan und Podolsky, 1966) modellmäßig nachzuempfinden. Die Modellsimulationen deuten auf eine Gleichartigkeit des Kontraktionsmechanismus im Skelettmuskel der Wirbeltiere und im Insektenflugmuskel.
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  • 11
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    Biological cybernetics 19 (1975), S. 1-18 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract Passive modification of the strength of synaptic junctions that results in the construction of internal mappings with some of the properties of memory is shown to lead to the development of Hubel-Wiesel type feature detectors in visual cortex. With such synaptic modification a cortical cell can become committed to an arbitrary but repeated external pattern, and thus fire every time the pattern is presented even if that cell has no genetic pre-disposition to respond to the particular pattern. The additional assumption of lateral inhibition between cortical cells severely limits the number of cells which respond to one pattern as well as the number of patterns that are picked up by a cell. The introduction of a simple neural mapping from the visual field to the lateral geniculate leads to an interaction between patterns which, combined with our assumptions above, seems to lead to a progression of patterns from column to column of the type observed by Hubel and Wiesel in monkey.
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  • 12
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    Biological cybernetics 19 (1975), S. 147-158 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract The functional properties of retinula cells of the fly Calliphora crythrocephala (wild type) have been determined through the application of nonlinear identification theory using white-gaussian stimulus functions. These results are also compared to similiar recordings of lamina cells. The accuracy of the resulting models is shown by the fact that those obtained from just 30 sec tests predict the actual total responses to the white-gaussian stimuli with a mean square error of about 5% and for a 2200 sec test the error is reduced to 2%. It has been shown that the second order kernels define all of the nonlinear properties and are the primary terms in the models for describing the variations in functional responses with illumination level, changing adaptation conditions and even variations in the conditions of the intracellular preparations.
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  • 13
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    Biological cybernetics 20 (1975), S. 51-59 
    ISSN: 1432-0770
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    Topics: Biology , Computer Science , Physics
    Notes: Abstract We analyse a model for neural activity including forgetting and self-inhibition processes. The analysis concerns the static and dynamic performances. The theoretical predictions are compared with previously published experimental data on the neural encoding in the ommatidia of the Limulus lateral eye. We conclude that the analysed model could reproduce the experimental behaviour, but different models cannot be excluded. Moreover an independent normalisation factor has to be assumed to account for the experimental gains.
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  • 14
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    Notes: Abstract The transformation of spatial patterns and their storage in short term memory by shunting neural networks are studied herein. Various mechanisms are described for real-time regulation of the amount of contrast with which a pattern will be stored. Parametric studies are described for the amount of contrast in the network responses to patterns presented at variable background or overall activity levels. Mechanisms for removing spurious peak splits and other disinhibitory responses are described. Furman's (1965) results on processing of patterns by shunting networks are generalized and reanalysed. Periodic responses (stable and unstable) corresponding to the time scale of slow cortical waves can be generated if a tonic input is set between two threshold activity levels. Their frequency as a function of tonic input size is unimodal. Order-preserving limit cycles are never found in STM; hence sustained slow oscillations as a mechanism for storing a pattern in STM are ruled out in favor of steady states (i.e., fast oscillations) with spatially graded activity levels. Such slow oscillations can, nonetheless, continuously retune the network's responsiveness to the patterns that perturb it.
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  • 15
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    Biological cybernetics 20 (1975), S. 137-143 
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    Notes: Abstract A package of methods and an experimental set-up for the analysis of dynamics of electrical signals from the brain. The methods described and discussed in this study allow detailed and multipurpose analysis of brain potentials both in the time and frequency domains. Special emphasis is given to a new computer-method introduced in this study: A posteriori selective averaging. The selective averaging method is compared with Wiener Filter Estimation of Evoked Potentials.
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    Biological cybernetics 20 (1975), S. 171-173 
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    Notes: Abstract The theoretical significance of the operation of convolution in processing signals in the nervous system is discussed. For a neural chain of perfect-integrator model neurones, information in a spike train is preserved when the p.d.f. is stable under convolution with itself. A hypothesis is proposed that stable interspike interval distributions characterize a simple information transmission pathway, and that non-stable interval distributions suggest more complex information processing functions.
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  • 17
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    Biological cybernetics 20 (1975), S. 145-160 
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    Notes: Abstract In this study, the simultaneously recorded and selectively averaged evoked potentials in some of the structures of the auditory pathway (acoustical cortex, medial geniculate nucleus, inferior colliculus), in the reticular formation and the hippocampus of the awake cat as well as the simultaneous amplitude frequency characteristics of these structures are given. The power spectral density functions computed from the simultaneously recorded spontaneous activity of these structures are also presented. Using these results, the following analyses are accomplished: (1) determination of the dynamics of potentials simultaneously obtained from various brain structures, in order to evaluate the common features of their system characteristics; (2) determination of the relationship (or interactions) between rhythmic activity and evoked potentials of the brain, and (3) elaboration of a working hypothesis for the dynamics of potentials of the brain. Some suggestions and comments are also made for investigators working toward theories or dynamic models of signal transmission in the brain.
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  • 18
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    Biological cybernetics 21 (1976), S. 85-95 
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    Notes: Abstract A demonstration is given that an orthogonalizing filter for patterns is formed adaptively and very rapidly in a network of neuron-like elements with internal feedback connections. It is here assumed that the feedback gain is variable, and proportional to the correlation matrix of the output pattern vectors. The time-dependent signal transfer properties of the complete system are described by a system matrix which satisfies a matrix Bernoulli differential equation; solutions of this equation are outlined. The asymptotic value of the system matrix is shown to correspond to the orthogonal projection operator on the space that is complementary to the space spanned by all of the earlier input pattern vectors. Such a system then acts as a filter, which optimally extracts the amount that is “new” in an input pattern with respect to all “old” patterns. It also has features that are directly attributable to a distributed associative memory that is optimally selective.
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  • 19
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    Biological cybernetics 21 (1976), S. 1-7 
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    Notes: Abstract The response of a space-clamped patch of Hodgkin-Huxley membrane to an applied current density ofA cos(2πft)+BμA/cm2 is computed for frequencies from 5 to 250 Hz. The train of action potentials generated is phase-locked to the driving cycle,N action potentials occurring at fixed phases inM cycles. For frequencies whereN/M is a simple ratio a describing function for the membrane is computed. The phase-locked behaviour and describing functions are similar to those obtained for a simple leaky integrator neurone model.
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  • 20
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    Biological cybernetics 21 (1976), S. 29-35 
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    Notes: Abstract If a binocular observer looks at surfaces, the disparity is a continuous vector field defined on the manifold of cyclopean visual directions. We derive this field for the general case that the observer is presented with a curved surface and fixates an arbitrary point. We expand the disparity field in the neighbourhood of a visual direction. The first order approximation can be decomposed into congruences, similarities and deformations. The deformation component is described by the traceless part of the symmetric part of the gradient of the disparity. The deformation component carries all information concerning the slant of a surface element that is contained in the disparity field itself; it is invariant for changes of fixation, differential cyclotorsion and uniform aniseikonia. The deformation component can be found from a comparison of the orientation of surface details in the left and right retinal images. The theory provides a geometric explanation of the percepts obtained with uniform and oblique meridional aniseikonia. We utilize the geometric theory to construct a mechanistic model of stereopsis that obviates the need for internal zooming mechanisms, but nevertheless is insensitive to differential cyclotorsion or uniform aniseikonia.
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  • 21
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    Biological cybernetics 21 (1976), S. 53-60 
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    Notes: Abstract A model is proposed for the frequency of firing of the chelonian muscle spindle in response to mechanical stretches. First an attempt is made to fit the response to a first and second order model and, after incorporating the encoding properties of the spindle, a simulation is obtained which predicts in terms of the instantaneous frequency of firing the response of the spindle to ramp and sinusoidal mechanical stretches.
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    Biological cybernetics 21 (1976), S. 97-101 
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    Notes: Abstract The interspike interval distribution of neuronal firing is analyzed by a model that assumes unit effect EPSP's lasting an exponential length of time. The model allows a general interarrival distribution; this contrasts with the numerous models requiring Poisson arrivals. The Laplace transform of the time to firing, modelled as the first passage time to a fixed arbitrary threshold level, is found. Comparisons are made for exponential and regular interarrivals using the first two moments of the time to firing. Surprisingly, the mean and variance of the time to reach any threshold level greater than one is greater for regular arrivals for any ratio of mean interarrival intervals to mean EPSP duration greater than 0.6.
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    Biological cybernetics 21 (1976), S. 139-144 
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    Notes: Abstract Reverberating neural activity is strictly defined and examined in continuous and discrete neuronal spaces with homogeneous structure. Reverberations start with a specific population of firing neurons called the initial excitation and spread out in waves of firing and refractory bands of neurons toward the periphery. The necessary and sufficient conditions for having reverberations are obtained for continuous space and discrete one-dimensional space. The excitation fronts of reverberating waves have stable shapes which depend only upon the structure of the neuronal space. The reverberatory processes in high-threshold discrete neuronal spaces show strongly nonlinear properties. Relation between reverberations and nervous functions is discussed.
