Journal of Neurophysiology

Schotland, J. L.; Rymer, W. Z.

doi: N/Apmid: 8509834

Abstract 1. To evaluate the hypothesis that the neural control of sensorimotor transformations may be simplified by using a single control variable, we compared the movement kinematics and muscle activity patterns electromyograms (EMGs) of the frog during flexion withdrawal and the hind limb-hind limb wipe reflex before and after adding an external load. In addition, the flexibility of spinal cord circuitry underlying the hind limb-hind limb wipe reflex was evaluated by comparing wipes before and after removal of one of the contributing muscles by cutting a muscle nerve. 2. The kinematics of the movements were recorded using a WATSMART infrared emitter-detector system and quantified using principal-components analysis to provide a measure of the shape (eigenvalues) and orientation (eigenvector coefficients) of the movement trajectories. The neural pattern coordinating the movements was characterized by the latencies and magnitudes of EMGs of seven muscles acting at the hip, knee, and ankle. These variables were compared 1) during flexion withdrawal and the initial movement segment of the limb during the hind limb-hind limb wipe reflex in both unrestrained movements and in movements executed when a load equal to approximately 10% of the animal's body weight was attached to a distal limb segment and 2) during the initial movement segment of the wipe reflex before and after cutting the nerve to the knee flexor-hip extensor, iliofibularis. 3. Addition of the load had no discernible effect on the end-point position of the foot during either reflex. However, during the loaded flexion reflex, the ankle joint did not move until after the hip and knee joints had moved to their normal positions. This delayed flexion of the ankle was accompanied by large increases in the magnitude of EMG activity in two ankle muscles that exceeded the levels found during unrestrained movements. Significant changes in the temporal organization of the EMG pattern accompanied the change in joint angle relations during flexion withdrawal. 4. Despite the addition of an external load, all animals successfully and reliably removed the stimulus during the wipe reflex, and the relative timing of both the EMG pattern and joint angle motion was preserved. 5. Immediately after section of the nerve to a single muscle (iliofibularis), all animals successfully and reliably removed the stimulus during the wipe reflex. The relative timing of muscle activation was preserved, accompanied by a reduction in the activity level of gluteus magnus, a muscle with action reciprocal to iliofibularis.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1993 the American Physiological Society

Chrachri, A.; Neil, D. M.

doi: N/Apmid: 8389820

Abstract 1. Extracellular and intracellular recordings from an isolated thoraco-abdominal preparation of the crayfish, Pacifastacus leniusculus, demonstrate that the swimmeret and the abdominal positioning systems can at times be spontaneously coordinated with each other. 2. Two forms of coordination were encountered between these two motor systems. First, some flexor and extensor motor neurons can burst in phase with the swimmeret power-stroke motor neurons. Second, when the flexor motor neurons displayed irregular bursting, the swimmeret rhythm was often inhibited. 3. Both of these two forms of coordination between the swimmeret and the abdominal positioning systems can be induced by depolarization of certain abdominal interneurons. 4. Bath application of oxotremorine increases the frequency of the swimmeret rhythm in a dose-dependent manner. The threshold concentration for this effect is 10(-8) M, and it persists for as long as oxotremorine is present in the bathing solution. 5. At a concentration of 10(-5) M, oxotremorine also induces slow rhythmic activity in the abdominal positioning system consisting of opposite firing between the flexor and extensor motor neurons. 6. Bath application of 10(-5) M oxotremorine also induces two types of interaction between these two abdominal motor systems. In cycle-by-cycle coordination the flexor motor neurons and one extensor motor neuron display rhythmic activity in phase with that of power-stroke motor neurons of the swimmeret system. A slow coordination also occurs with an inhibition of the swimmeret rhythm during the extensor bursts and an excitation during the flexor bursts. 7. Injection of similar doses of oxotremorine into the haemolymph of intact crayfish produces rhythmic abdominal movements that are comparable to the fictive pattern induced in the isolated preparation. Copyright © 1993 the American Physiological Society

