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@ -2,17 +2,15 @@
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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\chapter{Spiketrain analysis}
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\chapter{Spiketrain analysis}
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\selectlanguage{english}
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\enterm[Action potentials]{action potentials} (\enterm{spikes}) are
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the carriers of information in the nervous system. Thereby it is the
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\enterm[Actionspotentials]{Actionspotentials} (\enterm{spikes}) are
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time at which the spikes are generated that is of importance for
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the carriers of information in the nervous system. Thereby it is
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information transmission. The waveform of the action potential is
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mainly the time at which the spikes are generated that is of
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largely stereotyped and therefore does not carry information.
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importance. The waveform of the action potential is largely
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stereotyped and does not carry information.
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The result of the pre-processing of electrophysiological recordings are
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series of spike times, which are termed \enterm{spiketrains}. If
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The result of the processing of electrophysiological recordings are
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measurements are repeated we get several \enterm{trials} of
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series of spike times, which are then termed \enterm{spiketrains}. If
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measurements are repeated we yield several \enterm{trials} of
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spiketrains (\figref{rasterexamplesfig}).
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spiketrains (\figref{rasterexamplesfig}).
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Spiketrains are times of events, the action potentials. The analysis
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Spiketrains are times of events, the action potentials. The analysis
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@ -21,34 +19,32 @@ of these leads into the realm of the so called \enterm[point
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\begin{figure}[ht]
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\begin{figure}[ht]
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\includegraphics[width=1\textwidth]{rasterexamples}
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\includegraphics[width=1\textwidth]{rasterexamples}
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\titlecaption{\label{rasterexamplesfig}Raster-plot.}{Raster-plot of
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\titlecaption{\label{rasterexamplesfig}Raster-plot.}{Raster-plots of
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ten realizations of a stationary point process (homogeneous point
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ten trials of data illustrating the times of the action
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process with a rate $\lambda=20$\;Hz, left) and an inhomogeneous
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potentials. Each vertical dash illustrates the time at which an
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point process (perfect integrate-and-fire neuron dirven by
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action potential was observed. Each line displays the events of
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Ohrnstein-Uhlenbeck noise with a time-constant $\tau=100$\,ms,
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one trial. Shown is a stationary point process (left, homogeneous
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right). Each vertical dash illustrates the time at which the
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point process with a rate $\lambda=20$\;Hz, left) and an
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action potential was observed. Each line represents the event of
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non-stationary point process (right, perfect integrate-and-fire
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each trial.}
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neuron dirven by Ohrnstein-Uhlenbeck noise with a time-constant
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$\tau=100$\,ms, right).}
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\end{figure}
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\end{figure}
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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\section{Point processes}
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\section{Point processes}
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A temporal \enterm{point process} is a stochastic process that
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generates a sequence of events at times $\{t_i\}$, $t_i \in
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\reZ$.
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\begin{ibox}{Examples of point processes}
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\begin{ibox}{Examples of point processes}
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Every point process is generated by a temporally continuously
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Every point process is generated by a temporally continuously
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developing process. An event is generated whenever this process
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developing process. An event is generated whenever this process
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reaches a certain threshold. For example:
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crosses some threshold. For example:\vspace{-1ex}
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\begin{itemize}
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\begin{itemize}
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\item Action potentials/heart beat: created by the dynamics of the
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\item Action potentials/heart beat: created by the dynamics of the
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neuron/sinoatrial node
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neuron/sinoatrial node
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\item Earthquake: defined by the dynamics of the pressure between
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\item Earthquake: defined by the dynamics of the pressure between
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tectonical plates.
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tectonical plates.
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\item Evoked communication calls in crickets/frogs/birds: shaped by
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\item Communication calls in crickets/frogs/birds: shaped by
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the dynamics of nervous system and the muscle appartus.
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the dynamics of the nervous system and the muscle appartus.
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\end{itemize}
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\end{itemize}
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\end{ibox}
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\end{ibox}
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@ -60,43 +56,51 @@ generates a sequence of events at times $\{t_i\}$, $t_i \in
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$T_i=t_{i+1}-t_i$ and the number of events $n_i$. }
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$T_i=t_{i+1}-t_i$ and the number of events $n_i$. }
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\end{figure}
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\end{figure}
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In the neurosciences, the statistics of point processes is of
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A temporal \enterm{point process} is a stochastic process that
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importance since the timing of the neuronal events (the action
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generates a sequence of events at times $\{t_i\}$, $t_i \in \reZ$. In
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potentials) is crucial for information transmission and can be treated
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the neurosciences, the statistics of point processes is of importance
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as such a process.
