Effect of motor deprivation in early ontogeny on evoked potentials of the sensomotor and visual cortex in the rat

The influence of long-term (3 months) locomotor deprivation of rats in a month age, on the evoked potentials (EP) of the sensorimotor and the visual cortex was studied in conditions of presentation of single and paired stimuli. Changes were revealed in both cortical zones. An increase of peak latency of the initial positive EP phase in the sensorimotor cortex, and prolongation of the process of changes in excitability of neural elements, elicited by conditioning stimulus, was revealed both in the sensorimotor and the visual areas. The effect of deprivation on the dynamics of changes in neuronal systems excitability was greater in the visual evoked responses.     

Testing excitability in human firing motoneurons during the interspike interval

Monosynaptic testing of excitability in firing triceps surae muscle motoneurons activated during volitional contraction was performed using a technique for recording potentials from single motor units and by producing H-reflex. Motoneuronal excitability was assessed according to level of firing index. Motoneuronal firing index decreased during transition from a low background rhythmic firing rate of less than 6 spikes/sec to one of 6–8 spikes/sec. It hardly changed with a further rise in rate to 12 spikes/sec. The dependence between firing index and spike rate are put down to changes occurring in motoneuronal excitability during the interspike interval. Findings indicate that in the low frequency range of motoneuronal firing characteristic of natural muscle contraction, discharge rate may be considered one of the factors determining excitability in the motoneuron and hence its transmission qualities.Institute of Problems in Information Transmission, Academy of Sciences of the USSR. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 210–216, March–April, 1987.     

Primitive nervous systems: electrical activity in ventral nerve cords of the flatworm, Notoplana acticola.

Electrical activity evoked in the major cords of the ventral submuscular nerve plexus were measured. Recording and stimulation utilized suction electrodes attached directly to exposed nerve cords. Four categories of potentials have been recorded: (a) short latency spikes which have relatively high thresholds and appear to be single units; (b) short duration fast compound spikes that are made up of a small number of large units; (c) long duration compound potentials that are made up from a large number of smaller units, and (d) small amplitude potentials with long latencies and a characteristic shape. These can be conducted diffusely through the nerve plexus. The first two categories of spikes are called "fast" potentials because of their characteristic rise and fall times and the last categories are known as "diffuse" potentials. The spikelike fast potentials were only recorded from the main trunks (nerves VI), while diffuse potentials could also be recorded from side branches of these nerves. The diffuse potential appears to be conducted throughout the plexus but preferential conducting pathways occur around lesions. Both diffuse and fast potentials show facilitation of response to repeated stimulation. Facilitation can be demonstrated in the presence of high Mg2+ concentrations. Conductance of the diffuse potential also occurs in the presence of high ambient Mg2+. In Ca2+-free medium containing 1-mM EGTA one can also observe facilitatory events. The possibility of Mg2+-insensitive synapses is discussed.     

Stimulation at the cervicomedullary junction in human subjects.

In awake human subjects, corticospinal axons can be activated at the level of the cervicomedullary junction by electrical or magnetic stimulation. Such stimuli evoke single descending volleys which activate motoneurones and elicit responses in muscles of the upper limb. These responses (cervicomedullary motor evoked potentials, CMEPs) have a large monosynaptic component and can be used to test motoneurone excitability in a variety of tasks. CMEPs can be elicited in resting muscle and during all strengths of voluntary contraction. Examination of CMEPs during and after voluntary contractions reveals changes in motoneurone excitability but also suggests activity-dependent changes in the efficacy of the corticospinal pathway. Because they test the same subcortical pathway as transcranial magnetic stimulation, but are unaffected by altered excitability at a cortical level, CMEPs often offer the most appropriate comparison to allow interpretation of changes in motor evoked potentials. The advantages and disadvantages of stimulation at the cervicomedullary junction as a test of motoneurone excitability are reviewed.     

