Gain control in the electrosensory system: a role for the descending projections to the electrosensory lateral line lobe

The responses of E-cells, basilar pyramidal cells, of the electrosensory lateral line lobe (ELLL) were studied in normal animals (Apteronotus leptorhynchus) and in fish in which a component of the descending input from the midbrain n. praeeminentialis to the ELLL was interrupted by lesions or by application of local anesthetics. This treatment increased the responsiveness of these neurons by 100 to 300%. A method is described by which the animal's electric organ discharge (EOD) can be increased or decreased in amplitude. Responses of E-cells to a brief stationary electrosensory stimulus and to moving electrolocation targets were studied in normal and in lesioned animals with normal and altered EOD amplitudes. Large reductions in EOD amplitude, approximately 50%, result in no significant changes in the average size of E-cells' responses to either type of electrosensory stimulus in normal animals. Interruption of the descending input, however, results in a loss of the E-cells' ability to maintain constant response size when the EOD amplitude is reduced. Increases in EOD amplitude cause reductions in the size of E-cell responses to the moving electrolocation targets and to the stationary stimulus. The effects of increased EOD amplitude are present in normal animals and in animals in which the descending input is interrupted. The descending input to the ELLL seems to function as a gain control mechanism that is capable of compensating for losses in stimulus strength resulting from reduced EOD amplitude. The component of the descending input studied here does not seem to play a role in the response of the system to increases in EOD amplitude. These results are discussed in conjunction with the known details of the ELLL circuitry and its connections with other brain areas.     

Autocorrelation and spectrum of the modulated retinal discharge: A comparison of theoretical and experimental results

The theoretical autocorrelation of the cat ganglion cell discharge under stationary and dynamic conditions of light stimulus is compared with the autocorrelation determined experimentally. To obtain the theoretical autocorrelation, the stationary discharge is described by a stationary random point process of independent intervals equally distributed according to a gamma function, and the dynamic behaviour is described by a model defined in a previous paper. Comparison shows that the model predicts the experimental results. Finally, the power density spectrum is analysed and the relevance of the results to signal transmission by the retinal system is discussed.     

Patterns in the discharge of simple and complex visual cortical cells

The activity of visual cortical neurons (area 17) was recorded in anaesthetized cats in response to sinusoidal drifting gratings. The statistical structure of the discharge of simple and complex cells has been studied as a function of the various parameters of a drifting grating: spatial frequency, orientation, drifting velocity and contrast. For simple cells it has been found that the interspike interval distributions in response to drifting gratings of various spatial frequencies differ only by a time scale factor. They can be reduced to a unique distribution by a linear time transformation. Variations in the spatial frequency of the grating induce variations in the mean firing rate of the cell but leave unchanged the statistical structure of the discharge. On the contrary, the statistical structure of the simple cell activity changes when the contrast or the velocity of the stimulus is varied. For complex cells it has been found that the invariance property described above for simple cells is not valid. Complex cells present in their activity in response to visual stimuli two different firing patterns: spikes organized in clusters and spikes that do not show this organization ('isolated spikes'). The clustered component is the only component of the complex cell discharge that is tuned for spatial frequency and orientation, while the isolated spike component is correlated with the contrast of the stimulus.     

Properties of the feedback system controlling the coxa-trochanter joint in the stick insect Carausius morosus

The angle of the coxa-trochanter (C-T) joint in the stick insect Carausius morosus is controlled by a negative feedback mechanism. It is shown that the trochanteral hair plate alone functions as the feedback transducer and that the rhomboid hair plate is not involved in the feedback loop.The properties of the C-T control system were investigated by means of force measurements. The results cannot be adequately described in all details by either a fractional differentiator model, a model which fits many sensory systems, or a nonlinear bandpass filter, a model which fits the force response of the femur-tibia feedback loop. The fractional differentiator model adequately describes the frequency response of the open-loop system to sinusoidal stimulation with 34 deg stimulus amplitude. However, the responses to sinusoidal and steplike stimulation with 10 deg stimulus amplitude do not fit this model. They are better described by the model of a nonlinear bandpass filter.The possible contribution of mechanical properties of the musculature and the joint to the total force response is discussed. It is suggested that cocontractions occurring at higher stimulus frequencies alters the muscle properties and enables the animal to respond to stimulus frequencies above the upper corner frequency of the active feedback loop.     

