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Comparison of neuronal firing rates in somatosensory and posterior parietal cortex during prehension.

To evaluate their functional roles during prehension, single-unit recordings were made in the hand area of primary somatosensory areas 3b, 1 and 2 (S-I) and posterior parietal areas 5 and 7 (PPC) of the same animal. Response profiles of mean firing rate during performance of a multistage reach, grasp, and lift task were analyzed to determine the period(s) of peak firing and to measure statistically significant rises or falls in rate compared with baseline. We used the peak firing stage(s) to subdivide the population into classes tuned to single actions or two successive stages, or into multiaction groups that had sustained facilitation (BT) or inhibition (GI) during hand-object interactions. Four times as many neurons fired at peak rates during acquisition stages (approach, contact, grasp) than upon release, and their firing rates were higher. Grasping evoked the strongest responses, as grasp-tuned neurons had the highest peak rates in the population; BT, contact-grasp, and grasp-lift cells also fired maximally in the grasp stage. Grasping also coincided with maximal inhibition of GI cells, as well as of neurons tuned to approach or relaxation of grasp. Holding evoked the lowest mean rates, and had the fewest tuned cells. S-I and PPC showed significant differences in behaviors evoking peak firing as well as facilitation and inhibition; these correlated with input modalities in each area. Hand contact with the object and positioning of the fingers for grasp was the most strongly represented behavior in anterior S-I, where 61% received tactile inputs from glabrous skin. Nearly 60% were facilitated at contact, 38% fired at peak rates, and 10% were inhibited; release of grasp evoked peak firing in only 5% of 3b-1 neurons. In posterior S-I, where proportions of tactile and deep inputs were similar, positioning and grasping elicited peak responses in 38% and 31%, respectively; 80% were facilitated or inhibited during grasping. During lift and hold, inhibition rose to 43%, while excitation declined under 10%. PPC had the highest proportions firing at peak rates during hand preshaping before contact (28%) and had the most facilitated responses (38%) in this stage. Only 10% fired at peak rates during grasping. During later manipulatory actions, proportions of facilitated and inhibited responses in PPC were similar to those in posterior S-I. The data support models in which PPC plans hand movements during prehension rather than guiding their execution. Sensory monitoring of hand-object interaction occurs in S-I, where cells sense specific hand behaviors, signal stage completion, enable error correction, and may update grasp programs formulated in PPC. The results are discussed in relation to those obtained from lesion studies in humans.

Animals↗

Patterns of output firing generated by a many-input neuronal model for different model parameters and patterns of synaptic drive.

A relatively simple neuronal model with a large number of 'synaptic' inputs is described. The model has been extensively studied in order to investigate the influence which patterns of presynaptic firing may have upon the pattern of output firing. The model has been studied in two versions. In one the threshold remained constant, while in the other the threshold was increased following an output firing. This increase in threshold decayed with time and was analogous to the after-hyperpolarization of a real neurone. The effect on the frequency of output firing of changes in the threshold and of the overall input rate were studied in detail. The pattern of output firing was studied for two patterns of input firing. In one, each presynaptic input fired steadily, each at a different rate; in the other, the input firings were randomly distributed in time, being generated by a Poisson process. It was found that, for a given total input rate, the pattern of output firing was markedly more regular when the input processes were regular, even though the mean rate of output firing was not appreciably different for the two different distributions of input firings. It is shown that, with suitable parameters for the model, it is possible to mimic very closely the discharge of tonically discharging motoneurones under different experimental conditions. This suggests that the more general properties of the model may have considerable relevance to the way in which real neurones integrate their synaptic input.

Action Potentials↗

Diversity in periodic pattern of firing in human hippocampal neurons.

Firing periodicity was examined in human hippocampal neurons using autocorrelation analysis. Extracellular single-unit activities were recorded from the anterior hippocampus through fine platinum microelectrodes, and the typical firing pattern in an entire recording period was reconstructed statistically in autocorrelograms (average number of firings analyzed: 5639.0 +/- 968.1 SE, range: 1158 to 31,203; number of single-unit trains was 57). Three types of periodic firing were identified as highly consistent. The first pattern consisted of a random recurrence of high-frequency action potentials (100 to 300 Hz) and was observed as an intermittent burst. In this burst, the first 10 to 30 ms after the onset of the burst was the patterned firing of several action potentials, suggesting that the generation of this stereotyped portion of the burst is primarily due to intrinsic membrane characteristics. The second pattern was the continuous rhythmical firing with a lower frequency ranging from 1 to 30 Hz. The third pattern was a clustered rhythmical firing in which a series of short rhythmical firings recurred with regular intervals; the frequency of short rhythmical firing varied from 6.7 to 17 Hz between neurons, and the interval of the regular recurrence of these rhythmical firings ranged from 0.5 to 10 s among neurons. These firing periodicities not only cover a cellular rhythm in the theta frequency reported in the lower mammalian hippocampus but also appear to be more diverse than those previously reported for hippocampal neurons in the animal literature.