Egan, T. M.; Dagan, D.; Levitan, I. B.

doi: N/Apmid: 8389824

Abstract 1. Single calcium-activated potassium channels (KCa channels) were recorded from membrane patches of rat olfactory bulb neurons in culture. Only one kind of KCa channel was seen, and it was present in approximately 50% of detached patches. 2. This channel, like maxi-KCa channels of other tissues, had a single-channel conductance of 270 pS, a reversal potential (Erev) of 0 mV in symmetrical K+, and was highly selective for K+ over Na+ and Cl-. 3. The KCa channel was blocked by d-tubocurarine (d-TC) on the cytoplasmic side, and charybdotoxin (CTX) on the extracellular side. This pharmacology is identical to that of one type of KCa channel from rat brain, observed previously in artificial bilayers and called the type 1 KCa channel. 4. The probability that the channel was in the open state (Po) increased with membrane depolarization. The position of the Po versus transmembrane voltage (Vm) curve was shifted by changes in Ca2+i so that the channel was open more often in higher Ca2+i. The gating kinetics resembled those of the type 1 KCa channel observed in bilayers. 5. Po was increased after superfusion of the cytoplasmic membrane surface with the active catalytic subunit of cyclic AMP-dependent protein kinase (PK-A), together with MgATP. Phosphorylation altered the distribution of channel closed times but had little effect on open times. The results suggest that phosphorylation is an important molecular mechanism in modulating the activity of this KCa channel from mammalian brain. Copyright © 1993 the American Physiological Society

Thomas, S. A.; Hume, R. I.

doi: N/Apmid: 8389829

Abstract 1. In developing chick skeletal muscle, extracellular ATP activates an early excitatory current and a delayed potassium current. Previous work had shown that the potassium current elicited by ATP is sensitive to temperature and activates with a delay of nearly 1 s, suggesting that a second messenger is involved. The existence of a second messenger was confirmed by the observation that single potassium channels were activated in cell-attached patches, when ATP was applied outside of the patch pipette. 2. Two classes of ATP-activated potassium-channel currents were observed in cell-attached patches: one had a slope conductance of 23 pS, whereas the other had a slope conductance of 51 pS. 3. Pharmacological manipulations suggested that activation of the whole-cell potassium current by ATP did not require cyclic adenosine monophosphate (cAMP), cyclic guanosine monophosphate (cGMP), inositol 1,4,5-triphosphate (IP3), nitric oxide, or a rise in internal free calcium. Additional pharmacological experiments suggested that activation of the whole-cell potassium current might not require activation of a G protein and probably did not involve intracellular protein phosphorylation. 4. The ability of arachidonic acid and its metabolites to activate potassium channels in chick skeletal muscle was also tested. Arachidonic acid, several prostaglandins and several leukotrienes activated whole-cell potassium currents. However, results with several inhibitors suggested that arachidonic acid and its metabolites are not necessary for activation of the whole-cell potassium current by ATP. 5. In excised outside-out membrane patches, ATP activated a single class of potassium channels. The slope conductance of these channels indicated that they are likely to be identical to the smaller of the two classes of second messenger activated potassium channels observed in cell-attached patches. 6. The observation that the larger class of potassium channels observed in cell-attached patches was absent in excised patches suggests that activation of these channels by ATP requires a cytosolic factor that is easily dialyzed away. In contrast, the observation that the smaller class of potassium channels could still be activated by ATP in excised patches suggests that the two classes of potassium channels are activated by different mechanisms. These results also indicate that all the molecules involved in coupling ATP receptor activation to opening of the smaller class of potassium channels remain closely associated with an excised patch. One possible explanation is that there might be an intramembranous second messenger. Copyright © 1993 the American Physiological Society

Laurent, G.; Seymour-Laurent, K. J.; Johnson, K.

