A theory of photosensitivity of some lower animals

The two-factor nerve excitation theory is applied to the photoreactions of some lower animals along with the assumption that the threshold itself varies under the action of light, thus behaving formally like a third factor. The dependence of the reaction time upon the intensity of the stimulus is derived and is shown to follow the Bunsen-Roscoe law. The dependence of reaction time upon the time of exposure to light of a fixed intensity is derived and analyzed. The effect of dark adaptation upon the reaction time is analyzed quantitatively. Light adaptation and the effects of termination of exposure after light adaptation are discussed from the standpoint of this theory and the theory is found to be in agreement with observations.     

Determination of the inter-spike times of neurons receiving randomly arriving post-synaptik potentials

Stein's model for a neuron receiving randomly arriving post-synaptic potentials is studied from an analytic viewpoint, using some recent results in the theory of first passage times for temporally homogeneous Markov processes. The case when the only input is excitatory can be treated exactly. It is shown that the moments of the firing time are guaranteed to be finite so that the differential-difference equation for the expectation (and higher moments) of the time for the membrane potential to first reach threshold from resting level can be written down. Analytic solutions are obtained in a number of cases with main emphasis on the case when the threshold is twice the epsp magnitude. An invariance principie is formulated wherein at a given mean input frequency and for a given decay parameter, the distribution of firing times depends only on the ratio of threshld to epsp magnitude. For the case where this ratio is two, the variation in the mean discharge rate is obtained as a function of mean input frequency. The results are compared with the experimental data for the Poisson monosynaptic excitation of cat motoneurons by Redmanet al. Agreement between theoretical and experimental values is excellent at input frequencies near 102 sec^-1, and theory underestimates the firing rate below that input frequency. Reasons for the discrepancy are discussed at length including the uncertainties in the neuronal parameters and the dependence of epsp magnitude on mean input frequency. The problem of including an inhibitory input process together with excitation is treated by an approximation procedure when the inhibition is considerably weaker than the excitation. At the input frequency investigated it is shown that when inhibition “half as weak” as the excitation occurs, the mean discharge frequency is approximately halved. In the final section a method of estimating neuronal parameters from the moments of the experimental inter-spike time distribution is outlined.     

On the mathematical theories of excitation

The question of constancy of the velocity of propagation of excitation is discussed on the basis of the two-factor theory, and differences between this and Blair's theory are examined.     

Effect of stimulus (postsynaptic current) shape on fibre excitation.

Effects of variation of the stimulus pulse shape on the excitation of a nonmyelinated nerve fibre were studied using a mathematical model based on the Hodgkin-Huxley equations. Efficiency of smoothly changing pulses was compared with that of rectangular pulses. For pulses shorter than the time to excitation, the rate of the stimulus rise did not determine the ability of a smoothly changing pulse to excite the fibre. For a given stimulus duration, the main factor was the pulse area or the charge delivered by the pulse. The strength-duration curve for smoothly changing pulses was a nonmonotonic function, in contrast to the curve for rectangular pulses. The dependence of latency on changes in the pulse area was non-linear. It would be nonmonotonic when the pulse area variation were due to the stimulus duration or the stimulus rise duration. More that one propagating intracellular action potential (IAP) could arise upon fibre activation by a long smoothly changing threshold stimulus. Upon activation of relatively short fibres the IAP could arise not at the site of the smoothly changing stimulus injection. The rectangular pulses of long duration were more efficient than the corresponding smoothly changing ones. Irrespective of the shape, the pulses whose duration at the foot is 1-2 ms, are more suitable for a prolonged threshold fibre activation.     