Electric Stimulation↗

Methylenedioxymethamphetamine-induced inhibition of neuronal firing in the nucleus accumbens is mediated by both serotonin and dopamine.

Methylenedioxymethamphetamine (MDMA) is a mood-altering, legally restricted drug that has been reported to inhibit glutamate-evoked firing of cells in the nucleus accumbens. This study used extracellular recording combined with microiontophoresis to examine whether the inhibitory effect of MDMA on neuronal firing in the nucleus accumbens is mediated by serotonin and/or dopamine. Serotonin and serotonin agonists with relative selectivity for the receptor subtypes 5-HT1A, 5-HT1B, 5-HT2A/2C and 5-HT3 all significantly (P < 0.01) inhibited glutamate-evoked firing of cells in the nucleus accumbens compared to the effects of an acidic saline control solution (30-60 nA, 60 s ejection currents for all). The current (dose)-dependent inhibition produced by the serotonin agonists did not differ significantly from the inhibition produced by MDMA except for the 5-HT1A agonist 8-hydroxy-(2-di-n-propylamino) tetralin, which inhibited glutamate-evoked firing significantly more than MDMA or any of the other serotonin agonists. At the highest ejection current tested (60 nA, 60 s), glutamate-evoked firing was inhibited by MDMA in 94% of tested cells, by serotonin in 80% of tested cells and by the serotonin receptor subtype agonists in 95-100% of the tested cells. In addition to being mimicked by serotonin and serotonin agonists, MDMA-induced inhibition of glutamate-evoked firing in the nucleus accumbens was partially blocked by the serotonin antagonists ketanserin (100% of tested cells), methysergide (80% of tested cells), methiothepin (100% of tested cells) and WAY100135 (100% of tested cells). Furthermore, application of the serotonin uptake blocker fluoxetine, which prevents MDMA-induced serotonin release, also significantly attenuated MDMA-induced inhibition of glutamate-evoked firing in all of the cells that were tested. These observations suggest that MDMA-induced inhibition of nucleus accumbens cell firing is at least partially mediated by serotonin. Depletion of dopamine by pretreatment with the neurotoxin 6-hydroxydopamine and the synthesis inhibitor alpha-methyl-p-tyrosine blocked the inhibition of glutamate-evoked firing produced by MDMA applied with low ejection currents (30-40 nA, 60 s). However, this dopamine depletion had no effect on inhibition of glutamate-evoked firing produced by serotonin ejected with low or high currents (20-60 nA, 60 s). These results suggest that both dopamine release and an intermediate step of MDMA-induced serotonin release are necessary for the inhibitory effects of MDMA on neuronal excitability in the nucleus accumbens. The dopamine- and serotonin-mediated inhibitory effects of MDMA on glutamate-evoked firing of nucleus accumbens cells may play a role in the mood-altering properties of this increasingly popular drug.

Animals↗

Spontaneous firing rate of lateral septal neurons decreases after forced swimming test in Wistar rat.

