Responses of sagittally aligned Purkinje cells during perturbed locomotion: relation of climbing fiber activation to simple spike modulation

1992 ◽  
Vol 68 (5) ◽  
pp. 1820-1833 ◽  
Author(s):  
J. S. Lou ◽  
J. R. Bloedel

1. The purpose of these experiments is to test the hypothesis that the synchronous activation of sagittally aligned Purkinje cells by a physiologically relevant stimulus is associated with an increase in the simple spike responses of the same neurons. 2. This hypothesis was tested using a perturbed locomotion paradigm in decerebrate locomoting ferrets. The responses of 3-5 sagittally aligned Purkinje cells were recorded simultaneously in response to the intermittent perturbation of the forelimb during swing phase. A data analysis is introduced, the real time postsynaptic response (RTPR), that permits the quantification of the simple spike responses of Purkinje cells in a manner that can be related to their complex spike responses on a trial-by-trial basis. 3. The data support the above hypothesis by illustrating that the amplitude of the combined simple spike responses across the population of Purkinje cells is correlated with the extent to which their climbing fiber inputs are synchronously activated. These findings together with an analysis of the gain-change ratio support the view that the synchronous climbing fiber input may be responsible for mediating this increased responsiveness. 4. More generally, the data suggest that the task- and/or behaviorally dependent activation of sagittal strips of climbing fiber inputs may provide a mechanism whereby the responsiveness of discrete populations of Purkinje cells can be selectively regulated, specifying the groups of neurons that will be most dramatically modulated by mossy fiber inputs activated by the same conditions.

1995 ◽  
Vol 74 (6) ◽  
pp. 2573-2589 ◽  
Author(s):  
N. H. Barmack ◽  
H. Shojaku

1. The cerebellar uvula-nodulus receives vestibular projections from primary and secondary vestibular afferents as well as vestibularly related climbing fibers. It also receives visually related information from climbing fiber pathways. In this experiment we investigated how this information is mapped onto the uvula-nodulus. We studied the specificity, dynamics, and topographic distribution of climbing fiber responses (CFRs), simple spike responses, and mossy fiber terminal responses evoked by vestibular and optokinetic stimulation in rabbits anesthetized with alpha-chloralose. 2. Vestibularly evoked CFRs were found in the ventral uvula and nodulus. These responses were evoked during static roll tilt of the rabbit about a longitudinal axis and by sinusoidal oscillation about the longitudinal axis. Purely static responses were attributed to stimulation of the utricular otolith by the linear acceleration of gravity. CFRs that lacked a static component were attributed to activation of the semicircular canals. 3. Using a "null technique" we showed that the canal-sensitive CFRs were caused by stimulation of the anterior or posterior semicircular canals. Of the CFRs classified as canal related, 96% could be attributed to stimulation of the vertical semicircular canals. 4. Increases in CFRs were correlated with decreases in simple spike responses in half the Purkinje cells from which we recorded. These climbing-fiber-induced pauses in simple spikes occurred during spontaneous climbing fiber discharge as well as during climbing fiber discharge evoked by vestibular stimulation. The duration of this pause was inversely proportional to the spontaneous level of simple spikes before the occurrence of a CFR. In the other half of the recorded population of Purkinje cells, vestibularly driven CFRs did not alter the simple spike responses. 5. Vestibularly and visually mediated CFRs were topographically represented on the surface of the uvula-nodulus. CFRs driven by ipsilateral otolithic inputs were distributed over the entire mediolateral surface of the uvula-nodulus. CFRs driven by the ipsilateral posterior semicircular canal were distributed in a sagittal strip approximately 1.5 mm wide, extending laterally from the midline of the nodulus. CFRs driven exclusively by horizontal, posterior-->anterior optokinetic stimulation of the ipsilateral eye were distributed in a sagittal strip approximately 0.5 mm wide located 0.5-1.0 mm from the midline and restricted to the ventral nodulus. CFRs driven by the ipsilateral anterior semicircular canal were found in a sagittal strip approximately 1.0 mm wide extending 1.0-2.0 mm from the midline. 6. The sagittal, topographically arrayed climbing fiber strips effectively map a mediolateral gradient of possible postural responses based on vestibular and optokinetic information.


1983 ◽  
Vol 50 (1) ◽  
pp. 205-219 ◽  
Author(s):  
T. J. Ebner ◽  
Q. X. Yu ◽  
J. R. Bloedel