An analysis of the effects of stimulus transport and membrane charge on the salt, acid and water-response of mammals

We present an analysis of stimulus transport in mammalian tastereception emphasizing the coupling between hydrodynamic anddiffusive mass transfer. We show that flow-rate dependence inthe phasic portions of the gustatory response can be explainedby a flow-velocity dependent diffusionboundary layer in serieswith a flow-indifferent path length. Using data from the literaturewe show that the concentration dependence of the neural latencyto NaCl stimulation in the rat and the threshold concentrationcan be accounted for by the time course of stimulus arrivaland need not imply a uniquely slow sensory transduction process.We develop a generalized response function which describes aspectsof the early phasic neural response and shows that early eventsare governed solely by the local concentration of stimulus.This too is confirmed by data in the literature. The surfacepressure is characterized as an example of a nonmonotonic responsefunction which can account for the general properties of thesalt, acid and water response. In vitro studies on phopholipidmonolayers conform to the theory. It is suggested that surfaceactivity may be critical in transduction and that surface activeagents can have profound effects on taste reception.     

Dependence of the threshold of sinusoidal grating displacement on its spatial frequency characteristics

The threshold of detection of sinusoidal grating displacement was studied under conditions of the absence of apparent movement. It was shown that the threshold rises with the pattern size and depends on the number of grating cycles at a constant stimulus magnitude. The stimulus size in 2–3 periods is optimal for its localization. The results suggest that objects in the visual field are localized by mechanisms tuned to a certain spatial frequency and selective to a certain image size. The size of the analyzed part of the visual field requires respective frequency tuning of the mechanism.     

The rotation of autowaves as a result of their penetration through a system of unexcitable obstacles. A mechanism of arrhythmias associated with aging

The dependence of the frequency of occurrence of excitation vortices rotating around unexcitable obstacles on the size and the number of the obstacles and also on the medium excitation threshold was studied. It was shown that the vortex formation takes place in a wide range of the model parameter values. The assumption was formulated that the mechanism of formation of excitation vortices under study underlies the increase in the heart arrhythmias associated with aging.     

Is monomorphic tachycardia indeed monomorphic?

The dependence of some ECG characteristics on the excitation threshold was studied by mathematical modeling of cardiac arrhythmia in a 2D homogeneous excitable medium. It is shown that monomorphic tachycardias can arise both at elevated and at lowered excitability but the ECG characteristics in these cases are different.     

Chemotactic responses of Escherichia coli to small jumps of photoreleased L-aspartate

Computer-assisted motion analysis coupled to flash photolysis of caged chemoeffectors provides a means for time-resolved analysis of bacterial chemotaxis. Escherichia coli taxis toward the amino acid attractant L-aspartate is mediated by the Tar receptor. The physiology of this response, as well as Tar structure and biochemistry, has been studied extensively. The beta-2, 6-dinitrobenzyl ester of L-aspartic acid and the 1-(2-nitrophenyl)ethyl ether of 8-hydroxypyrene-1,3,6-tris-sulfonic acid were synthesized. These compounds liberated L-aspartate and the fluorophore 8-hydroxypyrene 1,3,6-tris-sulfonic acid (pyranine) upon irradiation with near-UV light. Photorelease of the fluorophore was used to define the amplitude and temporal stability of the aspartate jumps employed in chemotaxis experiments. The dependence of chemotactic adaptation times on aspartate concentration, determined in mixing experiments, was best fit by two Tar aspartate-binding sites. Signal processing (excitation) times, amplitudes, and adaptive recovery of responses elicited by aspartate jumps producing less than 20% change in receptor occupancy were characterized in photorelease assays. Aspartate concentration jumps in the nanomolar range elicited measurable responses. The response threshold and sensitivity of swimming bacteria matched those of bacteria tethered to glass by a single flagellum. Stimuli of similar magnitude, delivered either by rapid mixing or photorelease, evoked responses of similar strength, as assessed by recovery time measurements. These times remained proportional to change in receptor occupancy close to threshold, irrespective of prior occupancy. Motor excitation responses decayed exponentially with time. Rates of excitation responses near threshold ranged from 2 to 7 s-1. These values are consistent with control of excitation signaling by decay of phosphorylated pools of the response regulator protein, CheY. Excitation response rates increased slightly with stimulus size up to values limited by the instrumentation; the most rapid was measured to be 16 +/- 3 (SE) s-1. This increase may reflect simultaneous activation of CheY dephosphorylation, together with inhibition of its phosphorylation.     

Is monomorphic arrhythmia monomorphic?