doi: N/Apmid: 8389826

Abstract 1. The active properties of axonless nonspiking interneurons in the thoracic nervous system of the locust Schistocerca americana were studied in vivo with the switched current-clamp technique from dendritic impalements, and in vitro with the whole-cell variation of the patch-clamp technique. 2. In 20% of in vivo recordings, depolarization of a dendrite to potentials more positive than about -40 mV evoked resonant behaviour and/or regenerative potentials. The latter were slow (half width: 20-30 ms), small (base-to-peak amplitude: 25-35 mV), and were often followed by a pronounced after hyperpolarization (AHP). 3. The slow regenerative potentials sometimes had multiple peaks separated by incomplete repolarizations. The voltage envelope of such potentials was always broader than that of spikes with single peaks. In other recordings, a same depolarizing pulse could evoke several regenerative potentials with different waveforms. These results suggested the presence of multiple dendritic initiation sites separated by regions of inexcitable membrane, allowing decremental conduction and the passive fusion of spike envelopes. 4. Graded active responses could also be evoked on rebound from short hyperpolarizations such as inhibitory postsynaptic potentials (IPSPs) provided that the membrane was already depolarized to about -40 mV. IPSPs evoked by several presynaptic interneurons differed in their ability to evoke rebound potentials suggesting that some synaptic sites were electrically closer than others to regions of active membrane. 5. Patch-clamp recordings from somata of nonspiking neurons isolated from 75% embryos and grown in culture medium for 1-2 days revealed the presence of an inactivating inward current resistant to 0.5-1 microM tetrodotoxin (TTX). The inward current was carried equally well by Ba2+, and sensitive to blockade by Cd2+ (0.5 mM), Ni2+ (0.75 mM), or Co2+ (2.5 mM). 6. The current activated around -40 mV, with voltage-dependent activation (time-to-peak approximately 20 ms at -35 mV and 1-2 ms at 0 mV). Tail currents evoked upon repolarization were well fitted by a single exponential (tau = 1-2 ms). Deactivation time constants shorter than 300 microseconds, however, could not be measured. 7. The current inactivated rapidly in a voltage-dependent manner, following two-exponential kinetics. A very small persistent component could be explained by the overlap between activation and inactivation curves, greatest at approximately -20 mV. The voltage of half-inactivation was about -25 mV. At a resting potential of -58 mV, 90% of the current was available for activation. Recovery from steady-state inactivation followed the sum of two or more exponential processes.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1993 the American Physiological Society

Reyes, A. D.; Fetz, E. E.

doi: N/Apmid: 8389835

Abstract 1. The effects of excitatory postsynaptic potentials (EPSPs) on interspike intervals (ISIs) of neocortical neurons can be mimicked by pulse potentials (PPs) produced by current injection. The present report documents the dependence of the ISI shortening on the amplitudes of PPs and EPSPs and on the firing rate of the affected neuron. 2. In rhythmically firing necortical neurons, the ISI shortenings caused by PPs arriving at specific times in the ISI can be described by a shortening-delay (S-D) curve. The S-D curve yields three measures of the PPs' ability to shorten the ISI: 1) the mean ISI shortening, S; 2) the maximum shortening, Smax; and 3) the effective interval, defined as the portion of the ISI in which the PP consistently shortens the ISI. For PPs ranging between 80 microV and 3.6 mV (and cells firing at 25 imp/s), the mean shortening increased with amplitude h as S (ms) = 1.2*h (mV)1.24 (r = 0.94; P < 0.01). Smax increased linearly with amplitude as 4.9 ms/mV (r = 0.86, P < 0.01). The effective interval (as a percentage of the ISI) increased slightly with PP amplitude and had a mean value of 65 +/- 21% (mean +/- SD). 3. S-D curves obtained with stimulus-evoked EPSPs varied with EPSP amplitude in a manner similar to those of PPs. The relations obtained for stimulus-evoked EPSPs were not statistically different from those obtained for PPs in the same cells. 4. To determine the effect of firing rate. PPs were applied while neurons fired at frequencies ranging from 8 to 71 imp/s. Both S and Smax were approximately inversely proportional to the baseline firing rate (fo) and could be described as: S or Smax = kfo-m. The mean value of the exponent m (+/- SD) was 0.96 +/- 0.25 for S and 1.2 +/- 0.4 for Smax. These values were not statistically different from a value of 1 (1 group, 2-tailed t test). The effective interval did not vary significantly with firing rate. 5. The dependence of S on PP amplitude and baseline firing rate was incorporated into an expression for the average change in firing rate (delta f) produced by PPs occurring at rate fs: delta f = 0.03 h1.24 fs. The delta f increased with PP amplitude but did not vary significantly with the baseline firing rate. The values of delta f calculated from the S-D curves matched the values that were computed directly from the spike trains.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1993 the American Physiological Society

