Claims 46
From the paper mPFC spindle cycles organize sparse thalamic activation and recently active CA1 cells during non-REM sleep · claims file · checked 2026-11-06: ready
- fig1s3-spindles-not-theta
Spindle cycles detected in sleep carry spectral power concentrated near 10 to 15 Hz in the mPFC and CA1 LFP, unlike cycles detected by the same algorithm in quiet or active waking, which have theta-like profiles.
Spindle cycles detected during sleep are associated with spectral power concentrated in the 10–15 Hz band, suggestive of spindle (and not theta) detection
Figure 1, figure supplement 3 legendHow it will be judged
Spindle troughs by M-spindle, in sleep (M-sleep) and, separately, in waking time outside sleep. A wavelet scalogram of the mPFC and CA1 LFP averaged around troughs (plus or minus 0.5 s; Morlet wavelet, a standard choice, as the paper names no wavelet). Reproduced when the sleep trough-triggered mPFC scalogram peaks between 10 and 15 Hz and the waking CA1 scalogram peaks at a lower frequency than the sleep one.
- fig2-swr-phase-delta-spindle
CA1 SWRs are phase-locked to the mPFC delta peak (circular mean -0.04 rad) and to the trough and ascending phase of mPFC spindles (circular mean -1.72 rad).
SWRs preferentially occurred following and preceding KCs, phase-locked to the peaks of the delta-filtered LFP, and to the troughs and ascending phase of spindle cycles (circular mean and standard deviation of delta phase at the time of SWRs = −0.04 ± 0.43 rad; spindle phase = −1.72 ± 0.93 rad; n = 23 sessions, six rats
Results, opening section, paragraph 4; Figure 2B; Figure 2, figure supplement 1How it will be judged
SWRs by M-SWR in M-sleep; phases by M-phase. Per session the circular mean of the normalized phase distribution at SWR times; across sessions the circular mean and circular sd. Tolerances are the paper's circular sd. The paper reports no test for this finding. Reproduced when both means fall within tolerance; the per-session Rayleigh test (M-phase) is reported.
- fig2c-swr-around-kc
SWR probability rises within about 250 ms before and after mPFC K-complexes (4.65 sd at 208 ms before, 3.02 sd at 267 ms after), with its minimum about 10 ms after the KC.
SWR probability increased within the 250 ms around mPFC KCs (4.65 sd at 208.15 ± 90 ms before KCs, and 3.02 sd at 266.85 ± 100 ms after KCs; n = 23 sessions). This indicates a delta modulated increase in SWR probability in CA1, with the minimum rate of ripples following briefly after the trough of neocortical activity (10.33 ± 5.88 ms)
Results, Thalamic rebound following the down-state, paragraph 2; Figure 2CHow it will be judged
KCs by M-KC, SWRs by M-SWR, peri-KC SWR histogram by M-PETH, per session, then averaged over sessions. Peaks are the maximum z in -0.5 to 0 s and 0 to 0.5 s (the 0.5 s windows of the asymmetry index, Results, same section, paragraph 3). The window for the minimum is not given; readings (a) -0.25 to 0.25 s (the "250 ms around" of the Results), (b) -0.5 to 0.5 s. Open, whether the peak heights are of the session-averaged histogram or averages of per-session peaks (the times carry an sd, so they are per-session values); readings (i) mean of per-session peaks, (ii) peak of the mean histogram. Reproduced when, under a reading, both peaks are positive and all five values are within tolerance.
- fig3a-thalamic-rate-wake-sleep
Thalamic units fire less in sleep than in waking (10.10 against 7.73 spikes/s).
the firing rate of thalamic units changed from an average of 10.10 ± 4.26 spikes/s during wakefulness to 7.73 ± 3.09 spikes/s during sleep (n = 40 units; p=1.31 × 10 −6 , Wilcoxon signed rank
Results, Thalamic cells decrease their overall firing, paragraph 2; Figure 3AHow it will be judged
Sleep is M-sleep. Sessions with less than 5 min of waking are left out (Results, same paragraph). Waking is not defined; readings (a) all time outside the sleep periods, (b) time outside sleep when the smoothed velocity (M-sleep) exceeds 2 cm/s, a threshold the paper does not give. Rate is spike count over time in the state. Reproduced when, under a reading, sleep rate is lower in a Wilcoxon signed rank test with p < 0.05 and both means are within tolerance.
- fig3a-bursting-in-sleep
Thalamic units burst more in sleep; 25% of units have at least 5% of spikes in bursts in waking against 77.5% in sleep, and burst spikes rise from 4.60% to 18.65%.
we found that 25% of the thalamic cells had at least 5% of their spikes in bursts during wakefulness, while 77.5% did during sleep (average percentage of spikes in bursts during wakefulness 4.60 ± 7.97%, and during sleep 18.65 ± 15.3%; p=1.8 × 10 −7 ; Wilcoxon signed rank; n = 40 units, seven rats
Results, Thalamic cells decrease their overall firing, paragraph 4; Figure 3AHow it will be judged
Bursts by M-burst, states as in fig3a-thalamic-rate-wake-sleep, with its readings. Reproduced when, under a reading, the sleep percentage is higher (Wilcoxon signed rank, p < 0.05) and the four values are within tolerance. With fewer units than the paper, the two proportions are judged on direction and tolerance together.
- fig3a-bursting-around-swrs
Thalamic bursting is higher still in the 2 s around CA1 SWRs (25.52% of spikes in bursts).
