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explained_variance

ExplainedVariance

Bases: object

Explained variance measure for assessing reactivation of neuronal activity using pairwise correlations.

References

1) Kudrimoti, H. S., Barnes, C. A., & McNaughton, B. L. (1999). Reactivation of Hippocampal Cell Assemblies: Effects of Behavioral State, Experience, and EEG Dynamics. Journal of Neuroscience, 19(10), 4090-4101. https://doi.org/10/4090 2) Tatsuno, M., Lipa, P., & McNaughton, B. L. (2006). Methodological Considerations on the Use of Template Matching to Study Long-Lasting Memory Trace Replay. Journal of Neuroscience, 26(42), 10727-10742. https://doi.org/10.1523/JNEUROSCI.3317-06.2006

Adapted from https://github.com/diba-lab/NeuroPy/blob/main/neuropy/analyses/reactivation.py

Attributes:

Name Type Description
st SpikeTrainArray

obj that holds spiketrains

template EpochArray

time in seconds, pairwise correlation calculated from this period will be compared to matching period (task-period)

matching EpochArray

time in seconds, template-correlations will be correlated with pariwise correlations of this period (post-task period)

control EpochArray

time in seconds, control for pairwise correlations within this period (pre-task period)

bin_size float

in seconds, binning size for spike counts

window int

window over which pairwise correlations will be calculated in matching and control time periods, if window is None entire time period is considered, in seconds

slideby int

slide window by this much, in seconds

matching_windows array

windows for matching period

control_windows array

windows for control period

template_corr array

pairwise correlations for template period

matching_paircorr array

pairwise correlations for matching period

control_paircorr array

pairwise correlations for control period

ev array

explained variance for each time point

rev array

reverse explained variance for each time point

ev_std array

explained variance standard deviation for each time point

rev_std array

reverse explained variance standard deviation for each time point

partial_corr array

partial correlations for each time point

rev_partial_corr array

reverse partial correlations for each time point

n_pairs int

number of pairs

matching_time array

time points for matching period

control_time array

time points for control period

ev_signal AnalogSignalArray

explained variance signal

rev_signal AnalogSignalArray

reverse explained variance signal

plot function

plot explained variance

pvalue function

calculate p-value for explained variance by shuffling the template correlations

Examples:

Load data

>>> basepath = r"U:\data\HMC\HMC1\day8"
>>> st,cm = loading.load_spikes(basepath,brainRegion="CA1",putativeCellType="Pyr")
>>> epoch_df = loading.load_epoch(basepath)
>>> beh_epochs = nel.EpochArray(epoch_df[["startTime", "stopTime"]].values)

Most simple case, returns single explained variance value

>>> expvar = explained_variance.ExplainedVariance(
>>>        st=st,
>>>        template=beh_epochs[1],
>>>        matching=beh_epochs[2],
>>>        control=beh_epochs[0],
>>>        window=None,
>>>    )

Get time resolved explained variance across entire session in 200sec bins

>>> expvar = explained_variance.ExplainedVariance(
>>>        st=st,
>>>        template=beh_epochs[1],
>>>        matching=nel.EpochArray([beh_epochs.start, beh_epochs.stop]),
>>>        control=beh_epochs[0],
>>>        window=200
>>>    )

Get time resolved explained variance across entire session in 200sec bins sliding by 100sec

>>> expvar = explained_variance.ExplainedVariance(
>>>        st=st,
>>>        template=beh_epochs[1],
>>>        matching=nel.EpochArray([beh_epochs.start, beh_epochs.stop]),
>>>        control=beh_epochs[0],
>>>        window=200,
>>>        slideby=100
>>>    )
Source code in neuro_py/ensemble/explained_variance.py
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class ExplainedVariance(object):
    """Explained variance measure for assessing reactivation of neuronal activity using pairwise correlations.