Behavioral evidence of a latency code for stimulus intensity in mormyrid electric fish

Mormryid electric fish (Gnathonemus petersii) respond to novel stimuli with an increase in the rate of the electric organ discharge (EOD). These novelty responses were used to measure the fish's ability to detect small changes in the amplitude and latency of an electrosensory stimulus. Responses were evoked in curarized fish in which the EOD was blocked but in which the EOD motor command continued to be emitted. An artificial EOD was provided to the fish at latencies of 2.4 to 14.4 ms following the EOD motor command.Novelty responses were evoked in response to transient changes in artificial EOD amplitude as small as 1% of baseline amplitude, and in latency as small as 0.1 ms. Changes in latency were effective only at baseline delays of less than 12.4 ms.The sensitivity to small changes in latency supports the hypothesis that latency is used as a code for stimulus intensity in the active electrolocation system of mormyrid fish. The results also indicate that a corollary discharge signal associated with the EOD motor command is used to measure latency.Abbreviations EOD electric organ discharge - ELL electrosensory lateral line lobe - epsp excitatory post synaptic potential     

Energy dependence on discharge mode of Izhikevich neuron driven by external stimulus under electromagnetic induction

Energy supply plays a key role in metabolism and signal transmission of biological individuals, neurons in a complex electromagnetic environment must be accompanied by the absorption and release of energy. In this paper, the discharge mode and the Hamiltonian energy are investigated within the Izhikevich neuronal model driven by external signals in the presence of electromagnetic induction. It is found that multiple electrical activity modes can be observed by changing external stimulus, and the Hamiltonian energy is more dependent on the discharge mode. In particular, there is a distinct shift and transition in the Hamiltonian energy when the discharge mode is switched quickly. Furthermore, the amplitude of periodic stimulus signal has a greater effect on the neuronal energy compared to the angular frequency, and the average Hamiltonian energy decreases when the discharge rhythm becomes higher. Based on the principle of energy minimization, the system should choose the minimum Hamiltonian energy when maintaining various trigger states to reduce the metabolic energy of signal processing in biological systems. Therefore, our results give the possible clues for predicting and selecting appropriate parameters, and help to understand the sudden and paroxysmal mechanisms of epilepsy symptoms.     

Electrosensory phase sensitivity in the weakly electric fish Eigenmannia in the detection of signals similar to its own

The electric organ discharge (EOD) of the South American knifefish Eigenmannia sp. is a permanently present wave signal of usually constant amplitude and frequency (similar to a sine wave). A fish perceives discharges of other fish as a modulation of its own. At frequency identity (F = 0 Hz) the phase difference between a fish's own electric discharge and that of another fish affects the superimposed waveform. It was unclear whether or not the electrosensory stimulus-intensity threshold as behaviourally determined depends on the phase difference between a fish's own EOD and a sine-wave stimulus (at F = 0 Hz). Also the strength of the jamming avoidance response (JAR), a discharge frequency shift away from a stimulus that is sufficiently close to the EOD frequency, as a function of phase difference was studied. Sine-wave stimuli were both frequency-clamped and phase-locked to a fish's discharge frequency (F = 0 Hz). In food-rewarded fish, the electrosensory stimulus-intensity threshold depended significantly on the phase difference between a fish's discharge and the stimulus. Stimulus-intensity thresholds were low (down to 3 V/cm, peak-to-peak) when the superimposed complex wave changed such that the shift in zero-crossings times relative to the original EOD was large but amplitude change minimal; stimulus-intensity thresholds were high (up to 16.9 V/cm, peak-to-peak) when the shift in zero-crossings times was small but amplitude change maximal. Similar results were obtained for the non-conditioned JAR: at constant supra-threshold stimulus intensities and F = 0 Hz, the phase difference significantly affected the strength of the JAR, although variability between individuals was higher than that observed in the conditioned experiments.Abbreviations ACP active phase coupling - EOD electric organ discharge - JAR jamming avoidance response - F frequency (fish) — frequency (stimulus) [Hz] - p-p peak-to-peak     