Primitive nervous systems: Electrical activity in ventral nerve cords of the flatworm,Notoplana acticola

Electrical activity evoked in the major cords of the ventral submuscular nerve plexus were measured. Recordings and stimulation utilized suction electrodes attached directly to exposed nerve cords. Four categories have been recorded: (a) short latency spikes which have relatively high thresholds and appear to be single units; (b) short duration fast compound spikes that are made up of a large units; (c) long duration compound potentials that are made up from a large number of smaller units, and (d) small amplitude potentials with long latencies and a characteristic shape. These can be conducted diffesely through the nerve plexus. The first two categories of spikes are called “fast” potentials because of their characteristic rise and fall times and the last categories are known as “diffuse” potentials. The spikelike fast potentials were only recorded from the main trunks (nerves VI), while diffuse potentials could also be recorded from side branches of these nerves. The diffuse potential appears to be concluded throughout the plexus but preferential conducting pathways occur lesions. Both diffuse and fast potentials show facilitation of response to repeated stimulation. Facilitation can be demonstrated in the presence of high Mg²⁺ concentrations. Conductance of the diffuse potential also occurs in the presence of high ambient Mg²⁺. In Ca²⁺ free medium containing 1-mM EGTA one can also observe facilitatory events. The possibility of Mg²⁺-insensitive synapses is discussed.     

Effect of seduksen (diazepam) on cortical P300 potentials evoked by neutral and emotional words

The influence of seduxen (diazepam) on cortical potentials P300 evoked by neutral and emotional words was studied in adult subjects having life conflicts. The therapeutic dose of seduxen (10 mg) had no significant effect on the amplitude of the late positive cortical response P300 to neutral words. Seduxen depresses the emotional activation of the cerebral cortex; that is manifested in selective elimination of those changes in latency and amplitude of P300 wave which are observed in response to emotionally significant words. Under the action of seduxen, the interhemispheric difference in the latency of cortical response disappears due to latency increase in the right hemisphere.     

Effect of local surface cooling of the rat cortex on the impulse activity of neurons assumed to be the sources of corticofugal influences

In an experiment on albino rats with electrodermal stimulation of the forepaw evoked potentials (EP) in the neostriatum (NS), the cortical primary response (PR), and impulse reactions of neurons (mainly of layers V and VI of the cortex) were recorded. The zone of leading-off of the potentials in the cortex was subjected to local surface cooling, which led to an increase in the PR amplitude. This facilitation was accompanied by a change in the time parameters of the impulse reactions of the cortical neurons: the latency and duration increased, and a rhythmic organization of activity appeared or intensified (if it was already present). The increase in the PR amplitude and number of spikes in the response of the cortical neurons to stimulus presentation was far less intensive than the sharp increase in EP amplitude in the NS, and did not correspond to it fully in time. The data suggest that the activating influence of the corticofugal signal on EP in the NS is determined not so much by the intensity of the descending signal as by its temporal organization.I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 23, No. 2, pp. 181–189, March–April, 1991.     

Transcranial magnetic stimulation during resistance training of the tibialis anterior muscle.

During the first few weeks of resistance training, maximal voluntary contraction (MVC) force increases at a faster rate than can be accounted for by increases in protein synthesis. This early increase in MVC force has been attributed to neural mechanisms but the sources have not been identified. The purpose of this study was to measure changes in cortical excitability with transcranial magnetic stimulation during 4 weeks of resistance training of the tibialis anterior muscle. Ten individuals performed 6 sets of 10 MVCs 3 times per week for 4 weeks and ten participated as a control group. There were no changes in any parameters tested in the control group over the 4 weeks. In the training group, TA muscle strength increased significantly by 10% at week 2 and by 18% at week 4. As hypothesized, cortical excitability during resistance training also increased. The amplitude of the TA surface EMG motor evoked potential elicited by TMS during a low-level contraction increased by 32% after training with no change in the M-wave. These data indicate that there may be an increase in cortical excitability during the first few weeks of resistance training of the TA muscle.     

Effect of desglycine-arginine vasopressin on the excitability of the command neurons of the defensive reflex in the edible snail

Perfusion of the snail (Helix lucorum L.) CNS with DG-AVP (concentration 10(-6) M) in the course of low frequency intracellular stimulation (2-4-minute interval) of the defensive reflex command neurons led to an increase in the excitability. It was expressed both in the reduction of the spike generation latency, in the increased number of spikes in response to fixed stimuli, and in the activation of pacemaker potentials. If DG-AVP was added to the medium during endoneuronal habituation, there was no increase in the excitability. It is supposed that modification of the neuronal excitability may be caused by the DG-AVP effect on the pacemaker mechanism.     