    References
    -------
    1) Kudrimoti, H. S., Barnes, C. A., & McNaughton, B. L. (1999).
        Reactivation of Hippocampal Cell Assemblies: Effects of Behavioral State, Experience, and EEG Dynamics.
        Journal of Neuroscience, 19(10), 4090-4101. https://doi.org/10/4090
    2) Tatsuno, M., Lipa, P., & McNaughton, B. L. (2006).
        Methodological Considerations on the Use of Template Matching to Study Long-Lasting Memory Trace Replay.
        Journal of Neuroscience, 26(42), 10727-10742. https://doi.org/10.1523/JNEUROSCI.3317-06.2006

    Adapted from https://github.com/diba-lab/NeuroPy/blob/main/neuropy/analyses/reactivation.py

    Attributes
    ----------
    st : SpikeTrainArray
        obj that holds spiketrains
    template : EpochArray
        time in seconds, pairwise correlation calculated from this period will be compared to matching period (task-period)
    matching : EpochArray
        time in seconds, template-correlations will be correlated with pariwise correlations of this period (post-task period)
    control : EpochArray
        time in seconds, control for pairwise correlations within this period (pre-task period)
    bin_size : float
        in seconds, binning size for spike counts
    window : int
        window over which pairwise correlations will be calculated in matching and control time periods,
            if window is None entire time period is considered, in seconds
    slideby : int
        slide window by this much, in seconds
    matching_windows : array
        windows for matching period
    control_windows : array
        windows for control period
    template_corr : array
        pairwise correlations for template period
    matching_paircorr : array
        pairwise correlations for matching period
    control_paircorr : array
        pairwise correlations for control period
    ev : array
        explained variance for each time point
    rev : array
        reverse explained variance for each time point
    ev_std : array
        explained variance standard deviation for each time point
    rev_std : array
        reverse explained variance standard deviation for each time point
    partial_corr : array
        partial correlations for each time point
    rev_partial_corr : array
        reverse partial correlations for each time point
    n_pairs : int
        number of pairs
    matching_time : array
        time points for matching period
    control_time : array
        time points for control period
    ev_signal : AnalogSignalArray
        explained variance signal
    rev_signal : AnalogSignalArray
        reverse explained variance signal
    plot : function
        plot explained variance
    pvalue : function
        calculate p-value for explained variance by shuffling the template correlations

    Examples
    --------
    # Load data
    >>> basepath = r"U:\data\HMC\HMC1\day8"
    >>> st,cm = loading.load_spikes(basepath,brainRegion="CA1",putativeCellType="Pyr")

    >>> epoch_df = loading.load_epoch(basepath)
    >>> beh_epochs = nel.EpochArray(epoch_df[["startTime", "stopTime"]].values)


    # Most simple case, returns single explained variance value
    >>> expvar = explained_variance.ExplainedVariance(
    >>>        st=st,
    >>>        template=beh_epochs[1],
    >>>        matching=beh_epochs[2],
    >>>        control=beh_epochs[0],
    >>>        window=None,
    >>>    )

    # Get time resolved explained variance across entire session in 200sec bins
    >>> expvar = explained_variance.ExplainedVariance(
    >>>        st=st,
    >>>        template=beh_epochs[1],
    >>>        matching=nel.EpochArray([beh_epochs.start, beh_epochs.stop]),
    >>>        control=beh_epochs[0],
    >>>        window=200
    >>>    )

    # Get time resolved explained variance across entire session in 200sec bins sliding by 100sec
    >>> expvar = explained_variance.ExplainedVariance(
    >>>        st=st,
    >>>        template=beh_epochs[1],
    >>>        matching=nel.EpochArray([beh_epochs.start, beh_epochs.stop]),
    >>>        control=beh_epochs[0],
    >>>        window=200,
    >>>        slideby=100
    >>>    )
    """

    def __init__(
        self,
        st: SpikeTrainArray,
        template: EpochArray,
        matching: EpochArray,
        control: EpochArray,
        bin_size: float = 0.2,
        window: int = 900,
        slideby: int = None,
    ):
        """Explained variance measure for assessing reactivation of neuronal activity using pairwise correlations.