Bifurcation of periodic solutions of Hodgkin-Huxley model for the squid giant axon

The Hodgkin-Huxley model of the space-clamped squid giant axon is shown to admit unstable periodic solutions for current stimuli less than the stimulus at which the rest state becomes linearly unstable. The periodic solutions are demonstrated both by bifurcation theory and by numerical integration. The presence of subcritical unstable oscillations explains the discontinuous behaviour of the amplitude of the repetitive response as a function of current stimulus     

Manifestations of dynamic coding of the amplitude-modulated sounds on the level of auditory nerve fibres

Average firing rate of the auditory nerve fiber as function of the level of the tone with the frequency equal to characteristic frequency of the fibers, can be defined as an input-output characteristic. It is known that the steepening of the input-output characteristic of the real auditory nerve fiber is more, and the width is less than the spontaneous activity of the fiber. The latter characterizes fiber's ability to generate spikes, if the stimulus is absent. However it is known, that the real auditory nerve fibers with low spontaneous activity reproduce amplitude modulation of the signals much better, than the fibers with high spontaneous activity. From the results of simulation experiments, it follows that the dynamic properties of the auditory nerve fibers, providing fine tuning or adaptation of a fiber threshold under the stimulus level but not the static input-output characteristics, are the reason of fibers reproduction of stimuli amplitude modulations. However the auditory nerve fibers with high spontaneous activity due to abrupt input-output characteristic are capable to reproduce modulations of sounds whose levels are lower than a threshold of the fiber, if a weak signal adds to a weak broadband noise. This is a phenomenon of stochastic resonance found in the reactions of auditory nerve fibers.     

A novel mechanism for irregular firing of a neuron in response to periodic stimulation: irregularity in the absence of noise

Irregular firing of action potentials (AP's) is a characteristic feature of neurons in the brain. The variability has been attributed to noise from various sources. This study illustrates an alternative mechanism, namely, deterministic irregularity within a model of ionic conductances. Specifically, a model based on modern measurements of the Na⁺ and K⁺ current components from the squid giant axon fires irregularly in response to a continuous train of near-threshold current pulses. The interspike interval histogram from these simulations is multi-modal, a result which in other systems has been attributed to stochastic resonance. Moreover, the simulations exhibited short burst of spikes followed by relatively long quiescent periods, a result suggestive of patterned input to the model even though the input consisted of a train of regularly spaced current pulses. The variability of firing is attributable to variations in AP parameters, in particular AP amplitude. The action potential for squid giant axons is not all-or-none. Rather, it is fundamentally a continuous function of stimulus amplitude. That is, the membrane lacks a threshold. Variation in AP amplitude, and to a lesser extent, AP duration, can produce variations in the time to a subsequent AP, which represents a paradigm shift for understanding irregular neuronal firing. The emphasis is not as much on events prior to an AP as it is on the AP's themselves.     