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    Biological cybernetics 22 (1976), S. 21-31 
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    Notes: Abstract Discharges in cochlear nerve fibers evoked by low frequency phase-locked sinusoidal acoustic stimuli are synchronized to the stimulus waveform. Excitation and suppression regions of single units were explored using a stimulus composed of either a fixed intensity test tone at the characteristic frequency, a variable intensity interfering tone with a simple integer frequency relation to the characteristic frequency, or both. Compound period histograms were constructed from period histograms in response to normal and reversed polarity stimuli. Discharge patterns were characterized by Fourier components of the histogram envelopes. The two stimulus frequencies constituted the principal harmonics in the histogram envelopes and their combination accounted for observed rate changes. Suppression of the test tone harmonic as a function of interfering tone intensity was always seen; rate suppression was not. The harmonic was typically suppressed by 20–30 dB compared to the value for the test tone alone and often reached the 40–60 dB resolution limit of the experiment. Suppression plots were nearly linear on a power scale with an average slope of-0.8. The onset of suppression occurred for an interfering tone 9 dB greater on average than the test tone intensity. Information transfer through the peripheral system was described by the ratio of the principal harmonic amplitudes versus the ratio of the intensities of the two stimulus tones. These plots were nearly linear on a power scale with an average slope of 0.9. Neither the onset of suppression nor the slopes of the harmonic plots displayed strong dependence on characteristic frequency or interfering tone frequency. These features of harmonic behavior, however, are closely related to system nonlinearity. Comparison of measured harmonics to the predictions of two phenomenological models suggest the presence of complex nonlinear transformations in the peripheral auditory system.
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    Biological cybernetics 22 (1976), S. 203-211 
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    Notes: Abstract The response of many neurons in the nervous system is a nonlinear function of membrane potential. Nevertheless, if the membrane potentials are normally distributed then their covariance satisfies a linear equation. This suggests that information in the nervous system may be processed by correlations between membrane potentials, a hypothesis which is subject to direct experimental test. The acquisition, storage, and retrieval of information in the form of correlations is consistent with present knowledge of human information processing.
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  • 26
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    Notes: Abstract The general properties of the excitable membrane on molluscan pacemaker neurons can be described on the basis of a fair amount of experimental evidence available in the literature. The neuronal membrane exhibits under voltage clamp an initial inward current carried by both Na+ and Ca2+ ions, the time- and voltage-dependent characteristics of which are similar to that of other excitable structures. The conductance mechanism for the two ion species and the transport kinetics appear to be closely similar. The time course and amplitude of the delayed outward current carried by K+ ions shows a marked dependence on the membrane potential. Characteristic for the molluscan neurons is the existence of an additional fast transient outward current which is only activated by hyperpolarizing shifts from the membrane potential. A regular beating discharge over a wide range of frequencies can be predicted by making the assumption of a metabolically controlled driving of the Na+ conductance. Bursting pacemaker characteristics can be correctly simulated by the model if sinusoidal variations of an additional Na+ and Ca2+ conductances ≈g Na and ≈g Ca, and periodic variations of the K+ conductance ≈g K, governed by the known operation of a metabolic substrate cycle are introduced. The close approximation of experimentally observed impulse bursts requires that the actual inpulse-frequency and the amplitude of the after-spike hyperpolarization are determined by the temporal pattern of ≈g Na, while the spike amplitude is controlled by ≈g Na which (although of similar time course) is lagging in phase behing ≈g Na. The periodic changes in additional K+ conductance ≈g K, are responsible for burst termination and the changes in inter-burst interval, to the effect that spike doublets, triplets and multi-spike bursts can be simulated by a suitable choice for the time characteristics of ≈g K. The model makes use of the finding that the Ca2+ inflow associated with a spike discharge actually activates ≈g K, so that large postburst hyperpolarizations can be obtained in high-frequency bursts.
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    Biological cybernetics 23 (1976), S. 49-60 
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    Notes: Abstract Based on the available data for masking period patterns (MPP) produced by sinusoidal maskers of different frequencies and levels, a model is proposed which not only allows the peculiar shapes of the MPP of complex maskers to be understood, but also allows the MPP to tbe calculated. Several approximations have been made to describe the linear and nonlinear behaviour of the networks in the model. The agreement between the measured and calculated data is not exact, but in view of the incertainties of the physical parameters of the masker it is quite satisfactory.
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    Biological cybernetics 23 (1976), S. 73-82 
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    Notes: Abstract A simulated neuron was constructed in which effects on spike discharge of altering certain fundamental biophysical parameters could be studied. The simulation was performed by use of a digital computer. The simulation was tested by comparing performance of the simulated neuron with that of actual neurons. Rates and patterns of spike discharge were achieved for the simulated neuron that were comparable to those recorded from units in the motor cortex of awake cats. Altering biophysical parameters such as firing threshold, levels of synaptic input or rates of transverse and longitudinal current spread produced appropriate alterations of discharge rate. On the above basis it was possible to investigate interrelated effects on spike discharge of changing levels of synaptic input and rates of current spread within the simulated neuron. With low rates of longitudinal current spread, graded levels of synaptic input produced correspondingly graded levels of ouput discharge. With high rates of longitudinal current spread, the transfer properties of the neuron were markedly altered. The neuron became a bistable operator where synaptic inputs above a certain level were enhanced and all those below were suppressed. A linear relationship was found to exist between firing threshold and the level of synaptic input required to reach the transition from quiescence to near-tetanic rates of discharge. Alterations in behavior are increasingly thought to be subserved by changes in the efficacy of synaptic transmission or in the post-synaptic intergrative propeties of neurons. The results of our investigations describe the interplay between those two processes.
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    Biological cybernetics 23 (1976), S. 121-134 
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    Notes: Abstract This paper analyses a model for the parallel development and adult coding of neural feature detectors. The model was introduced in Grossberg (1976). We show how experience can retune feature detectors to respond to a prescribed convex set of spatial patterns. In particular, the detectors automatically respond to average features chosen from the set even if the average features have never been experienced. Using this procedure, any set of arbitrary spatial patterns can be recoded, or transformed, into any other spatial patterns (universal recoding), if there are sufficiently many cells in the network's cortex. The network is built from short term memory (STM) and long term memory (LTM) mechanisms, including mechanisms of adaptation, filtering, contrast enhancement, tuning, and nonspecific arousal. These mechanisms capture some experimental properties of plasticity in the kitten visual cortex. The model also suggests a classification of adult feature detector properties in terms of a small number of functional principles. In particular, experiments on retinal dynamics, including amarcrine cell function, are suggested.
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    Biological cybernetics 23 (1976), S. 149-156 
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    Notes: Abstract We applied the Wiener theory to analyse receptive field responses of L-cells in the carp and studied some dynamic properties of the receptive field of L-cells for monochromatic light stimuli. The L-cells were stimulated by each monochromatic light modulated in white-noise fashion. They responded almost linearly to all the monochromatic light stimuli. The impulse responses of the L-cells became larger in amplitude and faster in latency, peak response time, and repolarising phase as a spot of monochromatic light was enlarged. The L-cells seem to respond like a lowpass filter and the cutoff frequency of their gain characteristics increases with the enlargement of the monochromatic light spot. The relation between shift of cutoff frequency and spot diameter was monotonic increasing for each monochromatic light.
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    Biological cybernetics 21 (1976), S. 107-111 
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    Notes: Abstract Comparison of the discharge patterns of soleus motor units with associated changes in force exerted by foot during quiet stance have already demonstrated the following facts. In the initial stage of standing, the motor units exhibited stationary and asynchronous discharges. The force showed a sporadic presence of the high frequency oscillation in the 8–10 Hz band. After five to ten minutes of standing, the firing rate of individual motor unit discharges increased to about 10 spikes/sec and discharges of each motor unit were synchronized and phase-locked to each of the accompanying force oscillation. During this transitional stage, the discharges of motor units were characterized by spike dropouts from an otherwise regular spike train. To simulate the changes in the discharge characteristics, we have proposed a parallel feedback model of the stretch reflex arc. This was made of multiple α-motoneurons, motor units and muscle spindles. And motor units interact each other through group Ia afferent signals. As a result of simulation, motor units were found to exhibit stationary and asynchronous discharges when feedback gain was kept small. With an increase of feedback gain, the firing rate of individual motor units increased and finally the discharges of them were synchronized. During this transitional stage, the spike dropouts were observed in accordance with the experimental results. The neuronal mechanism of synchronization may partly be explained by the interactions of motor neurons through the above stated parallel feedback system.
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    Biological cybernetics 21 (1976), S. 227-236 
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    Notes: Abstract The probability of the joint occurrence of two statistically independent events is the product of the probabilities of the individual events. This fact is used to show that a neuron which detects coincident arrivals of spikes from two input neurons can function as a multiplier, i.e. its average output spike frequency is proportional to the product of the average input spike frequencies. The theoretical analysis is checked in two ways: (a) Computer simulations confirm the derived expressions for the output frequency and show that increasing the jitter in the input spike trains improves the operation of the multiplier by making the output spike train more regular (b) Experimentally recorded spike trains are used to demonstrate that the type and amount of jitter present in real spike trains is adequate for satisfactory operation of the proposed scheme for multiplication. The operating characteristics of the proposed multiplier make it an attractive candidate for the multiplicative mechanism that is involved in the optomotor response of insects.