        Parameters
        ----------
        st : SpikeTrainArray
            obj that holds spiketrains
        template : EpochArray
            time in seconds, pairwise correlation calculated from this period will be compared to matching period (task-period)
        matching : EpochArray
            time in seconds, template-correlations will be correlated with pariwise correlations of this period (post-task period)
        control : EpochArray
            time in seconds, control for pairwise correlations within this period (pre-task period)
        bin_size : float, optional
            in seconds, binning size for spike counts, by default 0.2
        window : int, optional
            window over which pairwise correlations will be calculated in matching and control time periods,
                if window is None entire time period is considered, in seconds, by default 900
        slideby : int, optional
            slide window by this much, in seconds, by default None
        """
        self.__dict__.update(locals())
        del self.__dict__["self"]

        self.__validate_input()
        self.__calculate()

    def __validate_input(self):
        """Validate input parameters."""
        assert isinstance(self.st, SpikeTrainArray)
        assert isinstance(self.template, EpochArray)
        assert isinstance(self.matching, EpochArray)
        assert isinstance(self.control, EpochArray)
        assert isinstance(self.bin_size, (float, int))
        assert isinstance(self.window, (int, type(None)))
        assert isinstance(self.slideby, (int, type(None)))

    def __calculate(self):
        """processing steps for explained variance calculation."""
        control_window_size, matching_window_size, slideby = self.__get_window_sizes()

        self.matching_windows = self.__get_windows_array(
            self.matching, matching_window_size, slideby
        )
        self.control_windows = self.__get_windows_array(
            self.control, control_window_size, slideby
        )
        self.__validate_window_sizes(control_window_size, matching_window_size)
        self.template_corr = self.__get_template_corr()
        self.__calculate_pairwise_correlations()
        self.__calculate_partial_correlations()

    def __get_window_sizes(self):
        """Get window sizes for control and matching periods."""
        if self.window is None:
            control_window_size = np.array(self.control.duration).astype(int)
            matching_window_size = np.array(self.matching.duration).astype(int)
            slideby = None
        elif self.slideby is None:
            control_window_size = self.window
            matching_window_size = self.window
            slideby = None
        else:
            control_window_size = self.window
            matching_window_size = self.window
            slideby = self.slideby
        return control_window_size, matching_window_size, slideby

    def __get_windows_array(self, epoch_array, window_size, slideby):
        """Get windows array for control and matching periods."""
        if slideby is not None:
            array = np.arange(epoch_array.start, epoch_array.stop)
            windows = np.lib.stride_tricks.sliding_window_view(array, window_size)
            windows = windows[::slideby, [0, -1]]
        elif np.array(epoch_array.duration) == window_size:
            windows = np.array([[epoch_array.start, epoch_array.stop]])
        else:
            array = np.arange(epoch_array.start, epoch_array.stop, window_size)
            windows = np.array([array[:-1], array[1:]]).T
        return windows

    def __validate_window_sizes(self, control_window_size, matching_window_size):
        """Validate window sizes."""
        assert (
            control_window_size <= self.control.duration
        ), "window is bigger than matching"
        assert (
            matching_window_size <= self.matching.duration
        ), "window is bigger than matching"

    def __get_template_corr(self):
        """Get pairwise correlations for template period."""
        self.bst = self.st.bin(ds=self.bin_size)
        return self.__get_pairwise_corr(self.bst[self.template].data)

    def __calculate_pairwise_correlations(self):
        """Calculate pairwise correlations for matching and control periods."""
        self.matching_paircorr = self.__time_resolved_correlation(self.matching_windows)
        self.control_paircorr = self.__time_resolved_correlation(self.control_windows)

    @staticmethod
    def __get_pairwise_corr(bst_data):
        """Calculate pairwise correlations."""
        corr = np.corrcoef(bst_data)
        return corr[np.tril_indices(corr.shape[0], k=-1)]

    def __time_resolved_correlation(self, windows):
        """Calculate pairwise correlations for given windows."""
        paircorr = []
        bst_data = self.bst.data
        bin_centers = self.bst.bin_centers

        for w in windows:
            start, stop = w
            idx = (bin_centers > start) & (bin_centers < stop)
            corr = np.corrcoef(bst_data[:, idx])
            paircorr.append(corr[np.tril_indices(corr.shape[0], k=-1)])

        return np.array(paircorr)

    def __calculate_partial_correlations(self):
        """Calculate partial correlations."""
        partial_corr, rev_partial_corr = self.__calculate_partial_correlations_(
            self.matching_paircorr, self.control_paircorr, self.template_corr
        )
        self.__calculate_statistics(partial_corr, rev_partial_corr)

    @staticmethod
    @jit(nopython=True)
    def __calculate_partial_correlations_(
        matching_paircorr, control_paircorr, template_corr
    ):
        """Calculate partial correlations."""

        def __explained_variance(x, y, covar):
            """Calculate explained variance and reverse explained variance."""