Sampling Time and Performance in Rat Whisker Sensory System

We designed a behavioural paradigm for vibro-tactile detection to characterise the sampling time and performance in the rat whisker sensory system. Rats initiated a trial by nose-poking into an aperture where their whiskers came into contact with two meshes. A continuous nose-poke for a random duration triggered stimulus presentation. Stimuli were a sequence of discrete Gaussian deflections of the mesh that increased in amplitude over time – across 5 conditions, time to maximum amplitude varied from 0.5 to 8 seconds. Rats indicated the detected stimulus by choosing between two reward spouts. Two rats completed more than 500 trials per condition. Rats'' stimulus sampling duration increased and performance dropped with increasing task difficulty. For all conditions the median reaction time was longer for correct trials than incorrect trials. Higher rates of increment in stimulus amplitude resulted in faster rise in performance as a function of stimulus sampling duration. Rats'' behaviour indicated a dynamic stimulus sampling whereby nose-poke was maintained until a stimulus was correctly identified or the rat experienced a false alarm. The perception was then manifested in behaviour after a motor delay. We thus modelled the results with 3 parameters: signal detection, false alarm, and motor delay. The model captured the main features of the data and produced parameter estimates that were biologically plausible and highly similar across the two rats.     

A comparative study of the physiological properties of the inner ear in Doppler shift compensating bats (Rhinolophus rouxi andPteronotus parnellii)

Summary Cochlear microphonic (CM) and evoked neural (N-1) potentials were studied in two species of Doppler shift compensating bats with the aid of electrodes chronically implanted in the scala tympani. Potentials were recorded from animals fully recovered from the effects of anesthesia and surgery. InPteronotus p. parnellii andRhinolophus rouxi the CM amplitude showed a narrow band, high amplitude peak at a frequency about 200 Hz above the resting frequency of each species. InPteronotus the peak was 25–35 dB higher in amplitude than the general CM level below or above the frequency of the amplitude peak. InRhinolophus the amplitude peak was only a few dB above the general CM level but it was prominent because of a sharp null in a narrow band of frequencies just below the peak. The amplitude peak and the null were markedly affected by body temperature and anesthesia. InPteronotus high amplitude CM potentials were produced by resonance, and stimulated cochlear emissions were prominent inPteronotus but they were not observed inRhinolophus. InPteronotus the resonance was indicated by a CM afterpotential that occurred after brief tone pulses. The resonance was not affected by the addition of a terminal FM to the stimulus and when the ear was stimulated with broadband noise it resulted in a continual state of resonance. Rapid, 180 degree phase shifts in the CM were observed when the stimulus frequency swept through the frequency of the CM amplitude peak inPteronotus and the frequency of the CM null inRhinolophus. These data indicate marked differences in the physiological properties of the cochlea and in the mechanisms responsible for sharp tuning in these two species of bats.     

Physiologically based mathematical model of transduction of mechanobiological signals by osteocytes

Developing mathematical models describing the bone transduction mechanisms, including mechanical and metabolic regulations, has a clear practical applications in bone tissue engineering. The current study attempts to develop a plausible physiologically based mathematical model to describe the mechanotransduction in bone by an osteocyte mediated by the calcium-parathyroid hormone regulation and incorporating the nitric oxide (NO) and prostaglandin E2 (PGE2) effects in early responses to mechanical stimulation. The inputs are mechanical stress and calcium concentration, and the output is a stimulus function corresponding to the stimulatory signal to osteoblasts. The focus will be on the development of the mechanotransduction model rather than investigating the bone remodeling process that is beyond the scope of this study. The different components of the model were based on both experimental and theoretical previously published results describing some observed physiological events in bone mechanotransduction. Current model is a dynamical system expressing the mechanotransduction response of a given osteocyte with zero explicit space dimensions, but with a dependent variable that records signal amplitude as a function of mechanical stress, some metabolic factors release, and time. We then investigated the model response in term of stimulus signal variation versus the model inputs. Despite the limitations of the model, predicted and experimental results from literature have the same trends.     