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    Biological cybernetics 22 (1976), S. 39-48 
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    Notes: Abstract Vestibular habituation was investigated in 6 adult cats submitted to repetitive alternating velocity steps (160°/sec). The progressive change of the nystagmic response was examined by constructing the diagrams of the slow cumulative eye position and slow phase eye velocity; then it was quantified by evaluating some characteristic parameters of these diagrams. Both acquisition during the series of stimulations delivered in the same day and retention after a rest period of one or four days were clearly observed. The experimental results were explained in terms of a mathematical model of the vestibulo-cerebellar interaction. The validity of the model was assessed by the results of simulation experiments.
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    Biological cybernetics 24 (1976), S. 111-119 
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    Notes: Abstract Nonlinear systems which have finite memories and are time invariant can be completely described by the Wiener functional expansion, in which a series of multidimensional kernels provide a polynomial approximation to the nonlinear behaviour. The kernels give a best fitting estimation to the total system behaviour in the least mean square sense and can therefore be used to describe systems in which the nonlinearities include discontinuous functions. A modification of the Wiener method described by Lee and Schetzen, which uses kernels defined in terms of cross correlation functions, has been used in most practical attempts to analyse nonlinear systems, but we have previously described how the cross correlations may be replaced with complex multiplications in the frequency domain. The speed of domain translation offered by the fast Fourier transform makes this method more efficient than time domain estimation. In this paper the practical implementation of the technique on a medium sized digital computer is described for nonlinear systems whose outputs are continuous or pulsatile signals. This description should be adequate to allow others to implement the analysis scheme. The technique is well suited to the analysis of nonlinear biological systems, particularly those encountered in neurophysiology, because of its generality, ability to deal with hard nonlinearities and ease of use with systems having pulsatile outputs.
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    Biological cybernetics 24 (1976), S. 59-59 
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    Biological cybernetics 24 (1976), S. 67-74 
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    Notes: Abstract The slowly adapting stretch receptor of the crayfish is inhibited via the large accessory neuron both by reflex activation of this inhibitory interneuron from the stretch receptor itself (autogenic inhibition) and by activation of the interneuron from stretch receptors in other abdominal segments (neighbourinhibition). Neighbour-inhibition increases proportionally with the increase in impulse frequency in the large accessory neuron produced by activity in neighbouring receptors and largely independently of the level of excitation in the stretch receptor itself. A simple model based on intracellular recordings from the receptor neuron predicts this behaviour fairly accurately. In this model each receptor impulse is followed by an IPSP after a delay proportional to the uninhibited interspike interval of the receptor (autogenic inhibition). The other IPSP's arrive randomly distributed in time (neighbour-inhibition). An alternative model in which all IPSP's arrive randomly produces similar results. This latter model can be modified to fit other neuronal systems.
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    Biological cybernetics 24 (1976), S. 85-101 
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    Notes: Abstract In the theoretical part of the present work the input-output relation for a multi-input system is developed into a functional power series. This is formally equivalent to a decomposition of the system into a sum of all possible combinations of 1-, 2-, 3-... input subsystems. The average response of the system to a uniformly moving patern is known to be a Fourier series with respect to spatial frequency. The coefficients of the series are linear combinations of the “weights” by which different subsystems contribute to the total reaction. If a system can be shown to have essential nonlinearities of no higher than second order it is possible to calculate, from a Fourier analysis of the average movement response, the “weight” by which the nonlinear interaction between any two input elements contributes to the total reaction. This interaction is termed “elementary movement detector”. By the analysis presented here the arrangement of the elementary movement detectors may be determined for a two-dimensional array of input elements and the strength of their contributions to the total movement reaction may be calculated. Special experimental methods have been developed which allow one to apply this analysis to the visual system of the fruitfly Drosophila. The preliminary data presented show that the direction sensitive optomotor response of Drosophila can be attributed predominantly to the contributions from two “elementary movement detectors” which interconnect neighbouring visual elements. The detectors are oriented in the hexagonal array of the compound eye at +30° and at-30° with respect to the horizontal line of symmetry. A weak contribution from a detector between neighbouring elements along the horizontal line of symmetry is suggested by the present data. In the course of the analysis the contrast transfer properties of the compound eye are characterized.
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    Biological cybernetics 22 (1976), S. 181-188 
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    Notes: Abstract The technique of subthreshold addition of sinusoidal gratings was used to analyse the visual system of man during the perception of edges, lines, and bars. The experimentally obtained sensitivity function varies in close relationship to the test pattern, and can be factorized into the conjugate complex spectrum of the test pattern at threshold and a pattern-invariant function of spatial frequency. Interpreting the sensitivity function as transfer function, which is possible under certain conditions, we can describe the visual system as a matched filter which extracts an input signal contaminated with noise of specified spectral energy density. Questions discussed refer to the spatial operations occurring in matched filters, the relationship between the modulation transfer function for sine-wave gratings and the pattern-invariant transfer component, the exact determination of elements within the theoretical concept, and the realization of matched filters by the nervous system.
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    Biological cybernetics 22 (1976), S. 235-238 
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    Notes: Abstract The stability of the EEG during eyes closed and eyes open were tested by the statistical method. The EEG records of 25 s were segmented into ten 2.5 s intervals for each state. The mean and variance and power spectra (2.5 to 17.5 Hz at resolution of 2.5 Hz) were calculated for each segment of 2.5 s interval. Thus, the sequences of mean values, variances and power at each frequency component were obtained for 25 s epoch. The stationarity of these sequences were tested by the run test and the trend test. The stationarity of mean, variance and power spectra were not rejected for 25 s records by any of two tests. In the records of 50 s, about 10–20% of records tested were rejected. The nonstationarity of the EEG appeared for 50 s records. This means that the EEG during period the eyes were closed, or opened can be regarded as stationary over time period as long as at least 25 s.
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    Biological cybernetics 24 (1976), S. 181-198 
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    Description / Table of Contents: Zusammenfassung Assoziative Informationsspeicherung und assoziativer Informationsaufruf stellen ein Prinzip der Funktion des menschlichen Gehirns dar. Auf der UNIVAC 1106 wurde ein Neuronennetzwerk aus 100 erregenden Bausteinen simuliert. Die Bausteine bestanden aus Verzögerungsglied, Kennlinienglied mit Schwellenwert und PD-Glied; sie entsprachen in ihrem Verhalten weitgehend dem von realen Neuronen mit verzögertem Anstieg und Adaptation des Ausgangssignals. Die Systemerregung wurde durch einen hemmenden Baustein konstant gehalten. Information wurde als Muster erregter Einzelbausteine codiert. Die Speicherung der Muster und ebenso der zeitlichen Reihenfolge der Muster erfolgte durch Kopplungskoeffizienten (Maß für die Signaldurchlässigkeit der Verbindungen zwischen den erregenden Bausteinen).—In stark vereinfachter Weise vermag das beschriebene System Leistungen des menschlichen Gehirns nachzuahmen: parallele Assoziation (vollständiger Aufruf eines Musters durch Eingabe eines Teils des Musters), serielle Assoziation (Aufruf einer zeitlichen Mustersequenz durch Eingabe des Anfangsmusters), Auswahl zwischen beiden Assoziationsarten durch ein Steuersignal, Zuordnung eines unbekannten (nicht gespeicherten) Musters, Zuordnung von Mustern aus zwei Systembereichen, Assoziation einer wahrscheinlicheren Musterfolge, Störung des Assoziationsvorganges “Eselsbrücke”, “Abstraktion des Gemeinsamen”, “Umlernen” und “produktiver Einfall”.—Das Prinzip der wechselseitigen bedingten Verknüpfung kann als Hypothese für den Lernvorgang d.h. die Abbildung von Informationsmustern bzw. zeitlichen Mustersequenzen durch Kopplungskoeffizienten angesehen werden.—Einzelbaustein, Systemstruktur und Funktionsprinzipien des beschriebenen Systems werden mit Strukturprinzipien des Gehirns verglichen.
    Notes: Abstract Associative information storage and associative information recall are fundamental principles of the human brain. A neuron network consisting of 100 excitatory elements was simulated on the UNIVAC 1106. The network elements consisted of a delay element, a characteristic with a threshold value and a PD-element. The element's behaviour was to a great extent analogues to that of real neurons with delayed increase and adaptation of the output signal. The total excitation of the system was controlled by an inhibitory component. Information was coded as a pattern of excitated elements. The information patterns and also the temporal sequence of patterns were stored in the coupling coefficients (measure of the signal transfer between the excitatory elements).—In a very simplified manner the system described above is able to imitate effects of the human brain, including parallel association (complete recall of the stored information pattern when a part of the pattern is offered at the system's input), serial association (recall of a temporal sequence of information patterns by input of the first pattern of the sequence), selection between the two association modes by means of an external signal, classification of an unknown (not stored) information pattern, coordination of patterns from two fields of the system, association of a more probable pattern sequence, disturbance of the association process, “memory aids”, “abstraction of common characteristics”, “reversal learning” and “productive ideas”.—The principle of the mutually conditioned connection may be regarded as a hypothesis for the learning act, that is for the representation of information patterns or temporal sequences of patterns through coupling coefficients.—The network elements, the structure and the function of the system are compared with structure and principles of the brain.
