Human hippocampal ripples signal encoding of episodic memories

Recent human electrophysiology work has uncovered the presence of high frequency oscillatory events, termed ripples, during awake behavior. This prior work focuses on ripples in the medial temporal lobe (MTL) during memory retrieval. Few studies, however, investigate ripples during item encoding. Many studies have found neural activity during encoding that predicts later recall, termed subsequent memory effects (SMEs), but it is unclear if ripples during encoding also predict subsequent recall. Detecting ripples in 124 neurosurgical participants performing an episodic memory task, we find insignificant ripple SMEs in any MTL region, even as these regions exhibit robust high frequency activity (HFA) SMEs. Instead, hippocampal ripples increase during encoding of items leading to recall of temporally or semantically associated items, a phenomenon known as clustering. This subsequent clustering effect (SCE) arises specifically when hippocampal ripples occur during both encoding and retrieval, suggesting that ripples mediate the encoding and future reinstatement of episodic memories.