The increase in bursting was more pronounced around the time of SWRs in CA1 (25.52 ± 17.29% of spikes in bursts in a 2 s window around SWRs; p=1.8 × 10 −7 Kruskall-Wallis
Results, Thalamic cells decrease their overall firing, paragraph 4; Figure 3AHow it will be judged
Spikes within plus or minus 1 s of SWRs (M-SWR) in sleep. Open, how overlapping windows count; readings (a) the union of windows, each spike once, (b) per SWR, a spike counted in every window it falls in. A Kruskal-Wallis test across the three groups (waking, sleep, SWR windows) as the paper names. Reproduced when the test gives p < 0.05, the SWR-window mean exceeds the sleep mean and is within tolerance.
- swr-shift-control
Compared with SWR times shifted by more than 2 s or 5 s, the 2 s windows around SWRs hold fewer thalamic spikes and a higher percentage of burst spikes.
In both cases (>2 and>5 s shifts), the average number of thalamic unit spikes was significantly lower in the SWRs windows compared to the shifted timestamps (p<10 −3 ), and the percent of spikes in bursts significantly higher (p=0.002 and p<10 −3 respectively after a 2 s or 5 s shift; Wilcoxon signed rank
Results, Thalamic cells decrease their overall firing, paragraph 4How it will be judged
Each SWR time shifted at random; the paper says only "at least 2 (or 5 s)". Readings for the shift (a) uniform in 2 to 3 s and in 5 to 6 s, (b) uniform in 2 to 10 s and in 5 to 10 s, forward. Per unit, spikes and burst-spike percent in plus or minus 1 s windows at true and shifted times, compared by Wilcoxon signed rank. Reproduced when, under a reading, both comparisons go the paper's way with p < 0.05 for both shifts.
- fig3c-thalamic-decrease-at-swrs
About 55% of thalamic units reduce their firing by more than 2 sd from the null in the 200 ms around SWRs (average -4.23 sd); 14.89% show a pure increase and 8.51% a rebound after a decrease.
55.32% of units in the thalamus showed a reduction in firing of more than two standard deviations (sd) from the null (average −4.23 ± 2.00 sd) in a 200 ms window around the time of SWRs occurrence ... A few thalamic units (14.89%) showed a pure increase in firing in the 200 ms window around CA1 SWRs ... Another four units (8.51%) had a rebound in firing
Results, Thalamic cells decrease their overall firing, paragraph 5; Figure 3B-CHow it will be judged
Peri-SWR histograms by M-PETH for each thalamic unit in sleep. The percentages fit 47 units (26, 7 and 4 of 47; Figure 4, figure supplement 1C gives n = 47). Open, how the window is read; readings (a) the minimum (or maximum) z in -100 to 100 ms beyond 2 sd, (b) the mean z over that window beyond 2 sd. A pure increase has a maximum above 2 sd and no bin below -2 sd before it; a rebound has a bin below -2 sd followed by a bin above 2 sd within the window. Reproduced when, under a reading, the decreasing units are the largest group and the four values are within tolerance; partially when only the direction holds.
- fig3s1-ca1-mpfc-at-swrs
Around SWRs, 90.18% of CA1 units increase their firing (by 19.85 sd on average), while in mPFC 30.95% of units increase and 40.48% decrease.
Instead, 90.18% of CA1 units increased their firing by 19.85 ± 18.65 sd in a 200 ms window around SWRs (4.91% of CA1 units showed an average decrease of 4.11 ± 1.59 sd), while the mPFC population included 30.95% of units showing an increase (average 6.22 ± 4.32 sd) and 40.48% a reduction in firing (average −3.22 ± 1.40 sd
Results, Thalamic cells decrease their overall firing, paragraph 5; Figure 3, figure supplement 1AHow it will be judged
As in fig3c-thalamic-decrease-at-swrs, with its readings, for CA1 and mPFC units. Reproduced when, under a reading, a large majority of CA1 units increase, mPFC holds both increasing and decreasing units, and the four values are within tolerance.
- glm-re-versus-other-nuclei
The size of the GLM effect is marginally larger in RE than in other thalamic nuclei (p=0.05), while the proportion of SWR-correlated units does not differ (p=0.159).
the size of the effect was marginally significant in RE compared to non-RE units (p=0.05). However, there was no difference between the proportion of SWR-correlated units in RE compared to non-RE units (p=0.159
Results, Thalamic cells decrease their overall firing, paragraph 6How it will be judged
Needs the nucleus of each thalamic unit (Table 1; from histology). The tests are not named. For the effect size, Mann-Whitney U on b1 (M-stats, unpaired). For the proportion, readings (a) Fisher exact test, (b) chi-square test, as M-stats names neither. The absence part is judged on that test at p < 0.05; reproduced when the proportion test does not reach p < 0.05 and the RE effect is the larger, with its p reported.
- fig3e-thalamic-rebound-after-kc
Thalamic units fire more after mPFC K-complexes than before (2.48 sd at about 275 ms after against 1.28 sd at about 263 ms before).
Units in the thalamus often fired asymmetrically with respect to KCs, with stronger firing in the few hundreds of milliseconds after the mPFC KC compared to before KCs (average of 2.48 standard deviations at 274.76 ± 135.33 ms after KCs, compared to 1.28 standard deviations at 263.31 ± 145.31 ms before KCs; n = 31 thalamic units
Results, Thalamic rebound following the down-state, paragraph 1; Figure 3D-EHow it will be judged
Peri-KC histograms by M-PETH (KCs by M-KC) per thalamic unit. Peaks are the maximum z in 0 to 0.5 s after and -0.5 to 0 s before KCs (the windows of the asymmetry index). The paper reports no test here; reproduced when the after peak exceeds the before peak on average and the four values are within tolerance.