Crepel, V.; Hammond, C.; Krnjevic, K.; Chinestra, P.; Ben-Ari, Y.

doi: N/Apmid: 8099607

Abstract 1. The effects of an anoxic-aglycemic episode (1-3 min) on the pharmacologically isolated N-methyl-D-aspartate (NMDA)-mediated responses were examined in CA1 pyramidal hippocampal neurons in vitro. 2. An anoxic-aglycemic episode induced a long term potentiation (LTP) of the NMDA receptor-mediated field excitatory post-synoptic potentials (EPSPs). This LTP, referred to as anoxic LTP, was observed in the presence of 1) a normal Mg2+ concentration +40.1 +/- 5% (mean +/- SE), 2) a low Mg2+ concentration (+52.2 +/- 10%), or 3) a Mg2+ free (+49 +/- 11%), 1 h after anoxia. 3. Bath application of D-2-amino-5-phosphonovaleric acid (D-APV, 20 microM, 15-21 min) before, during, and after the anoxic-aglycemic episode, which transiently blocked the synaptic NMDA receptor mediated response, prevented the induction of anoxic LTP. 4. The intracellularly recorded NMDA receptor-mediated EPSP was also persistently potentiated by anoxia-aglycemia (+47 +/- 4%). This potentiation was not associated with changes in membrane potential or input resistance. 5. These findings provide the first evidence that an anoxic-aglycemic episode induces an LTP of NMDA receptor-mediated responses. This potentiation may participate in the cascade of events that lead to delayed neuronal death. Copyright © 1993 the American Physiological Society

Trotter, Y.; Celebrini, S.; Beaux, J. C.; Grandjean, B.; Imbert, M.

doi: N/Apmid: 8509829

Abstract 1. Neural correlates of the permanent deficits in depth perception that occur when extraocular muscle proprioceptive (EMP) afferents are interrupted unilaterally in kittens were investigated by performing extracellular recordings in the primary visual cortex (area 17) in adulthood. Unilateral section of the ophthalmic branch of the trigeminal nerve (V1 nerve) were performed in 11 cats when they were between 5 and 12 weeks of age (uni-V1 group). Electrophysiological results were compared with those obtained in 17 normal adult cats (control group). 2. Binocular interactions were assessed by testing the sensitivity of cortical neurons to dichoptic presentations of moving sine-wave gratings whose interocular positional phase relationship was randomly varied. The amplitude modulation between the minimum and the maximum binocular responses defined the dynamic range. The degree of binocular suppression or facilitation was assessed by comparing these binocular response limits with the optimal monocular responses evoked through either eye at the best spatial frequency. The variability of both monocular and binocular responses was estimated by using the variation coefficient. 3. In uni-V1 cats, both the dynamic range and the degree of binocular suppression were significantly less pronounced than in controls, whereas binocular facilitation was not affected. The variability of the binocular responses was significantly increased, unlike monocular responses, whose variability was similar to control values. 4. From Fourier analysis of the poststimulus time histograms, two clear-cut categories of cells emerged that were differentially affected in the uni-V1 group. The "modulated" cells showed significantly less binocular suppression than in controls, and the "unmodulated" cells had binocular responses that were significantly more variable than in controls. Results from "simple" cells were similar to those of modulated cells, and results from "complex" cells were similar to those of unmodulated cells. However, in the unmodulated population, which was composed of both simple and complex cells, it was shown that the increase of variability was due to that of complex cells. 5. A nonparametric statistical test was applied on the interocular phase shift tuning curves to determine the minimum stimulus change necessary to elicit a significant change in the neural response. Two categories of cells were determined: the "discriminative" cells (80% in controls but 45% in uni-V1 cats) combined pronounced binocular suppression and dynamic range with relatively low variability. The reverse was true in the case of "nondiscriminative" cells (20% in controls and 55% in uni-V1 cats). 6. In uni-V1 cats, about half of the cells were monocularly activated.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1993 the American Physiological Society

