Relationship between place code and temporal coded

Published в Not reliable connection csgo betting | Октябрь 2, 2012

relationship between place code and temporal coded

interaural time difference, frequency modulation, non-sensory factors (i.e., rate-place vs temporal coding) of cochlear neurons. The basic idea of the temporal code is as simply stated as that of the rate code: information about the stimulus or action is contained in the. Together with the fact that unambiguous theoretical links can be made relating rate-place vs temporal coding) of cochlear neurons. SPORTS BETTING PROFESSOR NBA SYSTEM PLAYS

Illustration Tab On the Illustration tab, you can adjust these parameters: Play the Sound: check to start the animation and uncheck to stop the animation. Show Vertical Movement: when only a single hair cell is shown, allow or not the vertical motion of the hair cell that results from the basilar membrane. Show Rotation: when only a single hair cell is shown, show the rotation of the hair cell that results from the basilar membrane.

Frequency Hz : adjust to see how the cochlea and basilar membrane respond to different frequencies. Reset Pressing this button restores the settings to their default values and allows you to adjust speed and relative size. C Top, black dots represent all spikes of the cell in A as a function of position in cm x-axis and theta phase y-axis. Middle, scheme of spatial normalization.

Light gray dots represent spikes. The center of the place field was defined as the position of maximum firing rate. The orange horizontal scale on top shows the distance to the place field center in units of place field length.

Arrows highlight the relative distance of two spikes. Bottom, heat map representation of spike counts per relative distance and theta phase. In these maps, the phase distribution of place cell spiking cannot be properly visualized at space bins distant from the place field center because of their much lower spike count Fig.

To circumvent this, we normalized spike counts within each space bin a column slice by its mean over all phases Fig. This space-normalization allowed for assessing the phase distribution of spikes at positions away from the place field center.

We found that spikes concentrate near the theta peak before the animal enters the place field, at positions where place cells exhibit no major changes in firing rate compare Fig. Consistently, Fig. Notice that firing rate is comparably low before and after the place field, suggesting similar excitatory drive to place cells despite different coupling to theta at these regions. Figure 2 Place cells couple to theta phase before increasing their firing rate.

B Zoomed-out view of the data in A. C Mean firing rate as a function of relative distance. D Same data as in B , normalized by the mean number of spikes at each position. E Spike-phase coupling strength per relative distance solid line ; the dashed line reproduces the firing rate in C.

Notice that place cell spikes align to a preferred theta phase before major increases in firing rate arrowhead. Full size image To assess whether the results could be due to differences in spike counts among place fields, we next calculated the mean spiking phase for each place field per bin of relative distance. Figure 3A shows the spatial histogram of mean spiking phases.

Columns with non-uniform phase distributions indicate that place cells have similar mean spiking phase at these positions, with warmer colors denoting preferred theta phases. We next computed theta-phase coupling strength TPC using the distribution of mean spiking phases, and used a surrogate procedure to delimit statistically significant values black arrows in Fig. In fact, such a phenomenon of strong theta-phase coupling before a major increase in firing rate is also apparent in individual cells; for instance, notice that the example place cell in Fig.

S1 for other examples. This result also holds true when analyzing each of the three rats separately Supplementary Fig. S2 , when estimating theta phase by linear interpolation 13 Supplementary Fig. S3 , or when analyzing the first and later spikes per theta cycle separately 14 Supplementary Fig. Interestingly, and in contrast to previous views 15 , Figs 2E and 3B also demonstrate that the coupling of place cells to theta phase is spatially transient, which is to say that TPC — as FR — has a spatial receptive field.

Figure 3 Asymmetry of the temporal code for space by hippocampal place cells. A Histogram of mean spiking phase. At each position, individual place fields contribute with one count to the phase bin in which the mean spiking of their place cell occurs. Arrows mark the region where the distribution of mean spiking phases statistically differs from the uniform distribution, indicating coordinated theta-phase coupling across cells. D Mean relative distance of TPC and FR peaks, showing that place cells are maximally modulated by theta phase before their peak firing rate.

Full size image Asymmetry of the temporal code for space To further characterize differences in temporal and rate coding, we computed the peak and the full width at half maximum FWHM of the TPC and FR curves along space Fig. Moreover, while the portion of the FR curve above its half-maximum value was roughly symmetrical around the place field center see also Fig.