- kc-trough-timing
The trough of thalamic firing follows the KC (10.56 ms after), after the trough of mPFC units (-7.70 ms) and with the SWR trough (10.33 ms after).
(thalamic unit firing minimum at 10.56 ± 130.75 ms after KCs, n = 31; mPFC units −7.70 ± 72.3 ms, before LFP KCs, n = 46; SWRs 10.33 ± 5.88 ms after KCs
Results, Thalamic rebound following the down-state, paragraph 1How it will be judged
Time of the minimum z of each unit's peri-KC histogram (M-PETH), averaged over units. The paper gives no window for the minimum; readings (a) -0.25 to 0.25 s, (b) -0.5 to 0.5 s (the asymmetry index windows), neither stated for this measure. The wide tolerances are the paper's sd; reproduced when both means are within them and the thalamic mean is later than the mPFC mean.
- kc-fraction-of-thalamic-peaks
38.71% of thalamic units have a significant peak (above 2 sd) after KCs against 9.68% with a significant peak before KCs; 19.35% have both.
For the population of thalamic cells, 38.71% increased their activity above two sd (from the null distribution) following KCs, that is, in the down to up-state transition, compared to 9.68% of the cells with a significant peak preceding KCs, that is, in the up to down-state transition (19.35 % units had a significant peak both before and after KCs)
Results, Thalamic rebound following the down-state, paragraph 2How it will be judged
Peaks as in fig3e-thalamic-rebound-after-kc, over 31 units in the paper (12, 3 and 6 of 31). Open, whether 38.71% and 9.68% exclude the units with both; readings (a) they exclude them (the three groups are disjoint), (b) they include them. Reproduced when, under a reading, more units peak after than before and the values are within tolerance.
- kc-ca1-mpfc-symmetric
CA1 and mPFC units increase firing both before and after KCs, by similar amounts.
The modulation of SWRs was paralleled by a modulation of firing in CA1 units (n = 111), which increased their activity both before and after KCs (2.32 ± 1.49 sd at −233.7 ± 146 ms before KCs, and 2.35 ± 1.69 sd at 273 ± 140 ms after KCs) ... Activity in mPFC (n = 46 units) was modulated in a similar way, with increases in unit firing preceding and following the down-state of the slow oscillation (2.33 standard deviations at −267.30 ± 110.40 ms, before KCs; 2.31 standard deviations at 295.20 ± 120 ms, after KCs
Results, Thalamic rebound following the down-state, paragraph 2How it will be judged
As in fig3e-thalamic-rebound-after-kc for CA1 and mPFC units. Reproduced when the four values are within tolerance; the peak times are reported.
- kc-regions-compared
Peaks after KCs do not differ between thalamus, mPFC and CA1, while thalamic units fire significantly less before KCs than mPFC and CA1 units.
while units from the three areas showed no difference in the peak of the histograms in the time window following KCs (p=0.89, Kruskal-Wallis between regions), thalamic units had significantly lower firing before KCs compared to mPFC and CA1 units (p=0.007, Kruskal-Wallis
Results, Thalamic rebound following the down-state, paragraph 3How it will be judged
Peak z values as in fig3e-thalamic-rebound-after-kc, Kruskal-Wallis across the three regions. The absence after KCs is judged on that test at p < 0.05; the before effect is reproduced at p < 0.05 with the thalamic median lowest.
- fig3f-asymmetry-index
The KC asymmetry index is biased to positive values for thalamic units (0.24) and centred near zero for mPFC (-0.004) and CA1 (-0.06) units.
The distribution of this index ( Figure 3f ) was biased to positive values for thalamic units, but was uniformly distributed for mPFC and CA1 units (average 0.24 ± 0.22 in thalamus; −0.004 ± 0.47 in mPFC; −0.06 ± 0.5 in CA1
Results, Thalamic rebound following the down-state, paragraph 3; Figure 3FHow it will be judged
Index from the maximum firing rate in 0.5 s windows before and after KCs, after minus before, normalized to the maximum of the two windows (Results, same paragraph). Open, the histogram used and the normalization; readings (a) raw-rate peri-KC histogram (M-PETH bins), divided by the larger maximum, (b) the same divided by the sum of the two maxima, (c) z-scored histogram as in (a). Tolerances are the paper's sd. Reproduced when, under a reading, the thalamic mean is positive and within tolerance and the mPFC and CA1 means are within tolerance; a Wilcoxon signed rank test of the thalamic index against 0 is reported.
- spindle-mua-modulation
Multi-unit activity increases around mPFC spindle troughs in 84.62% of CA1 tetrodes, 53.3% of mPFC and 36% of thalamic tetrodes; individual units are less clearly modulated.
Across the sessions, 84.62% of the CA1 tetrodes showed an increase in spike activity (at least three standard deviations from null distribution in the −100 to 100 ms window of the peri-spindle-trough histogram), with 53.3% and 36% tetrodes showing a similar result in mPFC and thalamus
Results, Sparse thalamic firing, paragraph 1; Figure 3G-H; Figure 3, figure supplement 1CHow it will be judged
Peri-spindle-trough histograms of each tetrode's MUA by M-PETH (troughs by M-spindle); a tetrode counts when its maximum z in -100 to 100 ms is at least 3. Reproduced when the order CA1 > mPFC > thalamus holds and the values are within tolerance.
- fig4-units-not-rhythmic
Thalamic units do not fire rhythmically at spindle frequency; the autocorrelation peak is later for units (223.60 ms) than for thalamic MUA (119.30 ms), and falls in the spindle band for 63.3% of MUA curves against 17.02% of units.