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    Biological cybernetics 25 (1977), S. 83-92 
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    Notes: Abstract Double-layer neural networks with mutually inhibiting interconnections are analyzed using a continuous-variable model of the neuron. The first layer consists of excitatory neurons while the second layer consists of inhibitory neurons. Both feedforward and feedback interconnections exist between the two layers. An autonomous system of nonlinear differential equations is introduced to describe the network dynamics, and the stability conditions for some classes of equilibria are investigated in detail. Several simulation results are also presented. It is shown that even those networks which are formed with rather powerless synapses are capable of carrying out input pattern sharpening, temporary information storage, and periodic signal generation.
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    Biological cybernetics 25 (1976), S. 57-60 
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    Notes: Abstract A description is given of a non-parametric training program for constructing a non-linear functional which separates two disjoint sets G and P on the surface of the unit sphere in an arbitrary ℒ2-space. It is shown that if G and P are separable in a certain general sense, then the program is finitely convergent.
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    Biological cybernetics 25 (1977), S. 93-101 
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    Notes: Abstract The mathematical system analysis of structure and function of the human thermoregulatory system is introduced and compared to simplifying modelling techniques. In an adequate treatment the principle of locally and time dependent variables and parameters is taken into account. The complete equations are given in such a way that reference signals are not necessary. The problem of the set point is discussed. On the basis of a series of preliminary simplifications first results are presented concerning system-states under open and closed loop conditions, effectiveness of the controller and dynamics of radial temperature profiles. Quasistationary and dynamical experimental results are added for comparison. The results encourage us to continue the realisation of the complete system without the preliminary simplifying assumptions.
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    Biological cybernetics 25 (1977), S. 121-130 
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    Notes: Abstract Behavioral experiments are indispensable for the analysis of biological systems for cognition and recognition. When these are carried out as detection experiments three types of description can be used for the problem of visual pattern recognition which allow conclusions to be drawn on the operating function of the system. Provided that the signals to be recognized have additive noise superimposed on them, system description is possible: 1. on the basis on the probabilities of recognition and of mix-up,—2. through the analysis of the transformation of distribution densities of the noise,—3. by means of the measurable distances of the patterns from each other in feature space.-The analysis of the distribution densities shows that the human visual system acts like a linear classifier in the classification of six geometrical patterns. The independence of the classification from intensity as well as the human reaction to alteration in the power spectrum of the noise support this result. Simulation experiments on a computer show the efficacy of various biological relevant parameters for the linear classification and suggest that a narrow band and probably feature specific filtering precedes the classification.
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    Biological cybernetics 25 (1977), S. 163-176 
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    Notes: Abstract A scheme for constructing computer models of motorneurons is presented. These model neurons display both repetitive firing and action potentials of appropriate time course; the technique combines ideas of Dodge and Cooley (1973) and Kernell and Sjöholm (1972, 1973). This scheme is used to construct models of motorneurons of different input resistance (and hence different geometry) in order to examine Kernell's (1966) observations that (a) small motorneurons require a lower injected current to begin firing than do large motorneurons; but (b) the slope of the injected current-firing rate curve is less for small neurons than large. It is concluded that this behavior is not a simple consequence of the cell geometry, but requires different time courses for a slow conductance across the cell membrane. This observation is consistent with the experimentally observed differences in time course of afterhyperpolarization between large and small cells (Eccles et al., 1958). The models are used to study input-output relations for motorneurons when the input is a steady conductance change on the cell membrane, possibly in conjunction with an injected current. Comparisons are drawn with the experimental observations of Kernell (1965a, 1965b) and Chaplain and Schaupp (1973). It is shown that the observations of Henneman et al. (1965a, 1965b) and Milner-Brown et al. (1973a, 1973b) on the order of recruitment of motorneurons under conditions of natural stimulation may be explained by the following version of the size principle: a given input to a pool of motorneurons causes equivalent conductance changes on each cell of the pool—here equivalent means the input is distributed to corresponding portions of the soma-dendritic tree and is of equal magnitude in mhos. Hypotheses are offered as to how this distribution of input may be accomplished in Nature.
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    Biological cybernetics 25 (1977), S. 205-208 
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    Notes: Abstract A signal-flow diagram of the oculomotor control system has been derived which is able to describe the three modes of its action 1) pursuit movements 2) voluntary saccadic movements and 3) the passive non-innervated state of extraocular muscles which exists during sleep. It has been taken into consideration that in the smooth pursuit system there is a neural integrator in order to bring back to zero the error between the position of the eye and an external constant reference point. [Evidence for integration in oculomotor pathways we have from experiments by Cohen and Komatsuzaki (1972) who used stimulation of the pontine reticular formation.] All this is achieved by a system of variable structure with three states. In skeletomotor systems likewise there are smooth compensatory movements and voluntary movements and a state without any innervation. Some neurological diseases can be interpreted as an impairment of switching at special spots of the signal-flow diagram or as a disconnection of signalpathways, respectively. From this can be concluded that the signal-flow diagram derived for the rather lucid oculomotor control system should be able to describe the basic function of skeletomotor control systems, too.
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    Biological cybernetics 29 (1978), S. 11-18 
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    Notes: Abstract As compared to linear systems for which well-proven standardised procedures exist, the analysis of nonlinear systems based on random signals cannot be handled in a similar manner because of the lack of a nonlinear systems theory. A number of different procedures has been published, the most popular of them based on the application of the Volterra series, but also procedures based on the optimal linearisation of a given nonlinearity. This paper applies the procedure of optimal linearisation to a system with several imputs and one output. It is possible to extend the procedure to systems having several outputs. As an example, the parameters of the coupling function of a one-stage nerve-network will be computed. The practicability of the procedure and associated errors will be investigated.
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    Biological cybernetics 29 (1978), S. 127-136 
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    Notes: Abstract The nerve cells are believed to have such ability of self-organization that, given a number of input patterns, each cell tunes itself to become responsive to only one of the patterns, or to one subset of patterns having some features in common. The detectors of patterns or pattern subsets are formed in this manner. A simple but plausible mechanism of self-organization is proposed based on the two hypotheses: 1) Synaptic modification process is non-linear, activated when the output of a cell is positive. 2) Not only excitatory but also inhibitory synapses are modifiable. A rigorous mathematical analysis is given to elucidate the characteristics of modifiable synapses to form these detectors. The present model fits well most of the experiments on the developmental plasticity of the visual cortex such as the formation of orientation detecting cells, monocular and alternate monocular deprivation in normal and abnormal environments.
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    Biological cybernetics 29 (1978), S. 159-166 
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    Notes: Abstract An analysis of the model performance under normal and abnormal conditions is presented. The effects of changes in the alveolar ventilation rate on the inspiratory time, expiratory time, pause period, respiratory frequency, tidal volume, dead space volume and change in end expiratory level are considered. Moreover, theoretical predictions of the effects of added resistance loads are given. The results are displayed for three different values of the elastance. Comparisons with available experimental data show that the model predictions are in good accordance with observations under both normal and loaded conditions. None of the previous models have been able to reproduce the overall pattern of breathing as accurately as the present hierarchical model.
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    Biological cybernetics 29 (1978), S. 187-191 
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    Notes: Abstract A first time crossing problem for Gaussian stochastic process and monotonic time curve is considered and results are discussed with application to neural modelling. Using diffusion approximation of the stochastic process, integral equation for probability density function of the first time crossing has been obtained. Exact solution of the equation is given for two kinds of stochastic processes which have correspondingly infinitesimal and infinitely large correlation time; approximation methods are constructed for processes characterized by intermediate values of this parameter.
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    Biological cybernetics 30 (1978), S. 7-13 
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    Notes: Abstract A model for transmitter kinetics has been simulated by computer for a variety of inputs and it is shown how the parameters of the system determine its properties, such as phasic or tonic behavior, sensitivity and responsiveness. The results are discussed in relation to transmitter release and transmitter-receptor interactions.
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    Biological cybernetics 30 (1978), S. 99-108 
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    Notes: Abstract The mathematical relationship describing recurrent lateral inhibition is expressed as a linear operator equation. Under quite general conditions, the operator equation is shown to have a unique nonnegative solution. It is also shown that the linear operator for recurrent impedance is representable as an integral operator and that, when in application to physiological models it is interpreted as recurrent inhibition, the corresponding linear equation assumes a form more general than the well known Hartline-Ratliff equation. Finally, we introduce a class of impedance operators based on the probabilistic theory of Markov processes, solve the corresponding linear integral equation, and apply the theoretical properties of the solution to the analysis of physiological and psychophysical phenomena.