Physiological properties of sympathetic preganglionic neurones in the thoracic intermediolateral nucleus of the cat

Extracellular spikes were recorded from cell bodies of sympathetic preganglionic neurones in spinal segments T1-T3 of the cat. Each neurone was identified by its antidromic response to electrical stimulation of the sympathetic chain and was found in histological sections to lie within the intermediolateral nucleus. Physiological properties studied in detail included basal activity, spike configuration, and latency of antidromic activation. Also studied, in tests with paired stimuli, were the threshold interstimulus interval evoking two responses, as well as changes in amplitude and latency of the second spike which occurred at intervals near this threshold. Approximately 60% of the units studied were spontaneously active, the rest were silent. Spontaneous activity was characterized by a slow (mean = 3.1 +/- 2.6 (SD) spikes/s), irregular pattern of discharge. With approximately one-third of the cases there was a periodic pattern of discharge in phase with oscillations in blood pressure. This correlation of phasic activity suggests that many of the units studied were involved specifically in cardiovascular function. Silent and spontaneously active units could not be differentiated on the basis of latency of antidromic activation or threshold interstimulus interval; mean latency for the two groups was 7.2 +/- 4.9 ms, mean threshold interval was 6.4 +/- 4.7 ms. Thus, with the exception of basal activity, the physiological properties studied failed to indicate more than a single population of neurones. These results therefore suggest that the sympathetic preganglionic neurones in the intermediolateral nucleus subserving varied autonomic functions share overlapping physiological properties, and that functional differentiation of these neurones may be based on differences in synaptic inputs.     

The relationship between the soleus H-reflex amplitude and vibratory inhibition in controls and spastic subjects. II. Computer model

A computer model is presented that describes soleus H-reflex recruitment as a function of electric stimulus intensity. The model consists of two coupled non-linear transfer functions. The first transfer function describes the activation of muscle spindle (Ia) afferent terminals as a function of the electric stimulus intensity; whereas the second describes the activation of a number of motoneurons as a function of the number of active Ia afferent terminals. The effect of change in these transfer functions on the H-reflex recruitment curve is simulated. In spastic patients, a higher average maximal H-response amplitude is observed in combination with a decreased H-reflex threshold. Vibration of the Achilles tendon reduces the H-reflex amplitude, presumably by reducing the excitatory afferent input. Vibratory inhibition is diminished in spasticity. In the model, the afferent-motoneuron transfer function was modified to represent the possible alterations occurring in spasticity. The simulations show that vibratory suppression of the H-reflex is determined only in part by the inhibition level of the afferent input. With a constant level of presynaptic inhibition, the suppression of reflexes of different sizes may vary. A lowering of the motoneuron activation thresholds in spastic patients will directly contribute to a decrease of vibratory inhibition in spasticity.     

The role of dynamic stimulation pattern in the analysis of bistable intracellular networks

Bistable systems play an important role in the functioning of living cells. Depending on the strength of the necessary positive feedback one can distinguish between (irreversible) “one-way switch” or (reversible) “toggle-switch” type behavior. Besides the well- established steady-state properties, some important characteristics of bistable systems arise from an analysis of their dynamics. We demonstrate that a supercritical stimulus amplitude is not sufficient to move the system from the lower (off-state) to the higher branch (on-state) for either a step or a pulse input. A switching surface is identified for the system as a function of the initial condition, input pulse amplitude and duration (a supercritical signal). We introduce the concept of bounded autonomy for single level systems with a pulse input. Towards this end, we investigate and characterize the role of the duration of the stimulus. Furthermore we show, that a minimal signal power is also necessary to change the steady state of the bistable system. This limiting signal power is independent of the applied stimulus and is determined only by systems parameters. These results are relevant for the design of experiments, where it is often difficult to create a defined pattern for the stimulus. Furthermore, intracellular processes, like receptor internalization, do manipulate the level of stimulus such that level and duration of the stimulus is conducive to characteristic behavior.     