We found that the peak of the MUA autocorrelation occurs on average at 119.30 ± 78 ms, whereas the interval is longer for the units, 223.60 ± 113.30 ms (p<0.001, Wilcoxon Mann-Whitney). The peak of the autocorrelation occurred within the spindle band in 63.3% of the curves calculated with MUA (with subsampled spikes matched to the units) and in 17.02% of the units
Results, Sparse thalamic firing, paragraph 2; Figure 4A; Figure 4, figure supplement 1How it will be judged
Autocorrelation of sleep spikes; the peak is the maximum after the minimum that follows lag 0 (Results, same paragraph). MUA subsampled in blocks with the spike count of the sparsest unit of the session, averaged (Figure 4, figure supplement 1B). The 63.3% is of subsampled curves; for the 119.30 ms mean the paper plots MUA "all the spikes in sleep or a subsample" and does not say which it averages; readings (A) all MUA spikes, (B) the subsampled curves. Open, bin size, lag range and normalization; readings (a) 5 ms bins, (b) 10 ms bins, lags to 500 ms, normalized so lag 0 is 1 after removing it (coefficient), as the values 0.24 and 0.05 imply a normalized measure. Open, the spindle band in lags; readings (i) 71 to 167 ms (6 to 14 Hz), (ii) 50 to 150 ms (Results, same paragraph). Reproduced when, under a reading, unit peaks are later than MUA peaks (Mann-Whitney, p < 0.05), the spindle-band fraction is higher for MUA, and the lag means are within tolerance.
- xcorr-stronger-in-spindles
Pair cross-correlograms are stronger with spikes within spindle cycles (2.74 sd) than with sleep spikes between spindles (2.31 sd).
(average peak cross-correlograms with spindle spikes 2.74 ± 0.85 sd compared to 2.31 ± 1.18 sd with sleep spikes that occurred between detected spindles; p=0.015, Wilcoxon signed rank test
Results, Sparse thalamic firing, paragraph 5; Figure 4D; Figure 4, figure supplement 2B-CHow it will be judged
Correlograms as in fig4b-thalamic-pairs-correlated, with its readings, on spikes inside spindle cycles (M-spindle) and, separately, sleep spikes in periods with no spindle for at least 5 s. Open, the between-spindle periods; readings (a) gaps between consecutive troughs longer than 5 s, all their spikes, (b) only spikes more than 2.5 s from any trough. Reproduced when the spindle peak is higher (Wilcoxon signed rank, p < 0.05) and both values are within tolerance.
- fig5c-spindle-phase-locking
Most thalamic units (67.74%) fire preferentially on the descending phase of the mPFC spindle oscillation (circular mean 0.77 rad).
We found that the majority of thalamic units (67.74% of n = 31 units) showed preferential firing at the descending phases of the spindle-filtered mPFC LFP, as suggested by the positive values of the mean phase distribution (circular mean 0.77 ± 1.15 rad; p<0.05, Rayleigh test
Results, Phase-locking of thalamic units, paragraph 2; Figure 5CHow it will be judged
Spike phases by M-phase in sleep. The p < 0.05 is the per-unit threshold of the count (a selection, not recorded as a statistic). Open, what 67.74% counts; readings (a) units with a significant Rayleigh test, (b) units with positive mean phase, (c) both. Reproduced when, under a reading, the fraction and the circular mean are within tolerance.
- fig5c-delta-phase-locking
Thalamic units are phase-locked to mPFC delta near its positive peak (64.52% of units; circular mean 0.30 rad).
Thalamic units were phase-locked to the mPFC delta band, closer to the positive peak of the oscillation (64.52%, p<0.05, Rayleigh test; circular mean 0.30 ± 1.04 rad
Results, Phase-locking of thalamic units, paragraph 2; Figure 5CHow it will be judged
As in fig5c-spindle-phase-locking for the delta band, the fraction being units with a significant Rayleigh test. Reproduced when both values are within tolerance.
- phase-robust-to-spike-count
Preferred phases do not change when computed with 10% of the spikes.
the circular mean was not different when the histograms were calculated with 10% of the spikes (p=0.68 for spindle and 0.07 for delta; Wilcoxon signed rank
Results, Phase-locking of thalamic units, paragraph 2How it will be judged
A claim of absence. Per unit the circular mean from all spikes and from a random 10% of them (seed fixed, a standard choice), compared by Wilcoxon signed rank over units on the phase values as the paper names. Reproduced when p >= 0.05 for both bands; the median difference is reported.
- rebound-delta-phase-trend
The amplitude of the post-KC thalamic rebound tends to correlate with earlier preferred delta phase, though not significantly (r = -0.32, p=0.08); no such trend holds for firing before KCs.
The amplitude of the thalamic rebound after KCs (reported above) had a tendency to correlate with a preference for earlier phases of delta; although the correlation was not significant (p=0.08, r = −0.32), we did not observe such a trend when we regressed mean phase preference to the cell firing rate in a time window of the same length but before KCs (p=0.65, r = 0.08
Results, Phase-locking of thalamic units, paragraph 2How it will be judged
Rebound amplitude is the peak z after KCs (fig3e-thalamic-rebound-after-kc); the control is "the cell firing rate in a time window of the same length but before KCs". Open, the measure for both windows; readings (i) peak z in 0 to 0.5 s and in -0.5 to 0 s, (ii) mean firing rate in the same windows. The test is a linear regression ("regressed"), Pearson r. Open, whether the phase is circular; readings (a) phase in radians as a linear variable, (b) circular-linear correlation. Reproduced when, under a reading, r is negative with p >= 0.05 for the rebound and the before-KC regression gives p >= 0.05; the paper's absence of significance is judged at 0.05.