Tootle, J. S.

doi: N/Apmid: 8509831

Abstract 1. Spontaneous and visually evoked action potentials were recorded from single retinal ganglion cells in superfused retina-eyecups prepared from cats between postnatal day 5 (P5) and adulthood. The development of functional responses was studied quantitatively with contrast-modulated spots and sinusoidal grating stimuli. Some functionally characterized cells were impaled and injected intracellularly with horseradish peroxidase to permit direct structure-function comparisons during the period of synaptogenesis and the expression of transient morphological features. 2. Around the time of eye opening at postnatal day 7-postnatal day 10, the spontaneous discharge of ganglion cells was characterized by bursts of action potentials separated by periods of no activity lasting up to tens of seconds. This "burst-type" pattern gradually changed to a more regular spontaneous discharge during the first 2-3 postnatal wk. The burst-type discharge was completely blocked by 10 mM Mg2+, but was largely unaffected by bath application the excitatory amino acid receptor antagonist 6-cyano-7-nitroquinoxaline-2,3- dione at concentrations that eliminated both spontaneous and light-driven activity of cells that had developed a regular spontaneous discharge. DL-2-amino-7-phosphonoheptanoic acid, a competitive antagonist of the N-methyl-D-aspartate (NMDA) receptor, also had little effect on the burst-type discharge but did block the depolarizing effects of exogenous NMDA. These results demonstrate that the burst-type spontaneous discharge is a calcium-dependent process but probably is not mediated by synapses utilizing receptors for excitatory amino acids. 3. On P5, a minority of ganglion cells responded to a light stimulus delivered to the retina at the electrode tip. The percentage of responsive cells increased over time so that all cells responded by P10. During this transition period, the light responses were often weak, fatigued with repeated stimulation, and displayed poor temporal resolution. There was a rapid increase in the briskness of the response to a standard light stimulus during the second through fourth postnatal weeks. However, cells that responded sluggishly to visual stimulation were observed at all ages studied. 4. Both ON- and OFF-type receptive field (RF) centers were observed in approximately equal numbers as soon as the light response developed, and the proportion of the two types was independent of age. Three percent (10/342) of visually responsive ganglion cells had ON-OFF RFs, and the incidence of this type also appeared to be invariant with age.(ABSTRACT TRUNCATED AT 400 WORDS) Copyright © 1993 the American Physiological Society

Cohen, M. I.; Huang, W. X.; Barnhardt, R.; See, W. R.

doi: N/Apmid: 8389840

Abstract 1. In decerebrate paralyzed cats, we observed the responses of ventral and dorsal medullary inspiratory (I) neurons to two types of vagal afferent input that shorten neural I: lung inflation and vagal electrical stimulation. 2. A study population of 15 I neurons whose firing patterns suggested involvement in the inspiratory OFF-switch (IOS) was selected on the basis of two criteria: late onset of firing and excitation by vagal inputs. 3. Firing in relation to the end of I showed two types of response to vagal inputs. The pre-expiratory onset time (time from initial spike to end of I) was either unchanged (type 1 response in 5/15 neurons) or significantly changed (type 2 response in 10/15 neurons). 4. It is suggested that type 1 neurons, whose firing patterns remain closely locked to the end of I despite considerable changes of I duration, are involved in promoting the IOS, whereas type 2 neurons are either not involved (e.g., late-onset premotor neurons) or are involved at an earlier temporal processing stage. Copyright © 1993 the American Physiological Society