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    Biological cybernetics 30 (1978), S. 179-185 
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    Notes: Abstract Frequency variations in the human voice result from voluntary and involuntary changes in the parameters of the vocal system. The present work deals with involuntary frequency perturbations from two theoretical aspects: 1) the influence of pitch period variations on frequency changes in the band-limited signal which results from the resonant characteristics of the vocal tract; 2) the physiological parameters of the vocal system which are potentially able to govern involuntary frequency changes. It is shown that the modulation function of the vocal-cord wave can theoretically be derived from its harmonics using FM demodulation techniques, and that higher distortion may appear at higher harmonics. It is also shown that involuntary geometrical changes of the vocal tract and its terminal impedance as well as tension and initialarea changes of the vocal cord—changes well within the physiological range—can influence frequency changes in the human voice. The present results are correlated with our reported experimental findings on involuntary voice tremor, used in psychological stress evaluation. The role of the central nervous system, and possible mechanisms for these phenomena, are discussed.
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    Biological cybernetics 30 (1978), S. 221-230 
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    Notes: Abstract In order to study the motor unit action potential a computer simulation model was developed. It is based on the superposition of single muscle fibre potentials of the fibres belonging to the motor unit. The parameters which characterize each fibre (spatial position, diameter, and a dispersion of arrival time of the potential at the electrode) are chosen from statistical distributions which can be derived from anatomical and physiological data. The electrode type, position and dimensions can be specified. Simulated motor unit action potentials are presented in the time and frequency domain. The simulation results refer to (1) the influence of the electrode position and dimensions with respect to the motor unit territory, (2) the meaning of this model for the study of pathological phenomena, (3) the variability of some parameters characterizing the motor unit, (4) the selectivity of uni- and bipolar electrodes and finally (5) the influence of the geometrical situation of the motor end-plates within the muscle, on the shape of motor unit action potentials.
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    Biological cybernetics 31 (1978), S. 15-26 
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    Notes: Abstract A model of a local neuron population is considered that contains three subsets of neurons, one main excitatory subset, an auxiliary excitatory subset and an inhibitory subset. They are connected in one positive and one negative feedback loop, each containing linear dynamic and nonlinear static elements. The network also allows for a positive linear feedback loop. The behaviour of this network is studied for sinusoidal and white noise inputs. First steady state conditions are investigated and with this as starting point the linearized network is defined and conditions for stability is discovered. With white noise as input the stable network produces rhythmic activity whose spectral properties are investigated for various input levels. With a mean input of a certain level the network becomes unstable and the characteristics of these limit cycles are investigated in terms of occurence and amplitude. An electronic model has been built to study more closely the waveforms under both stable and unstable conditions. It is shown to produce signals that resemble EEG background activity and certain types of paroxysmal activity, in particular spikes.
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    Biological cybernetics 31 (1978), S. 49-54 
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    Notes: Abstract Some tumors of hormonal organs are clinically active, while others are not. The “silent” tumors may be discovered by accident or because of effects due to their increase in size. From a simple steady state analysis of hormonal feedback systems follows that hormonal cell multiplication does not significantly influence the systems steady state behaviour (hence the clinical silence). —Exceptions to this rule occur in three situations: when the gain of the system is low; when the growth concerns cells with isolated sensor or reference functions; or because of the growth of autonomous cells. In many biological systems the dangerous situation of clamping to low levels upon sensor cell multiplication has been prevented by lumping, such as the combination of sensor and comparator functions into sensor-comparator cells.
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    Biological cybernetics 31 (1978), S. 63-70 
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    Notes: Abstract In a simulated neuron with a dendritic tree, the relative effects of active and passive dendritic membranes on transfer properties were studied. The simulations were performed by means of a digital computer. The computations calculated the changes in transmembrane voltages of many compartments over time as a function of other biophysical variables. These variables were synaptic input intensity, critical firing threshold, rate of leakage of current across the membrane, and rate of longitudinal current spread between compartments. For both passive and active dendrites, the transfer properties of the soma studied for different rates of longitudinal current spread. With low rates of current spread, graded changes in firing threshold produced correspondingly graded changes in output discharge. With high rates of current spread, the neuron became a bistable operator where spiking was enhanced if the threshold was below a certain level and suppressed if the threshold was above that level. Since alterations in firing threshold were shown to have the same effect on firing rate as alterations in synaptic input intensity, the neuron can be said to change from graded to contrast-enhancing in its response to stimuli of different intensities. The presence or absence of dendritic spiking was found to have a significant effect on the integrative properties of the simulated neuron. In particular, contrast enhancement was considerably more pronounced in neurons with passive than with active dendrites in that somatic spike rates reached a higher maximum when dendrites were passive. With active dendrites, a less intense input was needed to initiate somatic spiking than with passive dendrites because a distal dendritic spike could easily propagate by means of longitudinal current spread to the soma. Once somatic spiking was initiated, though, spike rates tended to be lower with active than with passive dendrites because the soma recovered more slowly from its post-spike refractory period if it was also influenced by refractory periods in the dendrites. The experiment of comparing neurons with active and passive dendrites was repeated at a different, higher value of synaptic input. The same differences in transfer properties between the active and passive cases emerged as before. Spiking patterns in neurons with active dendrites were also affected by the time distribution of synaptic inputs. In a previous study, inputs had been random over both space and time, varying about a predetermined mean, whereas in the present study, inputs were random over space but uniform over time. When inputs were made uniform over time, spiking became more difficult to initiate and the transition from graded to bistable response became less sharp.
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    Biological cybernetics 32 (1979), S. 1-8 
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    Notes: Abstract A neural network model is proposed to explain the development of direction selectivity of cortical cells. The model is constructed under the following three hypotheses that are very plausible from recent neurophysiological findings. (1) Direction selectivity is developed by modifiable inhibitory synapses. (2) It results not from the direct convergence of many excitatory inputs from LGN cells but from cortical neural networks. (3) Direction-selective mechanism is independent of orientation-selective mechanism.—The model was simulated on a computer for a few kinds of inhibitory connections and initial conditions. The results were consistent with neurophysiological facts not only for normal cats but for cats reared in an abnormal visual environment.
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    Biological cybernetics 32 (1979), S. 25-33 
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    Notes: Abstract The optimal linear filters derived in the preceding paper can be thoroughly evaluated using computer simulations, based on the properties of mammalian sensory and motor nerve fibres. Using reasonable values for action potential waveforms, conduction velocity and electrode noise, good separation of motor and sensory signals can be obtained. The performance of the filters is degraded by 1) increasing the electrode noise, 2) introducing dispersion in the conduction velocities, or 3) variation in the waveform of the action potentials from that used in designing the filters. However, the variations needed to seriously degrade performance are quite large compared to those which are likely to be present in mammalian nerves. Use of these filters to distinguish different classes of sensory (or motor) signals based on conduction velocity is discussed.
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    Biological cybernetics 32 (1979), S. 49-62 
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    Notes: Abstract Ontogenetic development of ocularity domains — stripes, patches and layers in cortex, colliculus superior and lateral geniculate nucleus — is the result of organization that may either be intrinsic to the postsynaptic structure or induced to it by the afferents. A specific type of axonal growth behaviour that was recently proposed as a basis for ontogenetic development of retinotopy is sufficient to account also for ocularity domains. No intrinsic organization in the postsynaptic structure is required. The latter merely serves as a propagating medium for markers carried by the presynaptic terminals. Computer simulations demonstrate the mechanism to be complete and consistent.
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    Biological cybernetics 32 (1979), S. 85-93 
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    Notes: Abstract Two models for visual pattern recognition are described; the one based on application of internal compensatory transformations to pattern representations, the other based on encoding of patterns in terms of local features and spatial relations between these local features. These transformation and relational-structure models are each endowed with the same experimentally observed invariance properties, which include independence to pattern translation and pattern jitter, and, depending on the particular versions of the models, independence to pattern reflection and inversion (180° rotation). Each model is tested by comparing the predicted recognition performance with experimentally determined recognition performance using as stimuli random-dot patterns that were variously rotated in the plane. The level of visual recognition of such patterns is known to depend strongly on rotation angle. It is shown that the relational-structure model equipped with an invariance to pattern inversion gives responses which are in close agreement with the experimental data over all pattern rotation angles. In contrast, the transformation model equipped with the same invariances gives poor agreement to the experimental data. Some implications of these results are considered.
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    Biological cybernetics 32 (1979), S. 115-123 
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    Notes: Abstract The static and dynamic characteristics of phototransduction were studied in photoreceptors of the compound eye of the fly Phormia regina (Calliphoridae) using a green light emitting diode driven by a controlled current source. The LED provides sufficiently intense light to investigate the behaviour of the receptors over about half of the dark adapted range of the response versus log intensity curve. The effects of constant adapting light intensities upon the step response and upon the frequency response and coherence functions were examined. Using both methods the effect of light adaptation upon receptor sensitivity can be closely approximated by a similar linear dependence of log sensitivity upon log adapting intensity. However, there was no reliably detectable effect of light adaptation upon the time constant of the response over the range of adapting intensities used.