- fig5d-phase-lags-consistent
Phase lags between spikes of correlated thalamic pairs within spindles are less variable than those of uncorrelated pairs (sd 2.15 against 2.50).
The standard deviation of phase lag distributions within spindles was significantly lower for correlated pairs compared to uncorrelated pairs (2.15 compared to 2.50; p<0.001, Mann-Whitney U-test
Results, Phase-locking of thalamic units, paragraph 3; Figure 5D; Figure 5, figure supplement 1How it will be judged
For each spike of one unit in a spindle cycle, the spindle phase difference to the closest spike of the other, unwrapped to -2 pi to 2 pi (Results, same paragraph). Pairs correlated when the cross-correlogram peak exceeds 3 sd (fig4b-thalamic-pairs-correlated). Open, whether the sd is linear or circular; the -2 pi to 2 pi range implies linear. Open, whether the Mann-Whitney test is on pairs (one sd each) or on pooled lags; readings (a) pairs, (b) lags pooled per group. Reproduced when, under a reading, correlated pairs have the lower sd with p < 0.05 and both values are within tolerance.
- spindle-cycles-by-ripple-power
Spindle cycles with ripple power above the median are slightly shorter and larger, significant within most sessions but not across sessions.
Cycles with ripple power above the median were slightly shorter in duration (102.17 ± 2.01 ms compared to 103.22 ± 2.35 ms when ripple power was low; p<0.05 in 65% of the sessions, although not significant for the mean duration across sessions, p=0.11; Mann-Whitney U-test) and larger in amplitude (229.21 ± 102.14 µV versus 219.47 ± 94.32 µV; p<0.05 in 85% of sessions, p=0.54 across sessions; Mann-Whitney U-test
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 1; Figure 6, figure supplement 1How it will be judged
Cycles by M-spindle; ripple power is the mean squared ripple-filtered CA1 LFP over the cycle (Figure 6, figure supplement 1 title), split at the session median. Open, the amplitude; readings (a) mean of the two peaks minus the trough, (b) the larger peak minus the trough, on the spindle-filtered trace. Per session a Mann-Whitney U test; across sessions Mann-Whitney on session means. Reproduced when, under a reading, high cycles are shorter and larger within most sessions, the across-session tests give p >= 0.05, and the four values are within tolerance.
- fig6-modulation-with-ripple-power
The peri-spindle modulation index of most thalamic units (64.52%) is higher in spindle cycles with ripple power above the median; 22.58% are more modulated in low ripple power cycles.
This index increased in the majority of thalamic units in spindle cycles with ripple power above the median (64.52%, n = 31) ... In addition, a subset of units (22.58%, n = 31) showed stronger modulation with low ripple power spindles, with modulation values that were at least as high as the average for cells that increased their modulation with high ripple power
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 2; Figure 6A, DHow it will be judged
Modulation index is peak minus trough over the mean of the peri-spindle-trough histogram (M-PETH bins, raw rate) in a 200 ms window centred at 0 (Results, same paragraph), for high and low ripple power cycles. The 22.58% are units with a higher index in low cycles and a low-cycle index at least the mean high-cycle index of the increasing units. Reproduced when both values are within tolerance; partially when only a majority increases.
- modulation-amplitude-versus-ripple
The modulation index increases significantly from low to high amplitude spindles (p=0.005) but not from low to high ripple power spindles (p=0.25); absolute index values are similar under both classifications.
The absolute index values were similar in spindles classified based on spindle amplitude or ripple power (p>0.3, Wilcoxon signed rank), but increased significantly from low to high amplitude spindles (p=0.005) compared to low to high ripple power spindles (p=0.25). The modulation also increased, although less, when comparing spindles with low amplitude and low ripple power to spindles with high amplitude and high ripple power (p=0.014
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 4; Figure 6DHow it will be judged
Indices as in fig6-modulation-with-ripple-power, with cycles split at the session median of amplitude (readings of spindle-cycles-by-ripple-power), of ripple power, and of both jointly. Wilcoxon signed rank over units (Results names it for the first test; M-stats, paired, for the rest). The similarity of absolute values is a claim of absence, judged on the Wilcoxon signed rank test over units of the index under the amplitude split against the index under the ripple split at p < 0.05. Open, which cycles it compares; readings (a) high cycles of each split, (b) low and high cycles, each tested. Reproduced when that test gives p >= 0.05, the amplitude split gives p < 0.05 and the ripple split p >= 0.05, with the joint split p < 0.05.
- fig6e-quartile-synergy
In the highest amplitude spindles ripple power has no significant effect on modulation (p=0.94), while in the highest ripple power spindles amplitude does (p=0.04); 61.29% of units still increase modulation with ripple power in high amplitude spindles.
In the population of cells, we did not find evidence of a significant effect of ripple power on the modulation value of thalamic firing in high amplitude spindles (p=0.94; Wilcoxon signed rank), while higher spindle amplitude led to higher modulation values on high ripple power spindles (p=0.04). However, although the modulation values were not significantly different, a majority (61.29%) of units showed a further increase in modulation index with ripple power in high amplitude spindles (average increase 18.15 ± 21.94%); instead, 32.26% of the units increased their modulation with spindle amplitude (in the spindles with highest ripple power; average increase 30.00 ± 20.97%) ... The percent change in the modulation index with ripple power and spindle amplitude were positively correlated (Spearman correlation 0.51; p=0.003
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 4; Figure 6E; Figure 6, figure supplement 2How it will be judged
Cycles in the upper quartile of amplitude, split by ripple power; cycles in the upper quartile of ripple power, split by amplitude (Results, same paragraph). Open, the median for the split; readings (a) median within the quartile subset, (b) session median over all cycles. Index as in fig6-modulation-with-ripple-power. Reproduced when, under a reading, the ripple effect is absent (p >= 0.05), the amplitude effect present (p < 0.05), the Spearman correlation positive with p < 0.05, and the four values within tolerance.