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since the timing of neuronal events (action potentials, post-synaptic
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potentials, events in EEG or local-field recordings, etc.) is crucial
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Point processes can be described using the intervals between
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for information transmission and can be treated as such a process.
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successive events $T_i=t_{i+1}-t_i$ and the number of observed events
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within a certain time window $n_i$ (\figref{pointprocessscetchfig}).
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The events of a point process can be illustrated by means of a raster
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plot in which each vertical line indicates the time of an event. The
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The events originating from a point process can be illustrated in form
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event from two different point processes are shown in
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of a scatter- or raster plot in which each vertical line indicates the
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\figref{rasterexamplesfig}. Point processes can be described using
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time of an event. The event from two different point processes are
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the intervals between successive events $T_i=t_{i+1}-t_i$ and the
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shown in \figref{rasterexamplesfig}.
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number of observed events within a certain time window $n_i$
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(\figref{pointprocessscetchfig}).
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\begin{exercise}{rasterplot.m}{}
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Implement a function \code{rasterplot()} that displays the times of
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action potentials within the first \code{tmax} seconds in a raster
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plot. The spike times (in seconds) recorded in the individual trials
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are stored as vectors of times within a \codeterm{cell-array}.
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\end{exercise}
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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\section{Interval statistics}
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\section{Interval statistics}
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The intervals $T_i=t_{i+1}-t_i$ between successive events are real
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The intervals $T_i=t_{i+1}-t_i$ between successive events are real
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positive numbers. In the context of action potentials they are
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positive numbers. In the context of action potentials they are
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referred to as \enterm{interspike intervals}. The statistics of these
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referred to as \enterm{interspike intervals}. The statistics of
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are described using the common measures.
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interspike intervals are described using common measures for
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describing the statistics of stochastic real-valued variables:
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\begin{figure}[t]
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\begin{figure}[t]
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\includegraphics[width=0.96\textwidth]{isihexamples}\vspace{-2ex}
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\includegraphics[width=0.96\textwidth]{isihexamples}\vspace{-2ex}
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\titlecaption{\label{isihexamplesfig}Interspike interval
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\titlecaption{\label{isihexamplesfig}Interspike interval
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histogram}{of the spikes depicted in \figref{rasterexamplesfig}.}
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histograms}{of the spike trains shown in \figref{rasterexamplesfig}.}
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\end{figure}
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\end{figure}
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\begin{exercise}{isis.m}{}
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\begin{exercise}{isis.m}{}
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Implement a function \code{isis()} that calculates the interspike
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Implement a function \code{isis()} that calculates the interspike
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intervals from several spike trains. The function should return a
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intervals from several spike trains. The function should return a
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single vector of intervals. The action potentials recorded in the
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single vector of intervals. The spike times (in seconds) of each
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individual trials are stored as vectors of spike times within a
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trial are stored as vectors within a \codeterm{cell-array}.
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\codeterm{cell-array}. Spike times are given in seconds.
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\end{exercise}
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\end{exercise}
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\subsection{First order interval statistics}
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%\subsection{First order interval statistics}
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\begin{itemize}
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\begin{itemize}
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\item Probability density $p(T)$ of the intervals $T$
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\item Probability density $p(T)$ of the intervals $T$
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(\figref{isihexamplesfig}). Normalized to $\int_0^{\infty} p(T) \; dT
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(\figref{isihexamplesfig}). Normalized to $\int_0^{\infty} p(T) \; dT
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@ -138,10 +142,17 @@ and non-stationary return maps are distinctly different
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\begin{figure}[t]
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\begin{figure}[t]
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\includegraphics[width=1\textwidth]{returnmapexamples}
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\includegraphics[width=1\textwidth]{returnmapexamples}
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\includegraphics[width=1\textwidth]{serialcorrexamples}
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\includegraphics[width=1\textwidth]{serialcorrexamples}
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\titlecaption{\label{returnmapfig}Interspike interval analyses of a
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\titlecaption{\label{returnmapfig}Interspike interval
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stationary and a non-stationary pointprocess.}{Upper plots show the
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correlations}{of the spike trains shown in
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return maps and the lower panels depict the serial correlation of
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\figref{rasterexamplesfig}. Upper panels show the return maps and
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successive intervals separated by the lag $k$.}
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lower panels the serial correlations of successive intervals
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separated by lag $k$. All the interspike intervals of the
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stationary spike trains are independent of each other --- this is
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a so called \enterm{renewal process}. In contrast, the ones of the
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non-stationary spike trains show positive correlations that decay
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for larger lags. The positive correlations in this example are
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caused by a common stimulus that slowly increases and decreases
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the mean firing rate of the spike trains.}
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\end{figure}
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\end{figure}
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Such dependencies can be further quantified using the \enterm{serial
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Such dependencies can be further quantified using the \enterm{serial
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@ -170,17 +181,18 @@ with itself and is always 1.