The systemic or local administration of diverse antidepressants increases the neuronal firing rate of the lateral septal nucleus (LSN), whereas some stressful situations decrease its firing rate; however, any long-lasting effect exerted by the forced swimming (FS) test (15-min pretest and 5-min test 24 h later) on the firing rate of the LSN is unknown. Therefore, single-unit extracellular recordings were obtained from the LSN neurons of control rats (Ctrl, n=6) and FS rats (n=10) 2 h after the last swimming session. In other rats, spontaneous firing rate of cortical neurons was recorded under the same experimental conditions. The firing rate of the LSN neurons of the animals in the FS group was significantly lower (9.2+/-1.7 spikes/10 s; P<.004, n=35) in comparison with the Ctrl group (21.1+/-3.4 spikes/10 s, n=22). The reduced firing rate in the LSN after swimming tests was both evident and generalized given that approximately 83% of the total recorded neurons from the FS group fired below the mean+/-1 S.D. rate obtained from the Ctrl group. Accordingly, the mean first-order interval of neuronal firing rate in the FS group (621.3+/-22.6 ms) was significantly greater (P<.05) than that observed in the Ctrl group (391.5+/-29.2 ms), but no significant differences were found in the variation coefficient of these two experimental groups, illustrating regularity of firing. Nonsignificant differences or even an opposite trend were observed in the firing rate of cortical neurons in the FS group (26.3+/-8.4 spikes/10 s) as compared with Ctrl group (15.4+/-1.1 spikes/10 s). Accordingly, no differences were found in the variation coefficient (FS 55.3+/-7.2%, Ctrl 55.8+/-3.6%) or average first-order interval (FS 417.8+/-71.8 ms, Ctrl 494.1+/-64.5 ms). We conclude that the FS test constitutes a situation whose capacity for inducing long-lasting despair is reflected in a reduction in the firing rate of LSN neurons as it occurs in situations of anxiety and fear, contrary to the actions of antidepressant drugs.

Action Potentials↗

Burst-firing activity of presumed 5-HT neurones of the rat dorsal raphe nucleus: electrophysiological analysis by antidromic stimulation.

We recently reported raphe neurones which frequently fired spikes in short bursts. However, the action potentials were broad and the neurones fired in a slow and regular pattern, suggesting they were an unusual type of 5-hydroxytryptamine (5-HT) neurone. In the present study, we investigated whether these putative burst-firing 5-HT neurones project to the forebrain and whether all spikes fired in bursts propagate along the axon. In anaesthetised rats, electrical stimulation of the medial forebrain bundle evoked antidromic spikes in both burst-firing neurones and in single-spiking, classical 5-HT neurones recorded in the dorsal raphe nucleus. Although the antidromic spike latency of the single-spiking and burst-firing neurones showed a clear overlap, burst-firing neurones had a significantly shorter latency than single-spiking neurones. For both burst-firing neurones and classical 5-HT neurones, antidromic spikes made collisions with spontaneously occurring spikes. Furthermore, in all burst-firing neurones tested, first, second and third order spikes in a burst could be made to collide with antidromic spike. Interestingly, in a small number of burst-firing neurones, antidromic stimulation evoked spike doublets, similar to those recorded spontaneously. From these data we conclude that burst-firing neurones in the dorsal raphe nucleus project to the forebrain, and each spike generated by the burst propagates along the axon and could thereby release transmitter (5-HT).

Action Potentials↗

Plasticity and tuning of the time course of analog persistent firing in a neural integrator.

In a companion paper, we reported that the goldfish oculomotor neural integrator could be trained to instability or leak by rotating the visual surround with a velocity proportional to +/- horizontal eye position, respectively. Here we analyze changes in the firing rate behavior of neurons in area I in the caudal brainstem, a central component of the oculomotor neural integrator. Persistent firing could be detuned to instability and leak, respectively, along with fixation behavior. Prolonged training could reduce the time constant of persistent firing of some cells by more than an order of magnitude, to <1 s. Normal visual feedback gradually retuned persistent firing of integrator neurons toward stability, along with fixation behavior. In animals with unstable fixations, approximately half of the eye position-related cells had upward or unstable firing rate drift. In animals with leaky fixations, two-thirds of the eye position-related cells showed leaky firing drift. The remaining eye position-related cells, generally those with lower eye position thresholds, showed a more complex pattern of history-dependent/predictive firing rate drift in relation to eye drift. These complex drift cells often showed a drop in maximum persistent firing rate after training to leak. Despite this diversity, firing drift and the degree of instability or leak in firing rates were broadly correlated with fixation performance. The presence, strength, and reversibility of this plasticity demonstrate that, in this system, visual feedback plays a vital role in gradually tuning the time course of persistent neural firing.

Animals↗

Electrical coupling between model midbrain dopamine neurons: effects on firing pattern and synchrony.

