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Schuch- uniformly 0. Fisahn, E. Buhl, R. Dermietzel, U. Heinemann, the original paper Traub et al. Traub, unpublished data have obtained electrophysiolog- in the present study was as follows: 0. Why then would oscillations will be duly noted. This question was addressed by Traub et al.
This means that other, on average. By trial and error, either isolated or lie on small clusters. With low-connection densities, potentials. These could occur at low frequency as low as 0. A cluster is in some cases. For example, the large cluster has or by synaptic time constants, but rather by 1 topological struc- cells, and the next largest cluster is small, having only 13 cells.
The mean ture of the network and 2 the time for a spike in one axon to path length which measures the average length of the shortest gap induce a spike in a connected axon. A block diagram of the network model is shown in Figure 1. In the added As in previous studies Traub et al. Here, we and occurred once per 5 sec in interneurons; in pyramidal cells, in the illustrate how ripple oscillations might also be generated.
In our model of high-frequency oscillations in axonal networks Traub et al. The network model used in this paper is a hybrid of the models described Sharp waves were produced by applying a msec excitatory conduc- by Traub et al.
The slight differences are listed below. The shape of the conductance pulse defined f urther on. Further variability was introduced into the system by rons. Each neuron is multicompartmental and includes five axonal com- having different bias currents applied to the somata of the pyramidal cells partments Traub et al. The interneurons are divided into four classes of 96 cells each: msec.
Block diagram of network model. The model contains pyramidal cells and interneurons basket cells, axo-axonic cells, and dendrite-contacting cells , with intrinsic properties and connectivity de- scribed in Materials and Methods.
The figure highlights the conceptual distinction we make between the axons of the pyramidal cells and the soma—dendritic membranes. Interactions between components of the model are shown. Afferent inputs in the model are purely excitatory. For the sake of consistency, figures were made using data from the second sharp wave of any given simulation. Simulation programs saved the following types of data: somatic volt- ages of selected pyramidal cells some on the large cluster, some not and interneurons; voltages in the axon at the site of the gap junction of selected pyramidal cells; average signals, consisting of somatic voltages of nearby pyramidal cells or of 28 nearby interneurons; and total Figure 2.
Firing properties of pyramidal cells and interneurons during a GABAA synaptic conductance to a pyramidal cell, recurrently generated simulated ripple. A, Average voltage of nearby e-cell somata thick line reveals a pyramidal cell, and afferent AM PA conductance to an interneuron.
An inter- Autocorrelations and cross-correlations were computed using the middle neuron thin line fires at the same frequency. Spikes truncated. B, Single 75 msec of data from the simulated sharp wave.
The axon of the cell thin line gap junction. Axons of cells off the structions for a parallel computer. These latter programs were run on an large cluster do not fire at ripple frequency, and there are virtually no I BM SP2 with 12 nodes processors ; other programs ran on a single spikelets, although intracellular rippling still occurs data not shown. Some comments on numerical methods are discussed by Spikes truncated.
Signals in A and B are simultaneous. C, Autocorrelation Traub et al.
For f urther details, please contact Roger D. Traub at Hz. Cross-correlation of local average pyramidal cells with local average r. The basic properties of simulated ripples in our model using default parameters are shown in Figures 2 and 3. The afferent waves, distal axons are hyperpolarized by several millivolts, which input depolarizes the pyramidal cells and, to a lesser extent, the suppresses the ectopic spikes data not shown.
As the sharp wave interneurons Fig. The detailed oscillation is coherent Fig. During the sharp wave- mechanisms of this oscillation are analyzed by Traub et al. The data not shown. The period of the oscillation is Rippling arises in our model as follows.
Hyperpolarizing a pyramidal cell, during a ripple, shifts its phase relative to the local average signal.
Using data from the central portion of the ripple in the simulation of Figures 2 and 3, a pyramidal cell somatic signal was cross-correlated with the local average signal pyramidal cells thick line. The peak is near 0 msec 0. The simulation was repeated, with the index pyramidal cell hyperpolarized Figure 3.