            # Calculate covariance matrix
            n = len(covar)
            valid = np.zeros(n, dtype=np.bool_)
            for i in range(n):
                valid[i] = not (np.isnan(covar[i]) or np.isnan(x[i]) or np.isnan(y[i]))
            mat = np.empty((3, len(x)))
            mat[0] = covar
            mat[1] = x
            mat[2] = y
            cov = np.corrcoef(mat[:, valid])

            # Calculate explained variance
            EV = (cov[1, 2] - cov[0, 1] * cov[0, 2]) / (
                np.sqrt((1 - cov[0, 1] ** 2) * (1 - cov[0, 2] ** 2)) + 1e-10
            )

            # Calculate reverse explained variance
            rEV = (cov[0, 1] - cov[1, 2] * cov[0, 2]) / (
                np.sqrt((1 - cov[1, 2] ** 2) * (1 - cov[0, 2] ** 2)) + 1e-10
            )

            return EV, rEV

        n_matching = len(matching_paircorr)
        n_control = len(control_paircorr)
        partial_corr = np.zeros((n_control, n_matching))
        rev_partial_corr = np.zeros((n_control, n_matching))

        for m_i, m_pairs in enumerate(matching_paircorr):
            for c_i, c_pairs in enumerate(control_paircorr):
                partial_corr[c_i, m_i], rev_partial_corr[c_i, m_i] = (
                    __explained_variance(template_corr, m_pairs, c_pairs)
                )
        return partial_corr, rev_partial_corr

    def __calculate_statistics(self, partial_corr, rev_partial_corr):
        """Calculate explained variance statistics."""
        self.ev = np.nanmean(partial_corr**2, axis=0)
        self.rev = np.nanmean(rev_partial_corr**2, axis=0)
        self.ev_std = np.nanstd(partial_corr**2, axis=0)
        self.rev_std = np.nanstd(rev_partial_corr**2, axis=0)
        self.partial_corr = partial_corr**2
        self.rev_partial_corr = rev_partial_corr**2
        self.n_pairs = len(self.template_corr)
        self.matching_time = np.mean(self.matching_windows, axis=1)
        self.control_time = np.mean(self.control_windows, axis=1)

    @property
    def ev_signal(self):
        """Return explained variance signal."""
        return AnalogSignalArray(
            data=self.ev,
            timestamps=self.matching_time,
            fs=1 / np.diff(self.matching_time)[0],
            support=EpochArray(data=[self.matching.start, self.matching.stop]),
        )

    @property
    def rev_signal(self):
        """Return reverse explained variance signal."""
        return AnalogSignalArray(
            data=self.rev,
            timestamps=self.matching_time,
            fs=1 / np.diff(self.matching_time)[0],
            support=EpochArray(data=[self.matching.start, self.matching.stop]),
        )

    def pvalue(self, n_shuffles=1000):
        """
        Calculate p-value for explained variance by shuffling the template correlations.
        """
        from copy import deepcopy

        def shuffle_template(self):
            template_corr = deepcopy(self.template_corr)
            np.random.shuffle(template_corr)

            partial_corr, _ = self.__calculate_partial_correlations_(
                self.matching_paircorr, self.control_paircorr, template_corr
            )
            ev = np.nanmean(partial_corr**2, axis=0)
            return ev.flatten()

        if len(self.ev) > 1:
            print("Multiple time points, p-values are not supported")
            return

        ev_shuffle = [shuffle_template(self) for _ in range(n_shuffles)]

        ev_shuffle = np.array(ev_shuffle)

        n = len(ev_shuffle)
        r = np.sum(ev_shuffle > self.ev)
        pvalues = (r + 1) / (n + 1)
        return pvalues

    def plot(self):
        """Plot explained variance."""
        if self.matching_time.size == 1:
            print("Only single time point, cannot plot")
            return
        import matplotlib.pyplot as plt