The role of gap junctions between midbrain dopamine (DA) neurons in mechanisms of firing pattern generation and synchronization has not been well characterized experimentally. We modified a multi-compartment model of DA neuron by adding a spike-generating mechanism and electrically coupling the dendrites of two such neurons through gap junctions. The burst-generating mechanism in the model neuron results from the interaction of a N-methyl-D-aspartate (NMDA)-induced current and the sodium pump. The firing patterns exhibited by the two model neurons included low frequency (2-7 Hz) spiking, high-frequency (13-20 Hz) spiking, irregular spiking, regular bursting, irregular bursting, and leader/follower bursting, depending on the parameter values used for the permeability for NMDA-induced current and the conductance for electrical coupling. All of these firing patterns have been observed in physiological neurons, but a systematic dependence of the firing pattern on the covariation of these two parameters has not been established experimentally. Our simulations indicate that electrical coupling facilitates NMDA-induced burst firing via two mechanisms. The first can be observed in a pair of identical cells. At low frequencies (low NMDA), as coupling strength was increased, only a transition from asynchronous to synchronous single-spike firing was observed. At high frequencies (high NMDA), increasing the strength of the electrical coupling in an identical pair resulted in a transition from high-frequency single-spike firing to burst firing, and further increases led to synchronous high-frequency spiking. Weak electrical coupling destabilizes the synchronous solution of the fast spiking subsystems, and in the presence of a slowly varying sodium concentration, the desynchronized spiking solution leads to bursts that are approximately in phase with spikes that are not in phase. Thus this transitional mechanism depends critically on action potential dynamics. The second mechanism for the induction of burst firing requires a heterogeneous pair that is, respectively, too depolarized and too hyperpolarized to burst. The net effect of the coupling is to bias at least one cell into an endogenously burst firing regime. In this case, action potential dynamics are not critical to the transitional mechanism. If electrical coupling is indeed more prominent in vivo due to basal level of modulation of gap junctions in vivo, these results may indicate why NMDA-induced burst firing is easier to observe in vivo as compared in vitro.

Action Potentials↗

Persistent sodium currents and repetitive firing in motoneurons of the sacrocaudal spinal cord of adult rats.

Months after sacral spinal transection in rats (chronic spinal rats), motoneurons below the injury exhibit large, low-threshold persistent inward currents (PICs), composed of persistent sodium currents (Na PICs) and persistent calcium currents (Ca PICs). Here, we studied whether motoneurons of normal adult rats also exhibited Na and Ca PICs when the spinal cord was acutely transected at the sacral level (acute spinal rats) and examined the role of the Na PIC in firing behavior. Intracellular recordings were obtained from motoneurons of acute and chronic spinal rats while the whole sacrocaudal spinal cord was maintained in vitro. Compared with chronic spinal rats, motoneurons of acute spinal rats were more difficult to activate because the input resistance was 22% lower and resting membrane potential was hyperpolarized 4.1 mV further below firing threshold (-50.9 +/- 6.2 mV). In acute spinal rats, during a slow voltage ramp, a PIC was activated subthreshold to the spike (at -57.2 +/- 5.0 mV) and reached a peak current of 1.11 +/- 1.21 nA. This PIC was less than one-half the size of that in chronic spinal rats (2.79 +/- 0.94 nA) and usually was not large enough to produce bistable behavior (plateau potentials and self-sustained firing not present), unlike in chronic spinal rats. The PIC was composed of two components: a TTX-sensitive Na PIC (0.44 +/- 0.36 nA) and a nimodipine-sensitive Ca PIC (0.78 +/- 0.82 nA). Both were smaller than in chronic spinal rats (but with similar Na/Ca ratio). The presence of the Na PIC was critical for normal repetitive firing, because no detectable Na PIC was found in the few motoneurons that could not fire repetitively during a slow ramp current injection and motoneurons that had large Na PICs more readily produced repetitive firing and had lower minimum firing rates compared with neurons with small Na PICs. Furthermore, when the Na PIC was selectively blocked with riluzole, steady repetitive firing was eliminated, even though transient firing could be evoked on a rapid current step and the spike itself was unaffected. In summary, only small Ca and Na PICs occur in acute spinal motoneurons, but the Na PIC is essential for steady repetitive firing. We discuss how availability of monoamines may explain the variability in Na PICs and firing in the normal and spinal animals.

Animals↗

Jaw muscle afferent firing during an isotonic jaw-positioning task in the monkey.