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    Biological cybernetics 32 (1979), S. 187-199 
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    Notes: Abstract A theory dealing with the control of human, arbitrary movements is proposed. A schema is set up to suggest how the relevant information flows and what kind of operations affect it. A number of successive steps are distinguished in the production of a movement. It is assumed that the intended movement is carried out in the imagination, and that this imaginary movement is composed of a spatial trajectory and an intensity course, which are considered to be independent features of the intended movement. The spatial trajectory will be encoded in a special coding, which is related to the lengths of the muscles that effect the movement. From this special coding of the intended movement static and dynamic control signals can be derived. Because afferent and efferent signals are encoded in the same way in this schema, the evaluation and correction of the performed movement is quite simple. The higher levels in the control schema may function in an abstract way, i.e. the signals at these levels are barely concerned with details of the peripheral motor system. This abstract functioning of the higher levels is based on the numerous feedback mechanisms involved at all levels of control and in the peripheral motor system. Nevertheless, it is possible to incorporate specific peripheral properties in the generation of the control signals. The assumptions in this theory will be discussed and aspects of the proposed control schema will be compared with general control principles.
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    Biological cybernetics 32 (1979), S. 239-241 
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    Notes: Abstract Flies were filmed simultaneously from above and from the side. Their flight tracks were analyzed frame by frame. Male and female flies were found to chase other flies. But female chases are brief and poorly controlled as compared to male chases. Female flies use the lower frontal part of their visual field for tracking other flies. Male flies use the upper frontal part of their visual field for that purpose. Male flies are capable of controlling their forward velocity roughly proportional to the distance to their target. Implications for the function of recently found sexdimorph visual interneurones are discussed.
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    Biological cybernetics 33 (1979), S. 29-37 
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    Notes: Abstract Various observations and conjectures have been made about how we perceive three dimensional objects under monocular viewing conditions; in particular, when such objects are rotating. In this paper these proposals are reviewed and reinterpreted in the light of geometric properties of objects and their projections. Some results are also presented which support the conclusion that the visual system, in reconstructing the object from its projections, requires a fixed time to establish the objects curvature and torsion parameters. In the perceptual reconstruction process these parameters assume positive or negative values, and this equivocation is related to a form of the adjacency principle involving central projections and “perceptual geodesics’.
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    Biological cybernetics 32 (1979), S. 211-216 
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    Notes: Abstract It is argued that the internal model of any object must take the form of a function, such that for any intended action the resulting reafference is predictable. This function can be derived explicitly for the case of visual perception of rigid bodies by ambulant observers. The function depends on physical causation, not physiology; consequently, one can make a priori statements about possible internal models. A posteriori it seems likely that the orientation sensitive units described by Hubel and Wiesel constitute a physiological substrate subserving the extraction of the invariants of this function. The function is used to define a measure for the visual complexity of solid shape. Relations with Gestalt theories of perception are discussed.
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    Biological cybernetics 33 (1979), S. 1-7 
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    Notes: Abstract Sequential input adaptive system theory is applied to human eye tracking of targets with one-dimensional dimensional motion defined by sine functions, sums of sine functions and triangle functions. The sequential theory describes the mean eye tracking movements of two subjects for these inputs. The theory is predictive for irregular inputs in that when its parameters for a subject are identified and adjusted for a reference input, it then predicts mean eye tracking behavior for that subject for inputs of comparable complexity.
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    Biological cybernetics 33 (1979), S. 125-135 
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    Notes: Abstract A key problem in vision is to normalize one's lightness scale so that surface reflectances are always assigned the same gray value regardless of the illumination level. The solution requires an assessment of the relation between the strength of the illuminant and the strength of the image signal-information that is not available in the image alone. However, the level of scattered light in the optical system does provide an independent measure of the illuminant strength, and can be used to solve the lightness scale normalization problem. To do this requires a comparison between two imaging systems, each of which respond differently to the internal optical scatter. The rod and cone systems have properties that are ideally suited for such a role.
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    Biological cybernetics 33 (1979), S. 143-149 
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    Notes: Abstract The cooperative stereo algorithm of Marr and Poggio (1976) is modified empirically to provide an algorithm which exhibits many of the characteristics of the original algorithm but is more amenable to qualitative and quantitative interpretation.
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    Notes: Abstract Horizontal eye movements of the alert rhesus monkey resulting from both pseudorandom binary sequence (PRBS) and single frequency sinusoidal rotational stimulation were analyzed using a PDP 11/40 computer in order to generate gain, phase, and coherence estimates at discrete frequencies between 0.008 and 1.28 Hz. A computer simulation of vestibular induced eye movements was used to validate our analysis procedures and to determine the effects of digital noise. Frequency domain transfer functions derived from gain and phase estimates revealed that the responses to PRBS stimulation and to single frequency sinusoids were not appreciably different. PRBS testing was accomplished in approximately one third the time required for sinusoidal testing and yielded highly reproducible data. We conclude that PRBS stimulation is a reliable and efficient method for assessing linear system parameters of the horizontal vestibulo-ocular reflex. PRBS testing may be particularly and advantageous in studies of vestibulo-oculomotor plasticity in which rapid assessment of alterations in system dynamics is essential.
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    Biological cybernetics 33 (1979), S. 199-208 
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    Notes: Abstract Generally, the phase resetting experiments can be used to investigate the behaviors of the stable biological oscillators (e.g. circadian rhythms, biochemical oscillators, pacemaker neurons, bursting neurons). Winfree found that there are two types of phase transition curves (PTC) in the phase resetting experiments of biochemical oscillations. The one is the curve with an average slope of unity (Type 1) and is obtained for the small magnitude of perturbation. The other curve is that with a zero average slope (Type 0) and is obtained for the large magnitude of perturbation. Previously, we explained these results mathematically by the homotopy theory. In this paper, some properties of the human finger tapping neural network are studied psychologically using PTC on the basis of above theoretical results: assuming that an oscillatory neural network controls the human finger tapping, we performed two kinds of phase resetting experiments on the finger tapping. In the first experiment, we showed that the PTC was available to estimate the degree of functional interaction between the finger tapping neural network and that which controls another task. Three tasks (rapid key-pushing, rapid voicing and pattern discrimination) were chosen as the perturbation of the phase resetting experiment. Analyzing shapes of PTCs, it was found that the interaction with the key-pushing network was the largest, and that with the pattern recognition network was the smallest of the three. In the second experiment, we modified the first task as perturbation of the phase resetting experiments to investigate further the interactions between the left and the right hand motor systems. Consequently the following results were revealed. First, shapes of PTCs are very different according as subject's experiences of finger tapping. Second, the type of PTC for some subjects changes from Type 0 to Type 1 by learning. Third, the PTC tends to become Type 0 for shorter tapping periods. Fourth, neither changes of motor loads (the necessary force to push the key) nor an alternation of the tapping hand and the key-pushing hand affects the shape of PTCs.
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    Biological cybernetics 34 (1979), S. 31-34 
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    Notes: Abstract A formal representation of nerve spike trains in the form of a sum of rectangular functions is proposed. This formal instantaneous frequency function can be Fourier analyzed. The resulting algorithm has the useful properties of spike by spike calculations and an insensitivity to the mean (carrier) spike rate. The technique is also useful for producing a smooth (filtered) reconstruction of a spike train.
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    Biological cybernetics 34 (1979), S. 1-19 
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    Notes: Abstract This study is carried out on single (not averaged) recordings combining the spontaneous activity preceding the stimulus onset and the EP recorded upon acoustical stimulation. These recordings, which we call EEG-EPograms, are measured simultaneously from different subdural brain structures, such as the auditory cortex, medial geniculate nucleus, inferior colliculus, reticular formation and the hippocampus of awake cats. Using a combined analysis procedure (C.A.P.), the relevant frequency components of spontaneous EEG and EPs, recorded simultaneously from these brain nuclei, are analyzed according to the consistent selectivity bands depicted by the determined amplitude-frequency characteristics. These analyses provide us the following information: (1) there is an important congruency in the time courses of simultaneous response components in common frequency bands, especially in the α and β frequency ranges; (2) there exist significant coupling and synchrony between the evoked amplitude enhancements in the simultaneously recorded single response components; (3) the inter-nuclei coherency in the brain's electrical activity is enormously increased upon stimulation; (4) the evoked response magnitude can be predicted, with reasonable accuracy, from the spontaneous activity preceding the stimulation. The strong dependence of the response magnitude on the stimulus-preceding EEG is explained by means of a model network consisting of a population of relaxation oscillators, which can be brought to different states of synchrony and asynchrony. Some suggestions and comments are also made for investigators working toward theories of signal transmission in the brain.
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    Biological cybernetics 34 (1979), S. 21-30 
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    Notes: Abstract This study is carried out on single (not averaged) recordings combining the spontaneous activity preceding the stimulus onset and the EP recorded upon acoustical stimulation. These recordings, which we call EEG-EPograms, are measured simultaneously from different subdural structures, such as the auditory cortex, medial geniculate nucleus, inferior colliculus, reticular formation and the hippocampus of the cat brain during the slow wave sleep stage. Using a combined analysis procedure (C.A.P.), the relevant frequency components of spontaneous EEG and EPs, recorded simultaneously from these brain nuclei, are analyzed according to the consistent selectivity bands depicted by the determined amplitude-frequency characteristics for the SWS-stage. In parallel with the results which we obtained for the waking stage, these analyses provide also the following information: (1) there is an important congruency in the time courses of simultaneous response components in common frequency bands, especially in the alpha and beta frequency ranges; (2) there exist significant coupling and synchrony between the evoked amplitude enhancements in the simultaneously recorded single response components; (3) the inter-nuclei coherency in the brain's electrical activity is enormously increased upon stimulation; (4) the evoked response magnitude can be predicted, with reasonable accuracy, from the spontaneous activity preceding the stimulus. All these findings are discussed with reference to those obtained for the waking stage.