        fig, ax = plt.subplots(1, 1, figsize=(8, 3))
        ax.plot(self.matching_time, self.ev, label="EV")
        ax.fill_between(
            self.matching_time,
            self.ev - self.ev_std,
            self.ev + self.ev_std,
            alpha=0.5,
        )
        ax.plot(self.matching_time, self.rev, label="rEV", color="grey")
        ax.fill_between(
            self.matching_time,
            self.rev - self.rev_std,
            self.rev + self.rev_std,
            alpha=0.5,
            color="grey",
        )
        # check if matching time overlaps with control time and plot control time
        if np.any(
            (self.control_time >= self.matching_time[0])
            & (self.control_time <= self.matching_time[-1])
        ):
            ax.axvspan(
                self.control.start,
                self.control.stop,
                color="green",
                alpha=0.3,
                label="Control",
                zorder=-10,
            )
        # check if matching time overlaps with template time and plot template time
        if np.any(
            (self.template.start >= self.matching_time[0])
            & (self.template.stop <= self.matching_time[-1])
        ):
            ax.axvspan(
                self.template.start,
                self.template.stop,
                color="purple",
                alpha=0.4,
                label="Template",
                zorder=-10,
            )
        # remove axis spines
        ax.spines["right"].set_visible(False)
        ax.spines["top"].set_visible(False)

        ax.legend(frameon=False)
        ax.set_xlabel("Time (s)")
        ax.set_ylabel("Explained Variance")
        ax.set_title("Explained Variance")
        plt.show()

ev_signal property

Return explained variance signal.

rev_signal property

Return reverse explained variance signal.

plot()

Plot explained variance.

Source code in neuro_py/ensemble/explained_variance.py
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def plot(self):
    """Plot explained variance."""
    if self.matching_time.size == 1:
        print("Only single time point, cannot plot")
        return
    import matplotlib.pyplot as plt

    fig, ax = plt.subplots(1, 1, figsize=(8, 3))
    ax.plot(self.matching_time, self.ev, label="EV")
    ax.fill_between(
        self.matching_time,
        self.ev - self.ev_std,
        self.ev + self.ev_std,
        alpha=0.5,
    )
    ax.plot(self.matching_time, self.rev, label="rEV", color="grey")
    ax.fill_between(
        self.matching_time,
        self.rev - self.rev_std,
        self.rev + self.rev_std,
        alpha=0.5,
        color="grey",
    )
    # check if matching time overlaps with control time and plot control time
    if np.any(
        (self.control_time >= self.matching_time[0])
        & (self.control_time <= self.matching_time[-1])
    ):
        ax.axvspan(
            self.control.start,
            self.control.stop,
            color="green",
            alpha=0.3,
            label="Control",
            zorder=-10,
        )
    # check if matching time overlaps with template time and plot template time
    if np.any(
        (self.template.start >= self.matching_time[0])
        & (self.template.stop <= self.matching_time[-1])
    ):
        ax.axvspan(
            self.template.start,
            self.template.stop,
            color="purple",
            alpha=0.4,
            label="Template",
            zorder=-10,
        )
    # remove axis spines
    ax.spines["right"].set_visible(False)
    ax.spines["top"].set_visible(False)

    ax.legend(frameon=False)
    ax.set_xlabel("Time (s)")
    ax.set_ylabel("Explained Variance")
    ax.set_title("Explained Variance")
    plt.show()

pvalue(n_shuffles=1000)

Calculate p-value for explained variance by shuffling the template correlations.

Source code in neuro_py/ensemble/explained_variance.py
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def pvalue(self, n_shuffles=1000):
    """
    Calculate p-value for explained variance by shuffling the template correlations.
    """
    from copy import deepcopy

    def shuffle_template(self):
        template_corr = deepcopy(self.template_corr)
        np.random.shuffle(template_corr)

        partial_corr, _ = self.__calculate_partial_correlations_(
            self.matching_paircorr, self.control_paircorr, template_corr
        )
        ev = np.nanmean(partial_corr**2, axis=0)
        return ev.flatten()

    if len(self.ev) > 1:
        print("Multiple time points, p-values are not supported")
        return

    ev_shuffle = [shuffle_template(self) for _ in range(n_shuffles)]

    ev_shuffle = np.array(ev_shuffle)

    n = len(ev_shuffle)
    r = np.sum(ev_shuffle > self.ev)
    pvalues = (r + 1) / (n + 1)
    return pvalues