These experiments were designed to test the hypothesis that climbing fiber inputs evoked by a peripheral stimulus increase the responsiveness of Purkinje cells to mossy fiber inputs. This hypothesis was based on a previous series of observations demonstrating that spontaneous climbing fiber inputs are associated with an accentuation of the Purkinje cell responses to subsequent mossy fiber inputs (10, 12). Furthermore, short-term nonpersistent interactions between climbing and mossy fiber inputs have been an important aspect of many theories of cerebellar function (5, 7, 8, 12, 36). Extracellular unitary recordings were made from Purkinje cells in lobule V of decerebrate, unanesthetized cats. To activate mossy and climbing fiber inputs, the forepaw was passively flexed by a Ling vibrator system. A data analysis was developed to sort the simple spike trials into two groups, based on the presence or absence of complex spikes activated by the stimulus. In addition, during those trials in which complex spikes were activated, the simple spike train was aligned on the occurrence of the complex spike. For each simple spike response to the forepaw input, the average firing rate during the response was compared to background both in those trials in which complex spikes were activated and in those in which they were not. The ratio of the response amplitudes in the histograms constructed from these two groups of trials permitted a quantification of the change in responsiveness when climbing fiber inputs were activated. The results show that both excitatory and inhibitory simple spike responses are accentuated when associated with the activation of a complex spike. Using an arbitrary level of a gain change ratio of 120% as indicating a significant modification, 64% of the response components analyzed increased their amplitude when climbing fiber input was present. Simple spike response components occurring prior to complex spike activation were usually not accentuated, although in a few cells the amplitude of this component of the response increased. In addition, in a small number of cells the occurrence of complex spikes was associated with a new simple spike component. For excitatory responses, the magnitude of the gain change ratio was shown to be inversely related to the amplitude of the simple spike response evoked by the mossy fiber inputs. The data obtained is consistent with the hypothesis that the climbing fiber input is associated with an increase in the responsiveness of Purkinje cells to mossy fiber inputs. The increased responsiveness occurs whether the simple spike modulation evoked by the peripheral stimulus is excitatory or inhibitory. The change in responsiveness is short term and nonpersistent. It is argued that the activation of climbing fiber inputs to the cerebellar cortex is associated with an increase in the gain of Purkinje cells to mossy fiber inputs activated by natural peripheral stimuli.


1993 ◽  
Vol 69 (4) ◽  
pp. 1082-1090 ◽  
Author(s):  
Y. Sato ◽  
A. Miura ◽  
H. Fushiki ◽  
T. Kawasaki

1. Some scientists reported that the simple spike (SS) activity was transiently depressed after climbing fiber input, but others reported that predominant population of Purkinje cells increased their SS activity after the complex spike (CS). In the present study, SS activity after spontaneous CS was compared before and after the administration of pentobarbital sodium and of ketamine in high decerebrate cats. 2. Frequencies of spontaneous CS and SS firing were reduced (P < 0.001, t test) after pentobarbital administration of a total dose of 20-30 mg/kg. 3. In the peri-CS time histogram, the SS activity during a post-CS period of 10-110 ms with respect to that during a pre-CS period of -100-0 ms was reduced (P < 0.001) after the pentobarbital administration from, on average, 132.4 to 81.9%. In contrast, the SS activity during a post-CS period of 110-210 ms remained unchanged (P > 0.2). 4. In the pre-CS time histogram constructed after the pentobarbital administration, there were no significant differences (P > 0.01) between the SS activity during a pre-CS period of -600 to -500 ms and that during each of other pre-CS periods, suggesting that the barbiturate had little effect on the SS activity preceding the CS. 5. Analysis of raster diagrams revealed the variability of individual SS activity after the CS.(ABSTRACT TRUNCATED AT 250 WORDS)


1994 ◽  
Vol 71 (1) ◽  
pp. 401-419 ◽  
Author(s):  
E. De Schutter ◽  
J. M. Bower

1. Both excitatory and inhibitory postsynaptic channels were added to a previously described complex compartmental model of a cerebellar Purkinje cell to examine model responses to synaptic inputs. All model parameters remained as described previously, leaving maximum synaptic conductance as the only parameter that was tuned in the studies described in this paper. Under these conditions the model was capable of reproducing physiological recorded responses to each of the major types of synaptic input. 2. When excitatory synapses were activated on the smooth dendrites of the model, the model generated a complex dendritic Ca2+ spike similar to that generated by climbing fiber inputs. Examination of the model showed that activation of P-type Ca2+ channels in both the smooth and spiny dendrites augmented the depolarization during the complex spike and that Ca(2+)-activated K+ channels in the same dendritic regions determined the duration of the spike. When these synapses were activated under simulated current-clamp conditions the model also generated the characteristic dual reversal potential of the complex spike. The shape of the dendritic complex spike could be altered by changing the maximum conductance of the climbing fiber synapse and thus the amount of Ca2+ entering the cell. 3. To explore the background simple spike firing properties of Purkinje cells in vivo we added excitatory “parallel fiber” synapses to the spiny dendritic branches of the model. Continuous asynchronous activation of these granule cell synapses resulted in the generation of spontaneous sodium spikes. However, very low asynchronous input frequencies produced a highly regular, very fast rhythm (80–120 Hz), whereas slightly higher input frequencies resulted in Purkinje cell bursting. Both types of activity are uncharacteristic of in vivo Purkinje cell recordings. 4. Inhibitory synapses of the sort presumably generated by stellate cells were also added to the dendritic tree. When asynchronous activation of these inhibitory synapses was combined with continuous asynchronous excitatory input the model generated somatic action potentials in a much more stochastic pattern typical of real Purkinje cells. Under these conditions simulated inter-spike interval distributions resembled those found in experimental recordings. Also, as with in vivo recordings, the model did not generate dendritic bursts. This was mainly due to inhibition that suppressed the generation of dendritic Ca2+ spikes. 5. In the presence of asynchronous inhibition, changes in the average frequency of excitatory inputs modulated background simple spike firing frequencies in the natural range of Purkinje cell firing frequencies (30–100 Hz). This modulation was very sensitive to small changes in the average frequency of excitatory inputs.(ABSTRACT TRUNCATED AT 400 WORDS)


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