- fig6f-quartile-versus-median
Modulation values in the upper quartile are higher for spindle amplitude (1.16) than for ripple power (0.60), and the quartile split widens the spread of the index for amplitude but not for ripple power.
The increase in modulation index was higher in the upper quartile of spindle amplitude in 29% of thalamic units and with ripple power in 22.6% units. The absolute modulation values were higher for spindle amplitude compared to ripple power and positively correlated (average of 1.16 ± 0.57 in the upper quartile of spindle amplitude compared to 0.60 ± 0.26 in the upper quartile of ripple power; p=0.04, Mann-Whitney ... leading to a wider spread of the distribution (when spindles were classified based on quartiles compared to median; p=0.04; F-test) which was not the case with ripple power (p=0.36; F-test
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 5; Figure 6FHow it will be judged
Index as in fig6-modulation-with-ripple-power, in the upper quartile of amplitude and of ripple power against the remaining cycles, and against the median split. Open, what the quartile comparison is against; readings (a) upper quartile against the lower three quartiles, (b) upper against lower quartile. Mann-Whitney and F-tests as the paper names. Reproduced when, under a reading, the amplitude quartile index is the higher with p < 0.05, the F-test is significant for amplitude only, and the values are within tolerance.
- fig6g-spike-count-unchanged
The number of spikes of thalamic units does not change significantly between low and high ripple power spindles, nor between SWRs coupled and uncoupled to spindles.
the absolute number of spikes in single units did not change significantly with spindles low or high in ripple power, nor with SWRs coupled or uncoupled to spindles
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 6; Figure 6GHow it will be judged
A claim of absence, for which the paper names no test. Following M-stats (paired data), a Wilcoxon signed rank test over units at p < 0.05 of mean spikes per cycle in low against high cycles, and of mean rate in plus or minus 50 ms of SWRs in spindle cycles against an equal random sample of SWRs outside them. Reproduced when neither test is significant; median differences are reported.
- xcorr-pairs-ripple-modulation
In spindle cycles with high ripple power, the modulation index of pair cross-correlograms rises by 26.64% in 48.72% of thalamic pairs and falls by 20.90% in the rest.
We calculated the modulation index for the cross-correlograms of thalamic pairs and found that it increased by an average of 26.64% in 48.72% of the pairs in spindle cycles with high ripple power and decreased by 20.90% in the rest of the pairs (n = 39
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 7How it will be judged
Cross-correlograms of each pair with spikes in high and in low ripple power cycles (correlogram readings of fig4b-thalamic-pairs-correlated), modulation index as in fig6-modulation-with-ripple-power over plus or minus 100 ms of lag. Reproduced when the three values are within tolerance.
- thalamus-at-spindle-nested-swrs
Thalamic firing near SWRs is low and does not differ between spindle-nested and other SWRs in a 100 ms window (3.83 against 3.99 spikes/s), with a slight decrease for nested SWRs in windows of 1 s or more.
We found low firing rates but no significant difference in thalamic firing near SWRs that occurred within detected spindle cycles (mean firing 3.83 ± 3.49 spikes/s with spindle-nested SWRs, compared to 3.99 ± 3.39; p=0.18, Wilcoxon signed rank; n = 31); only when broadening the window of the cross-correlograms to at least 1 s around SWRs, did we find a slight decrease in thalamic firing with SWRs nested in spindles (mean firing 3.73 ± 3.43 spikes/s with spindle-nested SWRs, compared to 3.94 ± 3.35; p<10 −2 , Wilcoxon signed rank; n = 31
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 8How it will be judged
Nested SWRs fall inside a spindle cycle (M-spindle); the comparison set is an equal random sample of SWRs outside cycles (seed fixed, a standard choice). Mean rate in the central 100 ms and in the central 1 s of each unit's SWR cross-correlogram. Wilcoxon signed rank over units. The 100 ms absence is judged at p < 0.05; the 1 s decrease is reproduced at p < 0.05 with nested lower. Values within tolerance.
- mpfc-units-coupled-swrs
mPFC units as a population do not fire differently at SWRs coupled or uncoupled to spindles (z 0.17 against 0.24, p=0.46).
we found that there was no significant difference for the population of mPFC units between SWRs coupled or uncoupled to spindles (z-score 0.17 ± 1.26 for coupled-SWRs and 0.24 ± 0.86; p=0.46, Wilcoxon signed rank
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 9; Figure 6, figure supplement 3AHow it will be judged
Peri-SWR histograms of mPFC units in 2 ms bins over plus or minus 400 ms, coupled SWRs against an equal sample of uncoupled ones, z-scored to one shuffled distribution from randomized SWR times (Results, same paragraph); the value is the mean z in -40 to 40 ms (Figure 6, figure supplement 3A legend). Open, the randomization; readings (a) the M-PETH shifts, (b) uniform random times within sleep. A claim of absence, judged on Wilcoxon signed rank at p < 0.05.
- mua-coupled-swrs
mPFC and CA1 multi-unit activity is higher at SWRs coupled to spindles than at uncoupled SWRs, with no difference in thalamic tetrodes.