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    Biological cybernetics 34 (1979), S. 62-62 
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    Biological cybernetics 34 (1979), S. 107-110 
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    Notes: Abstract Angular wrist displacements in the monkey result in a short-latency (20–25 ms) “reflex” response of motor cortical cells in area 4 and separate peaks of activity in the gross EMG of the stretched muscles. Frequency domain analysis was carried out between (1) wrist position as input and motor cortical neuron response as output, (2) position as input and EMG response as output, and (3) motor cortical neuron response as input and EMG response as output. The results show that the dynamics of primary spindle afferents characterize the dynamics of the pathway to the motor cortex and that of the reflex loop. Results are discussed in terms of “long-loop” reflexes.
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    Biological cybernetics 34 (1979), S. 151-157 
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    Notes: Abstract A generalized analysis of the generator potential responses of R1-6 cells of Calliphora provides remarkable information on the visual properties for the Diptera. This shows that, although these cells have two peak response sensivities for monochromatic stimuli at 350 and 480 nm under single color stimulus conditions, and when the background illumination is either zero or in the region of 450–560 nm, the sensitivity to ultraviolet light is practically eliminated for background illumination in either the ultraviolet or the region around 600 nm or when any simultaneous dynamic stimulus in the region of 480–550 nm is also applied. These results seem somewhat perplexing to an understanding of the behavioral vision properties. It also is not consistant with the concept that the ultraviolet response is initiated by a sensitizing pigment within these cells that transfers energy to the rhodopsin-metarhodopsin process. However, it strengthens other evidence that the limited condition of ultraviolet responses comes from interaction from R7,8 cells but does not play an important behavioral role in the visual system fed from cells R1-6. As discussed in this paper, any high level pattern recognition controlling behavioral response to ultraviolet stimuli comes from the R7,8 cell system.
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    Biological cybernetics 34 (1979), S. 187-193 
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    Notes: Abstract Kinetics of the development of orientation tuning are inferred from quantitative analysis of extracellular recordings in the primary visual cortex of normally and dark reared kittens. 712 visual cells were classified in three functional groups: a) non-specific cells, and b) immature cells which are not as orientation selective as c) specific cells. Power regression and covariance analysis indicate that the “critical period” begins before 19 days and that the kinetics of the immature pool are the same in both rearing conditions. A catenary process of development of orientation selectivity is proposed, the immature compartment being a transit pool between non-specific and specific cells. Two sequential stages occur: 1) the realisation of an intrinsic programme of maturation, by which cortical specificity appears at eye opening and increases independently of visual experience 2) a phase of “epigenesis” beginning at 19 days, during which functional modification depends on visual experience.
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    Biological cybernetics 34 (1979), S. 227-232 
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    Notes: Abstract Repetitive firing of single tonic neurones is modeled to include in detail both membrane excitation kinetics and electrotonic effects due to membrane non-uniformities in the impulse encoder region. The model is evaluated dynamically and compared with similar data obtained from the crayfish stretch receptor neuron. Two dynamic techniques utilizing small amplitude sinusoidal signals are employed. One technique is used to fix the values of two parameters which relate to the electrotonic control of membrane potential in the interspike interval and to the relaxation time of the K-conductance during repetitive firing. The other technique is employed as a consistency check. The dynamics are particularly sensitive to the K-channel relaxation time in the interspike interval.
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    Biological cybernetics 34 (1979), S. 241-247 
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    Notes: Abstract In this paper the visual masking effect is interpreted on the basis of the transient characteristic in two dimensional neuronal networks. The study investigates the suitability of the effect for use as a measurement method. It is shown that the stimulus distribution in space can be scanned at different points in time and that various dynamic characteristic values of the system can be measured.
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    Biological cybernetics 31 (1978), S. 71-79 
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    Notes: Abstract Mammalian primary muscle spindle endings receive two inputs from fusimotor fibers and length changes of their extrafusal environment. Both inputs are also powerful noise sources disturbing the discharge patterns of the receptors. In this paper emphasis is laid on the extrafusal component. It is shown that extrafusal muscle activity can be viewed as a stochastic process which, acting as a common source, can correlate discharge patterns of adjacent muscle spindles. Some quantitative characteristics of these correlations and their relation to the underlying mechanical events are investigated in some detail in order to provide data for more theoretical considerations on their possible physiological importance.
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    Biological cybernetics 31 (1978), S. 97-98 
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    Notes: Abstract Recently, a model has been proposed to explain the statistics of the variability of interspike intervals of ganglion cells in the retina of goldfish under steady-state stimulation. In this note, it is shown that the dynamical behaviour of the model, both under steady-state and dynamical stimuli, is characterized by input-invariance of the output random process after a time transformation. This allows to study the signalprocessing properties of the model, and to test its applicability by a more complete experimental analysis.
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    Biological cybernetics 32 (1979), S. 77-83 
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    Notes: Abstract The perceptual difference between stimuli can be regarded as distance within the perceptual space of the bee. The author used this assumption to determine the specific distance function, on the basis of which the differences in the individual perceptual parameters constituted the perceptual difference between the complex stimulus and the reference stimulus. The perceptual differences can be deduced only indirectly from the choice frequency. Consequently, it was necessary to establish a “calibration curve”, to deduce quantitatively the perceptual difference from the choice frequency. The resulting hyperbolic curves for the parameters “brightness” and “size” were almost identical (Fig. 2). The perceptual difference between the complex stimulus and the reference stimulus is greater than one would expect in an Euclidean space. Rather it is the sum of the distances along the perceptual parameters which compose the complex stimulus (Fig. 3). Thus, the bee determines the perceived difference of composite stimuli which affect the perceptual parameters “brightness” and “size” in terms of the city-blockmetric.
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    Biological cybernetics 32 (1979), S. 107-113 
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    Notes: Abstract A computer (Fortran) model is proposed that describes the temporal and spatial coordination pattern of straight walking stick insects (Carausius morosus) for a broad speed range. It provides a stable pattern independent of the different starting positions. The model is based on six relaxation oscillators. The leading oscillator corresponds to a frontleg. Therefore the information flow runs from front to rear in contrast to earlier models (Graham, 1972; Wendler, 1968).
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  • 85
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    Notes: Abstract This paper deals with the problem of separating the spectra of signal and noise in ensembles where the signal can be considered as an invariant component and the noise as a stationary additive background. Several methods are discussed and compared on the basis of a statistical analysis of the first two moments of the estimators for signal and noise spectra. As a consequence a procedure is proposed which provides a flexible compromise between estimation accuracy and computational effort. The application of this procedure to a posteriori “Wiener” filtering is compared with a more common, but time consuming, technique.
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    Biological cybernetics 33 (1979), S. 113-120 
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    Notes: Abstract Time histograms of neural responses evoked by sinusoidal stimulation often contain a slow drifting and an irregular noise which disturb Fourier analysis of these responses. Section 2 of this paper evaluates the extent to which a linear drift influences the Fourier analysis, and develops a combined Fourier and linear regression analysis for detecting and correcting for such a linear drift. Usefulness of this correcting method is demonstrated for the time histograms of actual eye movements and Purkinje cell discharges evoked by sinusoidal rotation of rabbits in the horizontal plane. In Sect. 3, the analysis of variance is adopted for estimating the probability of the random occurrence of the response curve extracted by Fourier analysis from noise. This method proved to be useful for avoiding false judgements as to whether the response curve was meaningful, particularly when the response was small relative to the contaminating noise.
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    Biological cybernetics 33 (1979), S. 137-141 
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    Notes: Abstract The Hartline-Ratliff equation is a linear integral equation of the second kind and is employed in modeling inhibitory networks. Saturation of the inhibiting elements is commonly modeled as a function whose form is sigmoid; however, the resulting integral equation is nonlinear. Whenever the unknown function within the integral is hypothesized to be a nondecreasing nonlinear function, the Hartline-Ratliff equation becomes a nonlinear integral equation of the Hammerstein type. We present existence and uniqueness theorems for a Hammerstein equation which represents a further generalization of the Hartline-Ratliff equation.
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    Biological cybernetics 17 (1975), S. 1-6 
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    Notes: Abstract White noise modulated stimuli were employed for studying the transfer characteristics of the intracellularly measured retinula cell responses of the chalky mutant of the blowfly Calliphora erythrocephala Meig. Average depolarization levels varied between 6 and 20 mV. Response modulation depths of up to ±44 % could be achieved. The analysis of the stimulus-response relationship, which takes into account nonlinear properties of the receptor cell response, reveals, that this system is essentially linear for an intermediate frequency range between approximately 5 and 30 Hz. For higher as well as lower frequencies the system becomes increasingly nonlincar.