These results show that hippocampal place cells have an asymmetric temporal code for space. Figure 4 Asymmetric temporal coding irrespective of place field skewness. A Example place field. B Skewness of the firing rate curve at each threshold. C Percentage of the place field located before the center position of peak firing rate.

D Panels show the same as in Fig. Full size image It has been previously reported that rate-based place fields become skewed with experience: in familiar tracks, place cells spike more at locations preceding the location of their peak firing rate 16 , Importantly, however, the percentage of the spatial receptive field before the place field center was much larger for TPC than FR at all thresholds from the peak value Supplementary Fig.

S5 , which is to say that temporal coding has much greater asymmetry around the place field center than rate coding even at thresholds in which FR is negatively skewed. Interestingly, the joint dynamics of FR and TPC revealed that space coding by place cells occurs in three stages during trajectories through the place field, and that the sharp phase precession is an intermediate stage in their temporal organization Fig. In Fig.

Figure 5 Three-stage model of space coding by place cells. A Theta-phase coupling strength top , mean normalized firing rate middle and mean theta phase of spiking bottom per relative distance. Gray shaded area separates place cell activity along space into three stages, as labeled. B 2D plots of all pairwise combinations of the independent variables in A. C 3D plot of theta-phase coupling strength, firing rate and mean spiking theta phase.

Gray lines show 2D projections same curves as in B. D Mean change in theta-phase coupling strength left , mean theta phase of spiking middle , and mean change in spiking phase right during each stage. In A—C , the line color denotes the relative distance to the place field center and black dots mark stage boundaries. Full size image Figure 6 Schematic illustration of the three-stage spatial dynamics of place cell firing.

The illustration depicts the changes in firing rate and in theta-phase coupling strength as the animal crosses a place field top , as well as a schematic theta rhythm showing the probability of place cell firing within theta cycles along space bottom. Full size image Discussion Hippocampal place cells convey spatial information through both firing rate and spike timing. In the first case, firing rate increases as the animal approaches a specific location in space known as the place field of the cell 3 Fig.

In the present work, we revisited this debate by investigating the evolution of theta-phase coupling of spikes along space as the animal crosses the place field. Place cells code for space by precisely timing their firing at specific phases of the theta cycle 4 , 6 ; therefore, the study of temporal coding intrinsically relates to the study of theta-phase coupling.

Here we performed a simple, yet novel space normalization Fig. Together, the spatial changes of TPC, firing rate and spiking theta phase revealed a three-stage dynamics Figs 5 and 6. Our results show that place cell spikes are not always coupled to theta.

Rather, we found that spike-phase coupling is spatially transient during crossings of the linear track, that is, place cells only couple to theta in a bounded region of space around the place field center. Both the increase in the TPC curve and its later return to zero along space are non-trivial empirical findings, for they are not explained by the definition of the metric, nor by firing rate alone i. We also found that place cells align to theta phase much before they increase their firing rate, that is, much before the animal enters the rate-based place field Figs 2E and 3B.

This effect precedes the warm colors in phase-position firing maps i. These novel analyses show that hippocampal place cells display strong asymmetry between rate and temporal codes for space, and also within temporal coding itself with respect to the place field center. It should be noted that reliably measuring spike-phase coupling requires the sampling of several spikes 20 ; therefore, due to methodological constraints, the current results could only be achieved at the level of pooled spike counts or pooled place cell activity across theta cycles and trials.

In this sense, and contrasting to the sharp phase precession 21 , the stages of phase coupling and phase decoupling cannot be statistically inferred at the level of a single place cell activity in a single trial. Rather, the effect only becomes apparent when analyzing the same place cell across multiple traversals of its place field or when analyzing a population of place cells. Interestingly, it has been previously shown that, during phase precession of grid cells, the later spikes within theta cycles have larger phase variability than the first, leading spike Moreover, phase precession slope is steeper and better correlates with position when considering only the leading spikes It is therefore possible that the phase decoupling we observed to occur upon exiting the place field is due to the high phase variability of the later spikes.

However, this seems not to be the case, since we obtained similar results when analyzing the leading and later spikes per theta cycle separately Supplementary Fig.

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