Figure 1B shows the location of a typical CA1
recording site in one patient; Fig. 1C is a sche-
matic drawing of the depth electrode used in
our study (locations of hippocampal recording
sites in each patient are depicted in fig. S1).
Figure 1D shows typical SWR events as they
appear in the recordings. Our dataset included
8387 SWRs obtained from 15 patients (see Fig. 1,
E and F, for grand-average peri-ripple field po-
tential and spectral decomposition and Fig. 1G
for distribution of inter-ripple interval durations;
see table S1 for demographic information).
We analyzed four main conditions: (i) rest,
during which patients were instructed to rest
with eyes closed; (ii) viewing, during which pa-
tients inspected and memorized photographs of
famous faces and places; (iii) recall, referring to
times within the free-recall period in which pa-
tients verbally reported recalling a specific item
from the memorized set (i.e., epochs that began3 s before the onset until the offset of each
“recall event,”a total of 269.5 s on average);
and (iv) search, referring to all complementary
time intervals between recall events, in which
patients attempted to recall but did not report
any recalled item (330.5 s on average).SWR properties across cognitive states
We first examined whether the spectral signa-
tureoftheSWRsvariedacrossthedifferentNormanet al.,Science 365 , eaax1030 (2019) 16 August 2019 2of14
Time (sec)-3000300Hippocampalraw LFP(μV)-10010Ripple-band70-180Hz(μV)0 1 2 3 4 5010
4
Squaredenvelope(z score)detection thr.Voltage (μV)dB010-200-100 0 10020050100150200Frequency (Hz)Time (ms)-500 -250 0 250 500-40-20020Grand-average
n = 8387 ripples (15 patients)Time from ripple peak (ms)0510Mean = 2.4 sInter ripple interval (s)Probability
00.20.4Depth iEEG electrodes: 4.4mm
0.8mm
2mm
platinum contactsPatient 4: Left HippocampusCA1subiculumpresubiculum
parasubiculumHippocampal subfields:molecular layer
hippocampal fissureCA2/3fimbria
GC-DGrecording sitesAPRLRUN 1
time rest (200 s) viewing (130 s) counting1500 ms 750 msrecalling faces
(150 s)verbalized free-recallrecalling places
(150 s)time rest (200 s) viewing (130 s) counting recalling faces
(150 s)recalling places
(150 s)RUN 2
timeblindfold onE
A
B C
D
F G
Fig. 1. Experimental design and hippocampal SWR detection.
(A) Experimental design and stimuli ( 91 ). After viewing pictures of famous
faces and places (see methods), participants were asked to freely recall
and describe as many pictures as possible, targeting each category in
separate blocks. (B) Coronal slice and 3D reconstruction of a hippocampal
depth electrode in one representative patient. White arrow indicates
CA1 recording site used for ripple detection. (C) Schematic diagram of
depth iEEG electrodes used in our study. (D) Example of SWR events
as they appear in the recordings. From top to bottom: raw hippocampal
LFP; ripple-band filtered LFP (70 to 180 Hz); normalized ripple-band
envelope used for ripple detection. (EandF) Grand average peri-ripple
field potential and wavelet spectrogram centered on ripple peak
(n= 8279 SWR events from 15 patients). (G) Overall distribution of inter-
ripple intervals (n= 15 patients; error bars represent SEM).RESEARCH | RESEARCH ARTICLE