The influence of basic open-loop synaptic connections on the firing of simultaneously recorded neurons has been investigated with auto- and cross-correlation histograms, using experimental records and computer simulations. The basic connections examined were direct synaptic excitation, direct synaptic inhibition, and shared synaptic input. Each type of synaptic connection produces certain characteristic features in the cross-correlogram depending on the properties of the synapse and statistical features in the firing pattern of each neuron. Thus, empirically derived cross-correlation measures can be interpreted in terms of the underlying physiological mechanisms. Their potential uses and limitations in the detection and identification of synaptic connections between neurons whose extracellularly recorded spike trains are available are discussed.
SUMMARY1. Those features of a transmembrane current correlated with spike initiation were examined in Aplysia neurones using a Gaussian whitenoise stimulus. This stimulus has the advantages that it presents numerous wave forms in random order without prejudgement as to their efficacies, and that it allows straightforward statistical calculations.2. Stimulation with a repeating segment of Gaussian white-noise current revealed remarkable invariance in the firing times of the tested neurones and indicated a high degree of reliability of their response.3. Frequencies (< 5 Hz) involved in spike triggering propagated faithfully for up to several millimetres, justifying intrasomatic current injection to examine spike initiation at the trigger locus.4. Examination of current wave forms preceding spikes indicated that a wide variety could be effective. Hence, a statistical analysis was performed, including computation of probability densities, averages, standard deviations and correlation coefficients of pairs of current values. Each statistic was displayed as a function of time before the spike.5. The average current trajectory preceding a spike was multiphasic and depended on the presence and polarity of a d.c. bias. An early relatively small inward-or outward-going phase was followed by a large outward phase before the spike. The early phase tended to oppose the polarity of the d.c. bias.6. The late outward phase of the average current trajectory reached a maximum 40-75 msec before triggering the action potential (AP) and returned to near zero values at the moment of triggering. The fact that the current peak occurs in advance of the AP may be partially explained by a phase delay between the transmembrane current and potential. The failure of the average current trajectory to return to control values immediately following the peak argues for a positive role of the declining phase in spike triggering.H. L. BRYANST AND J. P. SEGUNDO 7. Probability densities preceding spikes were Gaussian, indicating that the average was also the most probable value. Although the densities were broad, confirming that spikes were preceded by a wide variety of current wave forms, their standard deviations were reduced significantly with respect to controls, suggesting a preferred status of the average current trajectory in spike triggering.8. The matrix of correlation coefficients between current pairs suggested that spikes tended to be preceded by wave forms that in part kept close to the average current trajectory and in part preserved its shape.9. The average first and second derivatives of spike-evoking epochs revealed that current slope and acceleration, respectively, were most crucial in the last 200 msec before spike triggering, and that these dynamic stimulus components were more important for a cell maintained under a depolarizing, rather than a hyperpolarizing bias.10. A simple model of the spike-triggering system, consisting of a linear filter (first-order Wiener kernel) followed by a threshold device with 'deadtime, was quite a...
The consequences of inhibitory or excitatory synaptic input between pacemaker neurons were predicted mathematically and through digital-computer simulations, and the predicted behavior was found to occur in abdominal ganglia of Aplysia and in stretch receptors of Procambarus. Discharge patterns under conditions that do not involve interneuronal feedback are characteristic and self-stabilizing. Paradoxically, increased arrival rates of inhibitory input can increase firing rates, and increased excitatory input rates can decrease firing rates.
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