Antidromic indentification of hypothalamic neurosecretory cells in rats

Characteristics of antidromic action potentials of neurons of the paraventricular and supraoptic nuclei of the rat hypothalamus were studied during stimulation of the hypothalamo-hypophyseal tract by stimuli of varied amplitude and frequency. Step-like changes were found in spike latency in response to an increase in strength (up to 1.5–2.5 thresholds) or frequency (over 100 Hz) of stimulation, as well as cases with variation of the degree of division of the peak into A and B components. Injection of leu-enkephalin analog into the third ventricle or intravenous injection of NaCl solution (1 M) caused reversible changes in the level of excitability of antidromically activated neurons: leu-enkephalin mainly increased the latent period and threshold of action potential generation and reduced the reproducible frequency of stimulation to 10 Hz, whereas NaCl had the opposite effect. The results indicate that when the adopted criteria of antidromic identification of neurosecretory cells are used the level of their excitability must be taken into account.A. A. Ukhtomskii Physiological Research Institute, A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 14, No. 6, pp. 585–591, November–December, 1982.     

A computational framework for cortical learning

Recent physiological findings have revealed that long-term adaptation of the synaptic strengths between cortical pyramidal neurons depends on the temporal order of presynaptic and postsynaptic spikes, which is called spike-timing-dependent plasticity (STDP) or temporally asymmetric Hebbian (TAH) learning. Here I prove by analytical means that a physiologically plausible variant of STDP adapts synaptic strengths such that the presynaptic spikes predict the postsynaptic spikes with minimal error. This prediction error model of STDP implies a mechanism for cortical memory: cortical tissue learns temporal spike patterns if these spike patterns are repeatedly elicited in a set of pyramidal neurons. The trained network finishes these patterns if their beginnings are presented, thereby recalling the memory. Implementations of the proposed algorithms may be useful for applications in voice recognition and computer vision.     

Coding with spike shapes and graded potentials in cortical networks

In cortical neurones, analogue dendritic potentials are thought to be encoded into patterns of digital spikes. According to this view, neuronal codes and computations are based on the temporal patterns of spikes: spike times, bursts or spike rates. Recently, we proposed an 'action potential waveform code' for cortical pyramidal neurones in which the spike shape carries information. Broader somatic action potentials are reliably produced in response to higher conductance input, allowing for four times more information transfer than spike times alone. This information is preserved during synaptic integration in a single neurone, as back-propagating action potentials of diverse shapes differentially shunt incoming postsynaptic potentials and so participate in the next round of spike generation. An open question has been whether the information in action potential waveforms can also survive axonal conduction and directly influence synaptic transmission to neighbouring neurones. Several new findings have now brought new light to this subject, showing cortical information processing that transcends the classical models.     

Occlusion of LTP-like plasticity in human primary motor cortex by action observation

Passive observation of motor actions induces cortical activity in the primary motor cortex (M1) of the onlooker, which could potentially contribute to motor learning. While recent studies report modulation of motor performance following action observation, the neurophysiological mechanism supporting these behavioral changes remains to be specifically defined. Here, we assessed whether the observation of a repetitive thumb movement--similarly to active motor practice--would inhibit subsequent long-term potentiation-like (LTP) plasticity induced by paired-associative stimulation (PAS). Before undergoing PAS, participants were asked to either 1) perform abductions of the right thumb as fast as possible; 2) passively observe someone else perform thumb abductions; or 3) passively observe a moving dot mimicking thumb movements. Motor evoked potentials (MEP) were used to assess cortical excitability before and after motor practice (or observation) and at two time points following PAS. Results show that, similarly to participants in the motor practice group, individuals observing repeated motor actions showed marked inhibition of PAS-induced LTP, while the "moving dot" group displayed the expected increase in MEP amplitude, despite differences in baseline excitability. Interestingly, LTP occlusion in the action-observation group was present even if no increase in cortical excitability or movement speed was observed following observation. These results suggest that mere observation of repeated hand actions is sufficient to induce LTP, despite the absence of motor learning.     