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% \includegraphics[width=0.48\textwidth]{poissoncounthist100hz100ms}
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% \includegraphics[width=0.48\textwidth]{poissoncounthist100hz100ms}
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% \titlecaption{\label{countstatsfig}Count Statistik.}{}
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% \titlecaption{\label{countstatsfig}Count Statistik.}{}
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% \end{figure}
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% \end{figure}
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The number of events $n_i$ (counts) in a time window $i$ of the duration $W$
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Counting the number of events $n_i$ (counts) in time windows $i$ of duration $W$
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yields positive integer random numbers that are commonly quantified
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yields positive integer random numbers that are commonly quantified
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using the following measures:
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using the following measures:
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\begin{itemize}
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\begin{itemize}
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\item Histogram of the counts $n_i$.
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\item Histogram of the counts $n_i$.
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\item Average number of events: $\mu_N = \langle n \rangle$.
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\item Average number of counts: $\mu_n = \langle n \rangle$.
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\item Variance of the counts: $\sigma_n^2 = \langle (n - \langle n \rangle)^2 \rangle$.
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\item Variance of counts: $\sigma_n^2 = \langle (n - \langle n \rangle)^2 \rangle$.
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\item \determ{Fano Faktor} (The variance divided by the average): $F = \frac{\sigma_n^2}{\mu_n}$.
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\item \determ{Fano Factor} (variance of counts divided by average count): $F = \frac{\sigma_n^2}{\mu_n}$.
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\end{itemize}
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\end{itemize}
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And in particular the average firing rate $r$ (spike count per time interval
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Of particular interest is the average firing rate $r$ (spike count per
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, \determ{Feuerrate}) that is given in Hertz \sindex[term]{Feuerrate!mittlere Rate}
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time interval , \determ{Feuerrate}) that is given in Hertz
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\sindex[term]{firing rate!average rate}
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\begin{equation}
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\begin{equation}
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\label{firingrate}
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\label{firingrate}
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r = \frac{\langle n \rangle}{W} \; .
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r = \frac{\langle n \rangle}{W} \; .
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@ -204,30 +216,30 @@ And in particular the average firing rate $r$ (spike count per time interval
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the distribution of spike counts observed in a certain time
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the distribution of spike counts observed in a certain time
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window. The function should take two input arguments: (i) a
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window. The function should take two input arguments: (i) a
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\codeterm{cell-array} of vectors containing the spike times in
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\codeterm{cell-array} of vectors containing the spike times in
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seconds observed in a number of trials and (ii) the duration of the
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seconds observed in a number of trials, and (ii) the duration of the
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time window that is used to evaluate the counts.
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time window that is used to evaluate the counts.\pagebreak[4]
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\end{exercise}
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\end{exercise}
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%%
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\section{Homogeneous Poisson process}
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\section{Homogeneous Poisson process}
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The Gaussian distribution is, due to the central limit theorem, the
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The Gaussian distribution is, because of the central limit theorem,
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standard for continuous measures. The equivalent in the realm of point
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the standard distribution for continuous measures. The equivalent in
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processes is the \enterm{Poisson distribution}.
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the realm of point processes is the \enterm{Poisson distribution}.
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In a \enterm[Poisson process!homogeneous]{homogeneous Poisson process}
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In a \enterm[Poisson process!homogeneous]{homogeneous Poisson process}
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the events occur at a fixed rate $\lambda=\text{const.}$ and are
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the events occur at a fixed rate $\lambda=\text{const}$ and are
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independent of both the time $t$ and occurrence of previous events
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independent of both the time $t$ and occurrence of previous events
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(\figref{hompoissonfig}). The probability of observing an even within a
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(\figref{hompoissonfig}). The probability of observing an event within
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small time window of width $\Delta t$ is given by
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a small time window of width $\Delta t$ is given by
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\begin{equation}
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\begin{equation}
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\label{hompoissonprob}
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\label{hompoissonprob}
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P = \lambda \cdot \Delta t \; .
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P = \lambda \cdot \Delta t \; .