The activity of jaw muscle receptors was studied by recording neurons in the mesencephalic nucleus of the trigeminal nerve in monkeys trained to control the position and movement of their mandible. Jaw position was measured by a weighted lever resting on the mandibular incisors. The force required to maintain the position of the lever was varied; in most cases it was either 25 or 360 g. Firing rates of neurons were related to stationary mandibular positions and to the velocity of movements during intervals when the movement velocity was constant. Of 49 neurons studied in detail, 21 fired at rates that were consistently and linearly related to static incisal openings. This static position sensitivity was typically about 5 spikes/mm of incisal opening. Most position-sensitive neurons fired at higher rates during opening movements and at lower rates during closing movements than would be accounted for by their position sensitivity. This sensitivity to the velocity of movement was not linear, however; slow closing movements sometimes did not produce a decrease in firing rate, and an actual increase during muscle shortening was seen in a few instances. The position sensitivity of eight neurons was evaluated during different loading conditions; in no case did it change substantially. Of the remaining 28 neurons, 26 fired at high rates during all opening movements and either stopped firing or fired at low, sporadic rates during closing movements. The static position sensitivity of these neurons was weak and variable both within and between neurons. The velocity sensitivity of these stretch-sensitive neurons was very nonlinear. Except for a range of slow movements (+/- 5 mm/s), the firing rate was maximal (200 spikes/s or higher) for most opening movements and zero for most closing movements. Maximal firing rates were higher when the loads being moved were increased from 25 to 360 g. The majority of position-sensitive neurons exhibited a large interspike-interval variability at wide incisal opening. In most of these neurons, this interspike-interval variability was periodic, usually at a rate of about 10 periods/s, and took the form of "saw-tooth" modulation on a record of instantaneous firing rate. Neurons that exhibited this modulation in a very prominent form also exhibited, in many instances, a substantial increase in firing rate during closing jaw movements.

Afferent Pathways↗

Spontaneous burst firing in cat primary auditory cortex: age and depth dependence and its effect on neural interaction measures.

1. Neural activity was recorded with two independent electrodes separated by 0.5-2 mm, aligned in parallel, and advanced perpendicular to the surface of the cat auditory cortex. Because the experiments were part of a study into laminar interaction the difference in recording depths for the two independently movable electrodes was never > 100 microns. Multi-unit activity on each electrode was separated on-line into single-unit spike-trains with a maximum variance spike sorting algorithm. Off-line controls on the quality of the spike-train separation were routinely performed. The first aim of this study was to describe the age dependence of spontaneous burst firing and to explore if and how it could be explained by age dependent changes in firing rate. The second aim was to investigate a potential layer dependence on burst firing. The third aim was to describe the effect of burst-removal procedures on the shape, strength, and width of the cross-correlogram and to investigate whether an age dependence in burst firing might account for the previously reported age dependence in correlation strengths. 2. Recordings were made from 237 single units from primary auditory cortex in nine adult cats and from 67 units in seven kittens age 10-52 days. The incidence of burst firing as a function of firing rate, age and depth of recording and unit characteristic frequency was investigated. In addition the effect of burst firing on the strength and width of the central peak in 471 neural pair correlograms was analyzed. 3. Burst firing could be distinguished at many different time scales; bursts lasting of the order of 10 s contained bursts with durations of the order of 1 s, which in turn contained bursts of 30-50-ms duration. The analysis in this paper was restricted to the short-duration bursts. 4. Burst firing on the short-time scale of 50 ms was characterized by relatively well defined intervals between the first two spikes (3-15 ms) followed by higher-order intervals with large spread (range 4-50 ms) but with increasing modal interval value. The typical adult five-spike burst template featured spikes at 0, 3.3, 14.6, 27.2, and 34.8 ms. Burst with fewer spikes showed larger intervals between the first three spikes. 5. The probability of occurrence of isolated spikes, pairs, triplets, etc. showed a power-law dependence on firing rate with a coefficient that was significantly lower than expected under Poisson firing conditions.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

Effect of stimulation on burst firing in cat primary auditory cortex.