A statistical analyzer for optic nerve messages

A statistical mathematical model of the discharge in a single optic nerve fiber is proposed, based on a discharge with intervals between impulses distributed independently according to a gamma distribution ("gamma discharge"). A light stimulus distorts the time axis of this discharge according to a "frequency function" which is characteristic of the stimulus. A linear filter is described which calculates the likelihood of a certain stimulus when the nerve fiber message is fed into it. This filter forms the basis of theoretical nerve message analyzers for three visual experiments: (a) The detection of the occurrence of a flash of light of known intensity and time of occurrence, (b) the detection of the time of occurrence of a flash of known intensity, and (c) The estimation of the intensity of a flash occurring at a known time. Possible neural mechanisms in the brain for analyzing optic nerve messages, based on the above mathematical models, are suggested. Changes of excitability or discharge frequency correspond to the output of the likelihood filter. Any such mechanism must be sufficiently plastic to have a response matched to each expected stimuus for most efficient vision near threshold.     

Sensitivity of different stimulus-timing strategies for the detection of small excitations in noisy spike train data

There are several different strategies to control the timing of a stimulus with respect to the ongoing discharge during the recording of neuronal stimulus-response characteristics. One possible strategy consists of delivering stimuli in such a way that a constant pre-stimulus spike density is reached. Another strategy enforces spike application with a constant stimulus latency after a spontaneous discharge. In this paper the sensitivity of these different strategies for statistical verification of small excitatory response components was investigated. It was found that the difference between observed poststimulus spike distribution and expected spike distribution under the null hypothesis of no stimulus effect was larger using a constant-stimulus-latency (CSL) strategy with an appropriate value for the stimulus latency. Thus, the statistical verification of neuronal response components is clearly facilitated if a CSL strategy is used. This superiority of the CSL strategy is marked, especially for small excitations at neurons discharging slowly with low discharge variability.     

Electrolocation in the presence of jamming signals: electroreceptor physiology

Responses of ampullary and tuberous electroreceptor afferents were studied using moving electrolocation targets and electrical modulations of the animal's electric organ discharge as stimuli. The ability of the electroreceptors to encode these stimuli was measured with and without various forms of electrical jamming signals. The goal of this study was to measure the deterioration in electroreceptor responses due to the jamming signals, and to compare these results with the behavioral measures of electrolocation under the same conditions of jamming as described in the preceding report (Bastian 1987). 1. Three types of jamming stimuli were used to interfere with the tuberous electroreceptor afferents' ability to respond to the test stimuli mentioned above: Broad-band noise, high frequency stimuli consisting of a sinusoidal waveform having a frequency maintained at a chosen difference frequency (DF) from the EOD frequency of the fish being studied, and 5 or 50 Hz sinusoidal stimuli. 2. The tuberous receptor afferents' spontaneous frequency was sensitive to continuous presentation of all but the 5 Hz jamming signals. The 4 Hz DF signal caused the largest increase in spontaneous activity, the 50 Hz stimulus was intermediate in effectiveness, and the noise stimulus caused the smallest increase. Estimates of the variability of the ongoing receptor activity were also made, and both the 4 Hz DF and the 50 Hz stimuli reduced the coefficient of variation of the receptor activity, but noise had no significant effect on this parameter. Noise, 4 Hz DF, and 50 Hz jamming signals also reduced the tuberous receptors' responses to a 100 ms EOD amplitude modulation, and the 5 Hz stimulus was again ineffective. 3. Noise and 4 Hz DF jamming were also effective in reducing tuberous receptor afferents' responses to a moving metal electrolocation target. The 4 Hz DF stimulus was most effective in reducing the receptor's ability to encode information about the target. Receptor responses showed about a three-fold larger decrease per 10 dB increase in DF jamming amplitude as compared to similar sized increases in noise amplitude. Threshold target distances were also determined with and without noise and DF jamming, and again, the noise stimulus was less effective in reducing the distance at which electrolocation targets were just detectable. 4. Recordings from ampullary receptor afferents confirmed that the galvanic potentials produced by metal electrolocation targets stimulate these receptors while EOD distortions caused by such objects probably do not.(ABSTRACT TRUNCATED AT 400 WORDS)