It is known that the properties of myocardium affect ECG. We studied the dependence of some ECG characteristics on the threshold of excitation by means of mathematical modeling of monomorphic arrhythmias in a homogeneous two-dimensional excitable medium. We found that monomorphic arrhythmias appear at both decreased and increased values of the excitation threshold, but there are some differences between ECG characteristics for such cases.     

Types of receptive fields in the lateral geniculate body and their functional model

In the Type I receptive fields (RFs) changes of the luminance leads to a shift of the curve relating the response and the stimulus area along the abscissa, in the Type II RFs the maximum of a response does not shift with changes of the luminance (Types I and II on classification by Glezer et al., 1971, 1972). The transient responses were observed in the Type I RFs and sustained responses in the Type II RFs. In the Type I RFs variation of the stimulus area and intensity brings about the change in the temporal and spatial frequency characteristics. This is produced by functional reorganization of the RF. In the Type II RFs there is no functional reorganization. The data obtained indicate that the Type I RFs are non-linear. By contrast, the Type II RFs are linear systems. The analysis of the model has shown that the distinctions in the dynamic characteristics of the responses of RFs belonging to different types is mainly due to different time constants for excitation and inhibition as well as inhibition coefficients. Distinctions in the mode of dependence of the RF response on stimulus parameters have been found to result from different relationship between delay time and stimulus parameters as well as different forms of the spatial weighting functions. It is shown that the Type I RFs transmit higher frequency components of the image spectrum, i.e. they emphasise the temporal and spatial contrasts. The Type II RFs transmit low frequency components of the spectrum including information about the intensity of an input stimulus.     

Initiation of high-voltage discharge in air by a plasma filament produced by an intense femtosecond laser pulse

High-voltage discharge initiated in air by a plasma filament produced by an intense femtosecond laser pulse was studied experimentally. It is found that the threshold of a laser-induced discharge decrease three-fold as compared to that of a discharge in undisturbed air. It is shown that the formation time of a laser-induced discharge decreases by almost three orders of magnitude as the applied voltage increases by a factor of 2. A numerical model of the discharge process is developed that adequately describes the experimental results. In particular, simulations reproduce the experimentally observed steep dependence of the formation time of a laser-induced discharge on the applied voltage, as well as typical values of the electric field required for such a breakdown.     

BLAIR'S "CONDENSER THEORY" OF NERVE EXCITATION

Blair''s recent theory of excitation is analysed with the following conclusions: 1. The theory is inapplicable to currents of long duration; i.e., slowly increasing currents and the opening excitation. 2. The theory is a modification of the condenser theory of excitation but the modification is to be rejected on three grounds: (a) The modification has no obvious physical significance. (b) It does not in fact remedy the divergence between calculation and observation. (c) It leads to certain conclusions of a surprising kind which are contrary to observed fact. 3. The qualitative value of the condenser theory is demonstrated by the fairly close agreement between calculation and observation over a considerable field of enquiry.     

On reinforcement and interference between stimuli

It is shown that the current “two-factor” theory of nerve excitation can account for sustained inhibition or enhancement by a sequence of stimulus pulses, and for the decrease in the reinforcement period with each successive pulse of the train.     

Unsteady beam-plasma discharge: II. Nonlinear theory

The mathematical model constructed in the first part of our paper is used to numerically investigate the development of the beam-plasma instability in a traveling-wave tube amplifier in the presence of a residual neutral gas. It is shown that the self-generation of ion acoustic waves in a plasma-filled amplifier can give rise to a modulation regime with a rigid excitation threshold. The dependence of the threshold for the self-modulation instability on the amplifier parameters is determined. The effect of self-modulation on the spectral and energy characteristics of the amplifier is analyzed.     