1. Neural activity was recorded extracellularly with two independent microelectrodes aligned in parallel and advanced perpendicular to isofrequency sheets in cat primary auditory cortex. Multiunit activity was separated into single-unit spike trains using a maximum variance spike sorting algorithm. Only units that demonstrated a high quality of sorting and a minimum spontaneous firing rate of 0.2 spikes/s were considered for analysis. The primary aim of this study was to describe the effect of periodic click train and broadband noise stimulation on short-time-scale (< or = 50 ms) bursts in the spike trains of single auditory cortical units and to determine whether stimulation influenced the occurrence, spike count, and/or temporal structure of burst firing relative to a spontaneous baseline. 2. Extracellular recordings were made in 20 juvenile and adult cats from 69 single auditory cortical units during click train stimulation and silence, and from 30 single units during noise stimulation and in silence. In an additional 15 single units the effect of both click train and noise stimulation was investigated. The incidence, spike count, and temporal structure of short-time-scale burst firing in the first 100 ms following stimulus presentation was compared with burst firing in the period starting 500 ms after stimulus presentation and with spontaneous burst firing. In addition, the serial dependence of interspike intervals within a burst was tested during periods of stimulation. 3. Burst firing was present in the stimulation, poststimulation, and spontaneous conditions. Longer bursts (consisting of > or = 3 spikes) were more commonly observed in the poststimulation and spontaneous conditions than in the stimulation condition. This effect was most pronounced during click stimulation. A period of elevated firing activity was present in a subset of units 0.5-1.5 s after stimulus presentation, indicating prolonged effects of stimulation on single-unit firing behavior. 4. For both stimuli, the proportion of single-unit responses composed of bursts was significantly greater in poststimulation and spontaneous periods than during stimulation. Burst rate was higher in post-click-train stimulation and spontaneous periods than during periods of click stimulation. The isolated spike rate was significantly higher during periods of noise and click stimulation than in the poststimulation and spontaneous periods. 5. An examination of the autocorrelograms and higher-order interspike interval histograms of single-unit responses during click train stimulation indicated that 25% of single-unit spike trains contained an excess of brief first-order intervals and 14% of spike trains contained a shortage of long higher-order interspike intervals relative to a spontaneous baseline. During noise stimulation, 10% of single-unit responses contained an excess of short intervals relative to baseline. Interspike intervals of short-duration bursts were not serially dependent during periods of stimulation. 6. A comparison of the autocorrelograms and higher-order interval histograms of single-unit responses in the poststimulation and spontaneous conditions indicated that 20% of single-unit spike trains contained an excess of short first-, second-, and third-order intervals following stimulation. This subgroups of single units could not be distinguished on the basis of the age of the animal or the depth at which the recording was made. 7. The low incidence of burst firing during stimulation opposes the view that bursts serve as a mechanism to emphasize or amplify particular stimulus-related responses in the presence of ongoing spontaneous activity in the primary auditory cortex. Moreover, there is little evidence to support the notion that brief bursts represent neural codes, because intraburst intervals are not serially dependent. It is suggested that pyramidal burst firing may be an effective way to evoke postsynaptic firing in inhibitory interneurons and subsequ

Acoustic Stimulation↗

Regularity of firing of neurons in the inferior colliculus.

The spike discharge regularity of 254 tonically firing units in the inferior colliculus (IC) of the anesthetized guinea pig was studied in response to tones presented at best frequency (BF) to the ear contralateral to the recorded IC. Regularity of firing was measured by calculating the coefficient of variation (CV) as a function of time over the course of a unit's response. Two hundred and fifteen units (56 under urethan and 159 under chloralose anesthesia) in the central nucleus of the IC (CNIC) were studied in detail. In response to tones at 15-25 dB above threshold, 80% of units in the urethan sample fired regularly (CV < 0.5) during their sustained response, and 46% were highly regular (CV < or = 0.35). For chloralose the values were 68% and 23%, respectively. Units recorded under urethan were significantly more regular than those recorded under chloralose. For units in the sample with a measurable onset CV, 63% were regular and 44% highly regular under urethan, and 73% were regular and 54% highly regular under chloralose. The units' peristimulus time histogram (PSTH) patterns were classified into subdivisions of four categories: choppers [9%: chop-sustained (Cs), chop-onset (Co)]; pausers [42%: pauser-chop-sustained (P/Cs), pauser-chop-onset (P/Co), pauser-no-chop]; ON-sustained (43%: primary-type, L-type, h-type); and sustained (6%). The presence of chopping was a reliable predictor of regularity: Cs and P/Cs units were highly regular throughout their response, whereas Co and P/Co units were highly regular at onset and became less regular. Some units in the other PSTH categories were highly regular despite the absence of chopping, and units with virtually identical PSTHs showed very different sustained CVs. Regularity was measured as a function of firing rate in 71 units. In 23%, regularity remained constant when firing rate changed with stimulus level. Forty-six percent fired more regularly as firing rate increased, 8% fired less regularly, and 23% of units showed no consistent relationship between CV and firing rate. Regularity did not correlate with the neurons' frequency response areas or BFs. Regular firing was also found in a smaller sample of units recorded in cortices surrounding the CNIC. We conclude that regular firing is a characteristic feature of most neurons in the IC. Regularity is a specific feature correlated with four PSTH types (Cs, Co, P/Cs, and P/Co). Other PSTH types may or may not exhibit regularity.