With this analysis we found stronger correlation of mPFC spikes with SWRs coupled to spindles (z-score 0.83 ± 1.34 compared to uncoupled, 0.09 ± 1.53; p=0.002). We also found stronger correlations with CA1 firing (55.81 ± 28.40, 50.40 ± 30.51; p<10–3; Wilcoxon signed rank), and no significant difference in the thalamic tetrodes (−64 ± 1.42, 0.34 ± 0.95; p=0.17
Results, Spindle activation of thalamic units modulated with CA1 SWRs, paragraph 9; Figure 6, figure supplement 3BHow it will be judged
As in mpfc-units-coupled-swrs, with its readings, per tetrode MUA. The thalamic coupled value is printed "−64 ± 1.42", which cannot be a z-score mean with that sd; it is read as -0.64 and not given an expected value. Wilcoxon signed rank over tetrodes (named for CA1, M-stats for the rest). Reproduced when mPFC and CA1 are higher for coupled SWRs at p < 0.05 and the thalamic test gives p >= 0.05.
- maze-swr-kc-increase
After maze exploration, SWR amplitude and rate and KC amplitude increase in sleep, while spindle number and amplitude do not change.
All the sessions showed an increase in the amplitude (peak-to-trough) and rate (per second) of SWRs during sleep after the maze exploration, as well as an increase in the amplitude of KCs (p=0.03, Wilcoxon signed rank), while spindles did not change significantly in number or amplitude (p>0.5
Results, Multi-region correlations after spatial exploration, paragraph 1How it will be judged
Needs the six maze sessions. Events by M-KC, M-spindle, M-SWR in sleep before and after the maze; per session the mean peak-to-trough amplitude of the ripple-filtered SWR, SWR count per second of sleep, mean KC amplitude, spindle trough count per second and amplitude. Wilcoxon signed rank over sessions (with n = 6, p = 0.03 means all six increase). Reproduced when every session increases in SWR amplitude, SWR rate and KC amplitude and the spindle tests give p >= 0.05.
- maze-control-sessions
In long sessions without the maze, SWR amplitude and rate do not differ between the first and second half.
We compared the SWRs amplitude and rate in the first half to the second half of these sessions, finding no difference in amplitude nor rate between the first and second half of the recording (p>0.2
Results, Multi-region correlations after spatial exploration, paragraph 1How it will be judged
A claim of absence. Four long sessions without maze from two of the maze rats, split at the midpoint of the recording; measures as in maze-swr-kc-increase; the paper names no test, so Wilcoxon signed rank (M-stats, paired) at p < 0.05. With n = 4 the smallest two-sided p is 0.125, so the test cannot find the effect and the claim is reproduced on it whenever the four sessions are present; the size and sign of each half-to-half change are reported, and whether all four share the sign of the maze effect is shown as a variant.
- maze-ripple-power-in-spindles
After maze exploration, CA1 SWR power within spindle cycles increases.
there was a significant increase in CA1 SWR power within spindle cycles (p=0.03, Wilcoxon signed rank
Results, Multi-region correlations after spatial exploration, paragraph 2How it will be judged
Mean ripple power in spindle cycles (as in spindle-cycles-by-ripple-power) per session, before against after the maze, Wilcoxon signed rank over sessions. Reproduced when it increases in all six sessions.
- fig7c-run-units-in-spindles
CA1 units active on the maze are more active in high ripple power spindle cycles than units not active on the maze, with independent effects of activity on the maze and of ripple power.
Units that were active during maze exploration were more likely to be active within spindle cycles with high ripple power than units not active on the maze (p=0.009, Mann-Whitney U-test for the mean number of spikes in spindle cycles and p=0.039 for the percent of spindle cycles with unit spikes ... found both factors to have a significant effect on the spike metrics (p<0.007 for ‘RUN compared to not in RUN’; p<0.01 for ‘low versus high ripple power’), with no interaction between the two factors (p>0.15
Results, Multi-region correlations after spatial exploration, paragraph 2; Figure 7A, CHow it will be judged
Needs the maze sessions. Run units have spikes on the maze; the paper classified them by visual inspection of clusters (Results, same paragraph), so the readings are (a) any labels the dataset gives, (b) units with at least 50 spikes in the 30 min maze epoch, (c) units with any spike in it (thresholds the paper does not give). Cycles from sleep after the maze, split at the median ripple power (spindle-cycles-by-ripple-power). Per unit the percent of high (and low) cycles with at least one spike and the mean spikes per cycle. Mann-Whitney U, two-way ANOVA (run by ripple power) and Tukey-Kramer as the paper names. Reproduced when, under a reading, both Mann-Whitney tests give p < 0.05 with run units higher and both ANOVA main effects p < 0.05.
- maze-ca1-rates-similar
CA1 units active and not active on the maze do not differ in firing rate in sleep after the maze.
These two groups of CA1 units did not differ in firing rate in the sleep post-behavior (calculated as the mean number of spikes over the total sleep time, or as the reciprocal of the mean inter-spike interval; p>0.1, Mann-Whitney U-test
Results, Multi-region correlations after spatial exploration, paragraph 2How it will be judged
A claim of absence. Rates both ways the paper gives, Mann-Whitney U at p < 0.05, under each classification reading of fig7c-run-units-in-spindles. Reproduced when neither rate differs; medians reported.
