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415 | 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()
|