Animals↗

Motor unit firing behavior in man.

Studies on motor unit firing behavior in man by the decomposition technique are described. The decomposition technique identifies motor unit firing with 100% accuracy at force levels of greater than 80% maximal voluntary contraction (MVC). In all muscles examined, the higher the recruitment threshold of the motor unit, the lower the rate at which it fired at the target level. Smaller muscles, such as those in the hand, recruit their motor units at 0-50% MVC and rely exclusively on firing rate increases to augment force output at 50-100% MVC. Larger muscles, such as those in the leg or arm, recruit motor units at least to 90% MCV, and possibly higher. Their firing rates have a relatively smaller dynamic swing. Thus, smaller muscles rely primarily on firing rate and larger muscles rely primarily on recruitment to modulate their force. High cross-correlation functions in firing rate behavior within a muscle were observed between individual motor unit firing rates at constant force isometric contraction. Thus the nervous system does not control the firing rate of motor units individually. Instead, it acts on the pool of homonymous motoneurons in a uniform fashion. Electrical stimulation of cutaneous receptors tends to increase the recruitment thresholds of low-threshold motor units and to decrease their firing rates, while high-threshold motor units generally exhibit a decrease in recruitment threshold and an increase in firing rate.

Action Potentials↗

Oscillatory firing of single human sphincteric alpha 2 and alpha 3-motoneurons reflexly activated for the continence of urinary bladder and rectum. Restoration of bladder function in paraplegia.

1. By recording with 2 pairs of wire electrodes from human sacral nerve roots (S3-S5) rhythmic as well as occasional firing was observed in alpha 2 and alpha 3-motoneurons in response to physiologic stimulation of the urinary bladder and the anal canal. The rhythmic firing consisted of periodically occurring impulse trains, most likely produced by true spinal oscillators which drove the motoneurons. 2. Alpha 2-motoneurons, innervating fast fatigue-resistant muscle fibres, were observed to fire with impulse trains of about 2 to 4 action potentials (Ap's). These impulse trains occurred every 110 to 170 msec (5-9 Hz). Alpha 3-motoneurons, innervating slow fatigue-resistant muscle fibres, fired about every 1400 msec (approximately 0.7 Hz) with impulse trains of about 11 to 60 Ap's. Alpha 1-motoneurons, innervating fast fatigue muscle fibres, and gamma-motoneurons were not observed in the continuous oscillatory firing mode. 3. Sphincteric motoneurons were observed most likely in the oscillatory firing mode in response to the sustained stretch (reflex) of the external and sphincter or to retrograde filling of the bladder (urethro-sphincteric guarding reflex), in order to preserve continence. A urethral sphincteric alpha 2-motoneuron increased its mean activity from 0.5 to 18 Ap's/sec during retrograde filling by changing its firing pattern from the occasional spike mode via the transient oscillatory firing mode to the continuous oscillatory mode. Up to a filling of the bladder of 500 ml the mean activity of the stretch receptors, measuring probably mural tension, increased roughly proportionally and the sphincteric motoneuron increased its activity to about 1 Ap/sec in the occasional spike mode. Up to 600 ml, the motoneuron responded in the transient oscillatory mode with mean activities of up to 5 Ap's/sec. With higher bladder fillings, the flow receptors afferents fired additionally, probably according to pressure symptoms, and the motoneuron switched into the continuous oscillatory firing mode and increased its activity up to 18 Ap's/sec at 700 ml. When the bladder was about 800 ml full, the stretch afferent activity decreased, the flow receptor activity increased strongly and the alpha 2-motoneuron activity decreased; the overflow incontinence had probably started. Micturition was not observed, probably because of brain death. 4. It is suggested that one adequate stimulus for an alpha 2-motoneuron of the external anal sphincter to jump into the oscillatory firing mode, was the activity from secondary spindle afferent (SP2) fibres from external anal sphincter muscle spindles.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗

[Effects of the firing temperature on the color and strength of metal bonded porcelain].