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    Biological cybernetics 17 (1975), S. 7-17 
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    Notes: Abstract 1. Intracellular recording were obtained from P-cells of the LGN of the cat. The impulse trains of a single presynaptic retinal ganglion cell and the postsynaptic P-cell were separated by band-pass-filtering and subsequent amplitude discrimination. 2. The rates of information and transinformation for the visual channel from the eye to a ganglion cell and to the connected P-cell were calculated. Input signals to the channel were trains of light flashes of different rate, luminance and spatial distribution. 3. Transinformation was calculated without restrictive assumptions for the code. 4. The transient behaviour of the system in response to a flash was fully considered for information calculations. Additionally, it was ensured that the state of the (adaptive) channel was considered correctly. 5. Information theory was applied in an extended way. The time courses of information transfer were calculated for various flash stimuli and compared with each other.
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    Biological cybernetics 27 (1977), S. 185-187 
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    Notes: Abstract This paper deals with the problem of self-controlling nets of Caianiello's type. These nets control themselves through their own elements.
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    Biological cybernetics 27 (1977), S. 215-220 
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    Notes: Abstract The process of selection of target points during voluntary eye movements when polygonal random shapes are observed was analysed in humans by means of an eye movement recording technique. A computer model was constructed with the aim to explain the empirical results. It has been found that the majority of fixation points were located at the angles. The marginal distribution over the x-axis of the individual angles depends on the angle's size: the maximal value of distribution was found for acute angles more distant from the vertex than in obtuse ones. The distribution of output activity of the “ganglion cells” in a computer model, reproducing some basic features of the retina, is in good agreement with the empirical results.
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    Biological cybernetics 34 (1979), S. 181-186 
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    Notes: Abstract The observation that the amplitude of vestibular nystagmus grows as gaze is increased in the direction of the nystagmus fast phase and diminished with gaze in the opposite direction is known as “Alexander's law”. We have developed an analog computer model to simulate Alexander's law in nystagmus secondary to dysfunction of a semicircular canal. The model utilizes relevant brainstem anatomy and physiology and includes gaze modulation of vestibular signals and push-pull integration to create eye positition commands. When simulating normally functioning semicircular canals, the model produced no nystagmus. When simulating total impairment of the canal on one side with gaze directed maximally in the opposite direction, the model produced a large amplitude nystagmus with linear slow phases directed toward the affected side. As gaze was changed from far contralateral to ipsilateral, the nystagmus gradually diminished to zero. When simulating partial impairment of one canal, the nystagmus was smaller in amplitude and absent in ipsilateral gaze.
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    Biological cybernetics 34 (1979), S. 215-225 
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    Notes: Abstract The role of the optokinetic reflex (OKR) is that of cooperating with the vestibulo-ocular reflex (VOR) in the task of image stabilization on the retina during head rotations in a stationary visual surround. Since the dynamics of VOR was already well established, it has been possible to make a broad estimation of what the dynamics of OKR should be in order to obtain the performances observed in normal subjects. A mathematical model of OKR has been presented, and the experimental results obtained by Raphan et al. (1977) in the monkey and by Collins et al. (1970) in man were used to validate the model and to obtain a precise estimation of its parameters.
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    Biological cybernetics 35 (1979), S. 21-37 
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    Notes: Abstract The olfactory bulb is the first central component in a highly sensitive yet markedly stable sensory system. It receives a surge of receptor activity with each inspiration and transmits output as a brief burst of oscillatory activity that is most clearly seen in the EEG. These properties together with the known anatomy and physiology of the bulb are used as design criteria to synthesize, evaluate and solve a set of nonlinear differential equations that represent lumped bulbar dynamics. According to the model bulbar processing is in two stages. In the outer layers the interneurons perform the operations of input range compression, integration, clipping, holding, and bias control. In the inner layers the input surge is converted to a burst, which is transmitted by the mitral cells as a pulse density wave. The phase, frequency, duration, and amplitude of the wave convey information centrally about both the input and the state of the system. The model suffices to replicate the forms of the EEG burst; the pulse probability distributions conditional on the EEG; the waveforms of averaged evoked potentials (AEPs) and post stimulus time (PST) histograms from the bulb and cortex; and the changes in waveform induced by behavioral control of attentiveness and habituation. It is inferred that with selective attention there is a permanent change in the strength of mutually excitatory connections among excitatory neurons, and that with habituation there is a reversible change in the effectiveness of excitatory synapses. The limitations and deficiencies of the model and the need for centrifugal controls of bulbocortical function are discussed.
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    Biological cybernetics 35 (1979), S. 55-62 
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    Notes: Abstract This communication describes a model for two “pacemaker” (i.e., regularly firing) nerve cells, such that one elicits IPSP's in the other. The assumptions involve essentially a linear dependence (“delay function”) of the postsynaptic interval lengthening (or “delay”) produced by the IPSP's on the position (or “phase”) with respect to the preceding spike of the latter's arrival. When the number of IPSP's in an interval increases, both the slope and intercept of the delay function increase, the former remaining under 2 and the latter unboundedly. Assumptions are more or less close to the actual biological reality, or are made for convenience. A recurrence equation for the phase can be calculated, as well as an expression for the “locking phase” (see below). Plots of postsynaptic vs presynaptic firing intensity averaged over steady conditions, e.g. of mean rates or intervals, are formed by a sequence of relatively broad“paradoxical” segments exhibiting positive slopes 1, 2, 1/2, 3, 1/3, ..., indicating that “inhibited” discharges are made more intense by those increases in “inhibitory” arrivals. These segments are separated by narrower “intercalated” segments where behavior is unclear except for a large overall negative slope, indicating that “inhibited” discharges are weakened markedly by other increases in inhibitory arrivals. Across the successive paradoxical segments that correspond to more and more intense presynaptic discharges (i.e., to higher rates or shorter intervals), postsynaptic intensities, though overlapping in part, become weaker and weaker. At the extremes, when the presynaptic discharge is very weak, or very intense, the postsynaptic cell tends to its natural undisturbed firing, or to not firing at all, respectively. The pre- and postsynaptic discharges inevitably achieve eventually an invariant relation, i.e., will “lock” at a constant phase, regardless of the phase of the first IPSP arrival. The characteristics of this behavior (e.g., the rate bounds of the paradoxical segments, or the magnitude of the locked phase) depend on such givens as presynaptic and postsynaptic pacemaker rates or intervals, and as the slope or intercept of the delay function.
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    Biological cybernetics 35 (1979), S. 131-136 
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    Notes: Abstract Two aspects of real systems-unidirectional corrector activity and non-negative signals-are analysed with regard to the steady state function and structure of negative feedback systems. Two classes of simple systems are distinguished: left-regulating systems (LRS) correcting decreases of the regulated variable, and right-regulating systems (RRS) acting against increases. The RRS-comparator differs in structure from the (classical) LRS-comparator: the signs of its inputs are reversed and the feedback path is positive. Opening the feedback path-which leads to maximal activity of a LRS-switches a RRS off.
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    Biological cybernetics 35 (1979), S. 175-181 
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    Notes: Abstract 300 years ago G. W. Leibniz has invented his binary number system, now widely used in computer sciences. The paper recapitulates some informations about the contents of the first private notes from 1679, partly reproduced in facsimiles, and the scientific background of that time. The circumstances and the genesis of the first treatise on this matter, published in 1703/5 by Leibniz at the Academy of Sciences, Paris, are discussed in detail.
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    Biological cybernetics 35 (1979), S. 235-241 
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    Notes: Abstract Model calculations are presented for the several properties of the development of the retinotectal projection in amphibians and fishes, using the Gierer-Meinhardt equations. One of these properties is the maintenance of topographic mapping between the retina and the tectum during their development despite the fact that the two tissues grow in morphologically different ways. Another is the existence of a “critical period”, at which the coordinates of the retina with respect to the tectum are irrevocably determined. It is assumed that the connections between the retinal and the tectal cells are made on the correspondence of positional markers which are given as a form of the distribution of a specific activator, the dynamics of which is described by the Gierer-Meinhardt equations. The monotonic distributions of the activator and the existence of the critical period are shown by a computer simulation of the proliferating retina. Several changes of the retinotectal projection after surgical operations on the retina or the tectum are also explained.
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    Biological cybernetics 17 (1975), S. 99-108 
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    Notes: Abstract Stability properties of gravity receptor transduction mechanisms were investigated by recording action potentials from the eighth nerve of cats maintained at two spatial positions both with and without an added click perturbation. The cells were demonstrated to present directional sensitivity, multivaluedness and wandering of mean rate trajectories. To the wandering were fitted appropriate stability boundaries corresponding to recent theories of finite time stability. Results supported the hypothesis that physiological stimuli result in local deformations of a flexible trampoline-like macula.
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    Biological cybernetics 17 (1975), S. 169-182 
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    Notes: Abstract In response to tone bursts of constant sound intensity, mammalian auditory-nerve fibers produce a maximum firing rate at onset, followed by an adaptation to a quasi-steady firing rate within about 150 msec. On the basis of two fundamental findings, it appears that the adaptation is additive, or linear, in nature and does not result from a multiplicative gain change. Basically, a given increment in stimulus intensity produces the same change in firing rate before and after the adaptation. In addition, the relative amount of adaptation, i.e. the ratio of driven onset firing rate to driven steady-state firing rate, is independent of tone intensity. Nonlinear effects that appear in the experimental results may be accounted for by two static nonlinearities, one preceding and one following the linear adaptation stage.
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