Motor cortex excitability does not increase during sustained cycling exercise to volitional exhaustion

The excitability of the motor cortex increases as fatigue develops during sustained single-joint contractions, but there are no previous reports on how corticospinal excitability is affected by sustained locomotor exercise. Here we addressed this issue by measuring spinal and cortical excitability changes during sustained cycling exercise. Vastus lateralis (VL) and rectus femoris (RF) muscle responses to transcranial magnetic stimulation of the motor cortex (motor evoked potentials, MEPs) and electrical stimulation of the descending tracts (cervicomedullary evoked potentials, CMEPs) were recorded every 3 min from nine subjects during 30 min of cycling at 75% of maximum workload (W(max)), and every minute during subsequent exercise at 105% of W(max) until subjective task failure. Responses were also measured during nonfatiguing control bouts at 80% and 110% of W(max) prior to sustained exercise. There were no significant changes in MEPs or CMEPs (P > 0.05) during the sustained cycling exercise. These results suggest that, in contrast to sustained single-joint contractions, sustained cycling exercise does not increase the excitability of motor cortical neurons. The contrasting corticospinal responses to the two modes of exercise may be due to differences in their associated systemic physiological consequences.     

Inhibition of spontaneous network activity in neonatal hippocampal slices by energy substrates is not correlated with intracellular acidification

Several energy substrates complementary to glucose, including lactate, pyruvate and β-hydroxybutyrate, serve as a fuel for neurons. It was reported recently that these substrates can substantially modulate cortical excitability in neonatal slices. However, complementary energy substrates (CES) can also induce an intracellular acidification when added exogenously. Therefore, action of CES on the neuronal properties governing excitability in neonatal brain slices may be underlain by a change in the cell energy status or by intracellular acidification, or both. Here, we attempt to elucidate these possibilities in neonatal hippocampus by recording neuronal population activity and monitoring intracellular pH. We show that a spontaneous network activity pattern, giant depolarizing potentials (GDPs), characteristic for the neonatal hippocampal slices exposed to artificial cerebrospinal fluid, is strongly inhibited by CES and this effect is unlikely to be caused by a subtle intracellular acidification induced by these compounds. Indeed, a much stronger intracellular acidification in the HCO(3) -free solution inhibited neither the GDP frequency nor the GDP amplitude. Therefore, modulation of neuronal energy homeostasis is the most likely factor underlying the effect of lactate, pyruvate and β-hydroxybutyrate on network excitability in neonatal brain slices.     

Dendritic sodium spikes endow neurons with inverse firing rate response to correlated synaptic activity

Many neurons possess dendrites enriched with sodium channels and are capable of generating action potentials. However, the role of dendritic sodium spikes remain unclear. Here, we study computational models of neurons to investigate the functional effects of dendritic spikes. In agreement with previous studies, we found that point neurons or neurons with passive dendrites increase their somatic firing rate in response to the correlation of synaptic bombardment for a wide range of input conditions, i.e. input firing rates, synaptic conductances, or refractory periods. However, neurons with active dendrites show the opposite behavior: for a wide range of conditions the firing rate decreases as a function of correlation. We found this property in three types of models of dendritic excitability: a Hodgkin-Huxley model of dendritic spikes, a model with integrate and fire dendrites, and a discrete-state dendritic model. We conclude that fast dendritic spikes confer much broader computational properties to neurons, sometimes opposite to that of point neurons.     

Excitability of the sensomotor cortex and red nucleus of rabbits at different levels of spatial synchronization of cortical potentials

To determine the excitability of the rabbit sensomotor cortex and red nucleus the animal's motor response to electrical stimulation of these structures at threshold strength was investigated. In computerized experiments the excitability of these structures was compared in situations characterized by different degrees of correlation of cortical potentials. An increase in the level of spatial synchronization of cortical potentials was shown to be accompanied by an increase in the excitability of the sensomotor cortex and red nucleus. This increase in excitability is evidently a neurophysiological mechanism of the increase in probability of appearance of an effector response to sensory stimulation when the level of spatial synchronization of cortical potentials is raised.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 9, No. 1, pp. 19–24, January–February, 1977.     