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\end{equation}
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\end{equation}
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In an \enterm[Poisson process!inhomogeneous]{inhomogeneous Poisson
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In an \enterm[Poisson process!inhomogeneous]{inhomogeneous Poisson
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process}, however, the rate $\lambda$ depends on the time: $\lambda =
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process}, however, the rate $\lambda$ depends on time: $\lambda =
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\lambda(t)$.
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\lambda(t)$.
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\begin{exercise}{poissonspikes.m}{}
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\begin{exercise}{poissonspikes.m}{}
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@ -249,7 +261,11 @@ In an \enterm[Poisson process!inhomogeneous]{inhomogeneous Poisson
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\begin{figure}[t]
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\begin{figure}[t]
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\includegraphics[width=0.45\textwidth]{poissonisihexp20hz}\hfill
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\includegraphics[width=0.45\textwidth]{poissonisihexp20hz}\hfill
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\includegraphics[width=0.45\textwidth]{poissonisihexp100hz}
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\includegraphics[width=0.45\textwidth]{poissonisihexp100hz}
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\titlecaption{\label{hompoissonisihfig}Distribution of interspike intervals of two Poisson processes.}{}
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\titlecaption{\label{hompoissonisihfig}Distribution of interspike
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intervals of two Poisson processes.}{The processes differ in their
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rate (left: $\lambda=20$\,Hz, right: $\lambda=100$\,Hz). The red
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lines indicate the corresponding exponential interval distribution
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\eqnref{poissonintervals}.}
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\end{figure}
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\end{figure}
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The homogeneous Poisson process has the following properties:
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The homogeneous Poisson process has the following properties:
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@ -267,7 +283,10 @@ The homogeneous Poisson process has the following properties:
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process is also called a \enterm{renewal process}.
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process is also called a \enterm{renewal process}.
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\item The number of events $k$ within a temporal window of duration
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\item The number of events $k$ within a temporal window of duration
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$W$ is Poisson distributed:
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$W$ is Poisson distributed:
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\[ P(k) = \frac{(\lambda W)^ke^{\lambda W}}{k!} \]
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\begin{equation}
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\label{poissoncounts}
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P(k) = \frac{(\lambda W)^ke^{\lambda W}}{k!}
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\end{equation}
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(\figref{hompoissoncountfig})
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(\figref{hompoissoncountfig})
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\item The Fano Faktor is always $F=1$ .
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\item The Fano Faktor is always $F=1$ .
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\end{itemize}
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\end{itemize}
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@ -286,8 +305,11 @@ The homogeneous Poisson process has the following properties:
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\begin{figure}[t]
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\begin{figure}[t]
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\includegraphics[width=0.48\textwidth]{poissoncounthistdist100hz10ms}\hfill
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\includegraphics[width=0.48\textwidth]{poissoncounthistdist100hz10ms}\hfill
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\includegraphics[width=0.48\textwidth]{poissoncounthistdist100hz100ms}
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\includegraphics[width=0.48\textwidth]{poissoncounthistdist100hz100ms}
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\titlecaption{\label{hompoissoncountfig}Count statistics of Poisson
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\titlecaption{\label{hompoissoncountfig}Distribution of counts of a
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spiketrains.}{}
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Poisson spiketrain.}{The count statistics was generated for two
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different windows of width $W=10$\,ms (left) and width $W=100$\,ms
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(right). The red line illustrates the corresponding Poisson
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distribution \eqnref{poissoncounts}.}
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\end{figure}
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\end{figure}
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@ -312,7 +334,7 @@ closely.
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\begin{figure}[tp]
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\begin{figure}[tp]
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\includegraphics[width=\columnwidth]{firingrates}
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\includegraphics[width=\columnwidth]{firingrates}
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\titlecaption{Estimating the time-dependent firing
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\titlecaption{Estimating the time-dependent firing
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rate.}{\textbf{A)} Rasterplot depicting the spiking activity of a
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rate.}{\textbf{A)} Rasterplot showing the spiking activity of a
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neuron. \textbf{B)} Firing rate calculated from the
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neuron. \textbf{B)} Firing rate calculated from the
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\emph{instantaneous rate}. \textbf{C)} classical PSTH with the
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\emph{instantaneous rate}. \textbf{C)} classical PSTH with the
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\emph{binning} method. \textbf{D)} Firing rate estimated by
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\emph{binning} method. \textbf{D)} Firing rate estimated by
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@ -529,5 +551,3 @@ kernel.
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the same size as the original stimulus contained in file
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the same size as the original stimulus contained in file
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\file{sta\_data.mat}.
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\file{sta\_data.mat}.
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\end{exercise}
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\end{exercise}
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\selectlanguage{english}
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