- fig7d-run-units-peri-swr
CA1 units active on the maze have higher peaks in their peri-SWR histograms (5.09 against 2.34 spikes/s), with no difference between SWRs nested in spindles and others.
the peri-SWR histograms of CA1 cells that had been active during run showed significantly higher peaks (mean 5.09 ± 4.23 spikes/s), compared to those units that were only active in the sleep post-behavior (mean 2.34 ± 2.09 spikes/s; p<10 −2 , Mann-Whitney U-test ... we did not find differences between the maximum nor mean firing in SWRs that were nested within spindle cycles compared to a sample of an equal number of SWRs not coupled to spindles (p>0.3 for both units in run and units not active in run; Wilcoxon signed rank
Results, Multi-region correlations after spatial exploration, paragraph 3; Figure 7D leftHow it will be judged
Peri-SWR histograms in raw rate (M-PETH bins) in sleep after the maze. The paper gives no window for the peak; readings (a) maximum in -100 to 100 ms (the 200 ms window of fig3c-thalamic-decrease-at-swrs), (b) maximum over the whole histogram (M-PETH extent). Nested and non-nested SWRs as in thalamus-at-spindle-nested-swrs. Reproduced when, under a reading of the classification, run peaks are higher at p < 0.05, the nested comparison gives p >= 0.05 in both groups, and both values are within tolerance.
- fig7b-run-units-spindle-phase
Spikes of maze-active CA1 units in spindle-nested SWRs are locked to spindle phase (circular mean -1.48 rad); not-run units are not, and Rayleigh Z is higher for run units and for coupled SWRs.
the spikes fired by units active in run during SWRs nested in spindles occurred at preferential phases of the spindle oscillation; phase distribution examples are shown in Figure 7b for the units in 7a (circular mean = −1.48 ± 1.33; concentration = 0.63 ± 0.32; mean p value = 0.027 ± 0.05, Rayleigh test; n = 27 units). The phase modulation in the same spindle-nested SWRs, was not significant for units not active in run (circular mean = −1.20 ± 1.55; concentration = 0.51 ± 0.29; mean p value = 0.218 ± 0.28; n = 14 units
Results, Multi-region correlations after spatial exploration, paragraph 3; Figure 7B, 7D rightHow it will be judged
Spindle phase (M-phase) of each unit's spikes in SWRs nested in spindles and in uncoupled SWRs, sleep after the maze. Open, which spikes are in an SWR; readings (a) spikes within the above-threshold period of the SWR (M-SWR), (b) spikes within plus or minus 50 ms of its timestamp. Units with too few SWR spikes are left out (one in the paper; no count given). Two-way ANOVA on Rayleigh Z and Tukey-Kramer as named. Reproduced when, under a reading, both main effects give p < 0.05, run units in coupled SWRs have the highest Z, and the run-unit mean Rayleigh p is below 0.05.
- fig8-thalamus-unchanged-by-maze
Thalamic units show no significant change after maze exploration in KC and spindle histograms, spindle phase, or spindle modulation index.
Thalamic units also had similar peri-event histograms with mPFC spindle cycles before and after behavior ( Figure 8b ), and no change in spindle phase distribution mean or concentration values (p>0.3, Mann-Whitney U-test); the spindle modulation index increased on average after behavior although not significantly (0.52 ± 0.30 before maze exploration and 0.84 ± 1.03 after; p=0.43 Wilcoxon signed rank test
Results, Multi-region correlations after spatial exploration, paragraph 4; Figure 8A-BHow it will be judged
A claim of absence, on the paper's tests. Phase by M-phase and modulation index as in fig6-modulation-with-ripple-power (all cycles), per thalamic unit in sleep before and after the maze. Reproduced when the Mann-Whitney tests and the Wilcoxon test give p >= 0.05; the changes are reported.
- fig8e-ca1-spindle-trend
CA1 units tend to take part more in high ripple power spindle cycles after maze exploration (60% fire more), without a significant before-after effect; ripple power has a significant effect.
60% of the CA1 units fired more spikes in high-ripple power spindle cycles after exploration compared to before (p=0.12), and fired in a larger proportion of high-ripple power spindle cycles (p=0.08; Wilcoxon signed rank ... finding a significant difference in engagement in spindles with ‘Low-High ripple’ power (p=0) and no significant effect for the ‘Before-After’ factor (p>0.06
Results, Multi-region correlations after spatial exploration, paragraph 4; Figure 8C-EHow it will be judged
Measures as in fig7c-run-units-in-spindles for all CA1 units, before and after the maze. Open, the median for the ripple split; readings (a) per epoch, (b) over both epochs together. Reproduced when, under a reading, the fraction is within tolerance, the ANOVA ripple effect gives p < 0.05, and the before-after tests give p >= 0.05.
- thalamus-not-selective-for-run-cells
Thalamic firing does not differ between spindle cycles that hold spikes of maze-active CA1 units and those that hold spikes of other CA1 units, nor with cycles in which co-modulated CA1 pairs are active.
We found no significant differences between the peak or the average firing values of the cross-correlograms between the two types of spindles and thalamic units (p>0.3, Wilcoxon signed rank), nor any differences between the mean number of thalamic spikes per spindle cycle or the percent of spindle cycles with spikes from thalamic units (p>0.1
Results, Multi-region correlations after spatial exploration, paragraphs 5 and 6How it will be judged
A claim of absence. Spindle cycles after the maze containing spikes of run units against cycles with spikes of not-run units; peri-spindle thalamic histograms (M-PETH) and per-cycle measures as in fig7c-run-units-in-spindles; Wilcoxon signed rank over thalamic units (named for the first, M-stats for the rest). CA1 pairs co-modulated during the maze were chosen by eye (Figure 7, figure supplement 1B); reading, pairs whose maze cross-correlogram peak within plus or minus 100 ms exceeds 3 sd (the pair rule of fig4b-thalamic-pairs-correlated, not stated for these pairs). Reproduced when no test gives p < 0.05; repeated on thalamic MUA as the paper did.
Pipeline
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