Causal relationships between frequency bands of extracellular signals in visual cortex revealed by an information theoretic analysis

Characterizing how different cortical rhythms interact and how their interaction changes with sensory stimulation is important to gather insights into how these rhythms are generated and what sensory function they may play. Concepts from information theory, such as Transfer Entropy (TE), offer principled ways to quantify the amount of causation between different frequency bands of the signal recorded from extracellular electrodes; yet these techniques are hard to apply to real data. To address the above issues, in this study we develop a method to compute fast and reliably the amount of TE from experimental time series of extracellular potentials. The method consisted in adapting efficiently the calculation of TE to analog signals and in providing appropriate sampling bias corrections. We then used this method to quantify the strength and significance of causal interaction between frequency bands of field potentials and spikes recorded from primary visual cortex of anaesthetized macaques, both during spontaneous activity and during binocular presentation of naturalistic color movies. Causal interactions between different frequency bands were prominent when considering the signals at a fine (ms) temporal resolution, and happened with a very short (ms-scale) delay. The interactions were much less prominent and significant at coarser temporal resolutions. At high temporal resolution, we found strong bidirectional causal interactions between gamma-band (40–100 Hz) and slower field potentials when considering signals recorded within a distance of 2 mm. The interactions involving gamma bands signals were stronger during movie presentation than in absence of stimuli, suggesting a strong role of the gamma cycle in processing naturalistic stimuli. Moreover, the phase of gamma oscillations was playing a stronger role than their amplitude in increasing causations with slower field potentials and spikes during stimulation. The dominant direction of causality was mainly found in the direction from MUA or gamma frequency band signals to lower frequency signals, suggesting that hierarchical correlations between lower and higher frequency cortical rhythms are originated by the faster rhythms.     

Mechanically induced reflex responses in human triceps brachii

The short and long latency reflex responses of human triceps brachii muscle were recorded in 14 healthy volunteers. An electromechanical hammer was used to stretch the muscle and recordings were made from a surface electromyogram. The monosynaptic tendon reflex occurred at a mean latency of 12.5 ms (SE 0.7 ms). Later responses were observed in activated conditions (weak force production, preparatory period) at a mean latency of 62.8 ms (SE 3.5 ms). The amplitude of the short latency reflex increased during weak tension, the long latency reflex amplitude seemed to increase during the preparatory period testing. The amplitude increases can be attributed to increased lower motoneuron excitability even during weak voluntary activity. The tendency towards an increased amplitude during the preparatory period may be connected with the higher regulation of the long latency reflex.     

Selective increase in corticospinal excitability in the context of tactile exploration

In this study, we compared changes in corticomotor excitability under various task conditions engaging the index finger of each hand. Functional demands were varied, from simple execution to demanding sensory exploration. In a first experiment, we contrasted facilitation in the first dorsal interosseus (FDI) by monitoring changes in motor evoked potentials (MEPs) when participants (young adults, n = 18) performed either a simple button pressing (BP) task or a more demanding tactile exploration (TE) task (i.e., discrimination of raised letters). This experiment showed a large effect of task conditions (p < 0.01) on MEP amplitude but no effect of “Hand”, while latency measurements were unchanged. In fact, MEPs were on average 40% larger during TE (2410 ± 1358 µV) than during BP (1670 ± 1477 µV). The two tasks produced, however, different patterns of electromyographic (EMG) activity, which could have accounted for some of the differences observed. A second experimental session involved a subset of participants (10/18) tested in third task condition: finger movement (FM). The latter task consisted of scanning a smooth surface with the tip of the index finger to reproduce the movements seen with the TE task. The addition of this third condition task confirmed that MEP facilitation seen during TE reflected task-specific influences and not differences in background EMG activity. These results, altogether, provide further insights into the effect of task conditions on corticomotor excitability. Our findings, in particular, stress the importance of behavioural context and tactile exploration in leading to selective increase in corticomotor excitability during finger movements.