The cross-correlation of the oscillates at high frequency and phasically drives interneurons. A, Inter- index cell with the local average thin line now reveals a minimum near neuron thin line, spikes truncated fires at ripple frequency. The phasic waves of synaptic excitation lead the action poten- local average pyramidal signal, the peak occurred near 0 msec tials.
When the same cell was hyperpolarized tory output axonal network thin line , plotted as the number of axons depolarized above 70 mV from rest.
Data from simulation the local average signal Fig. Ylinen et al. This is consistent with the notion that the ripple waves are predominantly IPSPs. The imposed on the afferent excitatory conductance. In contrast, fluc- oscillation period is approximately the crossing time multiplied by tuations in GABAA receptor conductance were 25—50 nS. There the expected length of such chains. Our model predicts that small was, additionally, a slower component to the GABAA conduc- decreases in junctional conductance, by prolonging the time it tance, produced by the dendrite-contacting interneurons whose takes for spikes to cross gap junctions, should slow the period, ISPCs are relatively slow; see Materials and Methods.
The be sufficiently large, consistent with in vivo data showing that latter effect is expected because, in a more connected network, an halothane suppresses ripples Ylinen et al. First, antidromic propagation into pyramidal cells lying on orthodromic. When Fig. With the strength of IPSCs used, however, the gap junctional conductance was reduced only 2. Spikelets are also [Note that for pyramidal cells not lying on the large cluster, to be noted in the pyramidal cell soma.
Finally, for comparison, spikelets will not be seen; instead, the ripple appears purely as Figure 5C illustrates rippling, using data from the simulation of synaptic potentials data not shown.
When the gap junctional conductance was in- orthodromic output of the axonal plexus induces, after a very creased an additional Note that ortho- Traub et al.
Blocking GABAA receptors allows a ripple frequency oscilla- tion to persist, but pyramidal cell firing increases. Figure 5. Oscillatory behavior in the model depends on axo-axonal gap Traces in A—C are simultaneous. Cells on the large cluster as in C exhibit junction conductance.
Each panel shows the soma of a pyramidal cell 10 —20 mV partial spikes arrows , deriving from partially blocked anti- lying on the large cluster thick line and the axon of the cell thin line.
Many of the full action potentials are also antidromic; the Spikes are truncated. A, With a pyramidal cells antidromically, as well as orthodromically. Compare the small gap junctional conductance, the axonal network cannot sustain high-frequency oscillation superimposed on CA1 epileptiform field po- high-frequency oscillations.
B, At a higher gap junction conductance, there is evidence of sustained axonal network activity; the soma fires only once, but the axon fires seven times, and spikelets occur in the soma.
Individual pyramidal cells express a high- not shown , and pyramidal cells Fig. That axonal activity frequency oscillation as well, with a synaptic component, and, if the cell is on the large cluster, spikelets occur also. This is a type of event that occurs in to rippling as the conductance of pyramidal IPSCs is reduced. Figure 6 demonstrates Other parameters are as in the simulation of Fig.
This type of activity occurs in a given briefly four possible models of rippling and attempt to identify cell provided it lies on the large cluster at the same frequency as experiments that could distinguish between the different models. Of course, these models could possibly work in conjunction with Rippling in the model also requires that IPSCs on pyramidal one another in various combinations, but it is premature to enter cells not be excessively large.
Figure 7 illustrates a simulation with into the resulting complexities. Africa and the Middle East; Hyoscyamus niger L.
Schmidt, H. Koch, H. Tallos de hasta 50 cm, ramificados. Corola de mm, dividida hasta c.
Estambres exertos, con filamentos vilosos, al menos en la mitad inferior. Semillas de ,4 x 0,,2 mm, alveolado-crestadas, grises. Florece de Enero a Mayo. Flora Iberica: Hierba perenne, bienal o anual, densamente pubescente, con pelos pluricelulares largos, blancos y pelos cortos glandulosos y eglandulosos. Tallos hasta de 80 cm, ramificados, tomentosos.