Vector representation of associative learning

I. P. Pavlov [12] has shown that conditioned reflexes are selective both with respect to conditioned stimuli and to conditioned reflexes elicited by those conditioned stimuli. At the neuronal level selective aspects of conditioned stimuli are based on detectors selectively tuned to respective stimuli. The selective aspects of conditioned reflexes are due to command neurons representing specific unconditioned reflexes. It can be assumed that conditioned reflexes result from association between selective detectors and specific command neurons. The detectors activated by a conditioned stimulus constitute a combination of excitations--a detector excitation vector. The detector excitation vector acts on a command neuron via a set of plastic synapses--a synaptic weight vector. Plastic synapses are modified in the process of learning making command neuron selectively tuned to a specific conditioned stimulus. The selective tuning of a particular command neuron to a specific excitation vector referred to a conditioned stimulus is a basis of associative learning. The probabilities of conditioned reflexes elicited by conditioned and differential stimuli implicitly contain information concerning excitation vectors that encode respective stimuli. Contribution of the vector code to associative learning was explored combining differential color conditioning with intracellular recording from color-coding neurons. It was shown that colors in carps and monkeys are represented on a hypersphere in the four-dimensional space similar to human color space. The basis of the color space is constituted by red-green, blue-yellow, brightness and darkness neurons.     

Correct interpretation of heme protein spectra allows distinguishing between the heme and the protein dynamics

It is shown by using the vibronic approach that the iron displacement out of the porphyrin plane in deoxyheme proteins intermixes the porphyrin pi and axial iron-histidine sigma electronic subsystems. This intermixing explains the substantial coupling of the iron-histidine vibration to the heme Soret excitation, the appearance of the iron-histidine band in the corresponding resonance Raman spectra, and a number of other experimental data, including the dependence of the iron-histidine vibrational frequency on the extent of the iron displacement out of the porphyrin plane. This dependence implies that there is an anharmonic coupling between the corresponding vibrations, which is shown to be the cause of the specific temperature dependence of the iron-histidine band. The anharmonic coupling and the dependence of the dipole transition moment of the charge transfer optical absorption band III on the iron-porphyrin distance cause the anomalous temperature and pressure dependencies of this band. It is shown that the change in both the magnitude and the distribution of the iron-porphyrin distance is expected to affect the band III intensity. Consequently, the stationarity of the band III intensity can be considered as a signature of the stationarity of the iron-porphyrin distance and its distribution in deoxyheme proteins, whereas the band III position and width could be also affected by the change in the protein electric field, caused by the protein globule dynamics.     

Two phases of gramicidin photoinactivation in bilayer lipid membranes in the presence of a photosensitizer

The kinetics of the light-induced decrease in the gramicidin-mediated current across a bilayer lipid membrane in the presence of a photosensitizer has been shown to include a slow phase with a characteristic time of the order of 1 s and a fast phase. Based on the dependence of the slow phase relative amplitude and characteristic time on the gramicidin-mediated stationary conductance we concluded that the slow phase reflected the establishment of an equilibrium between gramicidin monomers and dimers in the membrane after the distortion of this equilibrium resulting from modification of a portion of gramicidin molecules by reactive oxygen species generated upon excitation of the photosensitizer. The dependence of the fast phase contribution to the overall kinetics on the stationary conductance allowed us to conclude that the fast phase is associated with transition of gramicidin dimers into a nonconducting state. The characteristic time of the fast phase measured with nanosecond laser excited pulses is 1.5 ms. The slow phase of the decrease in the gramicidin-mediated current was considerably decelerated in the presence of Rose Bengal. The results obtained indicate that adsorption of Rose Bengal on the bilayer interface leads to a reduction of the dipole potential drop at the membrane-solution boundary, similarly to the action of phloretin.     

Analysis of Possible Involvement of Local Bioelectric Responses in Chilling Perception by Higher Plants Exemplified by <Emphasis Type="Italic">Cucurbita pepo</Emphasis>

Macroelectrodes and electrophysiological setup were used in experiments with stems of 15-day-old pumpkin (Cucurbita pepo L.) seedlings for computer-assisted data recording. It is shown that local bioelectric responses induced by graded local chilling are similar to the receptor potentials of animals. These responses increase gradually with stimulus strength and trigger the action potential generation when a temperature threshold is attained. The excitation threshold of cells in seedling stems displays the phenomenon of accommodation. Parameters of local bioelectric responses induced by intermittent cooling can undergo changes similar to sensitization-habituation patterns. The results indicate that local electrical responses may be involved in early stages of cooling perception in cells of higher plants devoid of locomotive functions.