There are many reports on the optical properties of natural teeth and fused porcelain. However, only few studies have been reported on the relationship between the firing temperature, and the optical and mechanical properties of metal bonded porcelain. This is a study of the effects of the firing temperature on the optical and mechanical properties of two groups of porcelain, each of which has five kinds, opaque, body, incisal, transparent and glaze respectively. The results were as follows: 1. The chief ingredient of each porcelain was leucite (K2O . Al2O3 . 4SiO2). Some other components were metal oxides used as pigment and opaque, for regulating the recrystallization of leucite in the fused porcelain, for controlling the thermal expansion, and for diffusing in the both sides of alloy and porcelain. 2. About 40% of each porcelain powder consisted of granular powder, smaller than 10 microns in diameter. 3. Measured by using a color analyser, L, a, and b values of all opaque porcelain did not depend upon the firing temperature. L value of the other porcelain showed decrease in proportion to the increase of the firing temperature. And it slightly decreased over 900-940 degrees C. 4. The firing temperature did not affect the transparency of the opaque porcelain. The transparency of the other porcelain increased in proportion to the increase of the firing temperature. And it also showed slight increase over 900-940 degrees C. The firing temperature did not affect the Heize values of the opaque porcelain. The Heize values of the other porcelain decreased in proportion to the increase of the firing temperature. 5. The compressive and diametral tensile strength of all the porcelain increased in proportion to the increase of the firing temperature to 910-940 degrees C. However, it decreased in proportion to the increase of the firing temperature over 910-940 degrees C. 6. It was confirmed by the SEM observations, microphotographs and X-ray diffraction of the fused porcelain that fully fusing between particles occurred at the firing temperature of 910-940 degrees C. And it was also confirmed by surface roughness.

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Morphine and methionine-enkephalin: different effects on spontaneous and evoked neuronal firing in the mesencephalic reticular formation of the rat.

The technique of microiontophoresis was used to determine the effects of morphine and methionine-enkephalin (met-enkephalin) on spontaneous neuronal firing and on firing evoked by a nociceptive stimulus (evoked firing). Morphine and metenkephalin produced one of the three following patterns of effects on single units in the mesencephalic reticular formation: 1) morphine but not met-enkephalin blocked evoked firing; 2) met-enkephalin but not morphine blocked evoked firing; and 3) both morphine and met-enkephalin blocked evoked firing. For neurons exhibiting each of these three patterns, the mean T100, a modification of the T50 which is analogous to a dose-response curve, correlated with the effects of the drugs on evoked firing. There appear to be differences in cell size and location which are associated with different effects of the drugs. The difference in the effects of the drugs on evoked firing cannot be explained by differences in transport number, diffusion or degradation of the drugs, nor by different locations of the drug ejection barrels. Naloxone, administered intravenously or micriontophoretically, antagonized the drug-induced blockade. The effects of morphine and met-enkephalin on spontaneous firing did not correlate with their effect on evoked firing. Furthermore, in the majority of cases, the effects of morphine and met-enkephalin on spontaneous firing were not the same. These data indicate that there may be more than one type of opiate receptor in the mesencephalic reticular formation.

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Mortality and cancer incidence in Stockholm fire fighters.

Fire fighters are exposed to irritating, asphyxiating, and toxic gases and aerosols, to psychological stress, and to physically demanding work. Due to differences in fire fighting techniques, exposure conditions for fire fighters differ among different countries. The purpose of this investigation was to study cancer incidence and mortality in fire fighters who have been working with fire fighting methods used in Sweden from the beginning of this century onwards. All male fire fighters employed for at least 1 year in the City of Stockholm during 1931-1983 were traced, and an index of the number of fires fought was calculated for each individual. The mortality during 1951-1986 (among 1, 116 fire fighters) was lower than expected (SMR = 82; 95% confidence interval 72-91) compared with local mortality rates, with a low mortality in circulatory diseases, obstructive lung diseases, violent deaths, and suicides. The cancer incidence in 1958-1986 was equal to the expected (SMR = 100; 95% confidence interval 83-119). However, an excess of stomach cancer (18 observed vs. 9.37 expected; SMR = 192, 95% CI 114-304) was observed. There was also a tendency for higher incidence and mortality in stomach and brain cancer with increasing number of fires. There were four deaths from brain cancer compared to 0.8 expected (SMR = 496; 95% CI 135-1270) in the highest exposure category. Fire fighters are, however, not systematically exposed to known stomach or brain carcinogens, and the results need confirmation in further studies with extensive exposure evaluations.

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