Title : Examining consolidation-dependent schema updating using continuous 1 memory measures 2 3

21 Memory schemas allow us to make efficient judgments about the world without the 22 cost of memorizing every detail of previous experiences. It has long been known that 23 schemas can enhance long-term memory for related information. The usefulness of 24 schemas, however, critically depends on their adaptability: how flexibly a schema can 25 be updated according to changing environmental conditions. It is difficult to track the 26 dynamic updating of individual memories with traditional memory measures. Using a 27 continuous report paradigm we show that schematization effects increase with 28 consolidation, that a group which was given time to consolidate showed increased 29 evidence of schema updating compared to a group that was not given time to 30 consolidate, and that the strength with which schemas are initially established 31 predicts to what degree participants let schemas guide their responses in a later test. 32 These results provide a novel perspective on the dynamic knowledge structures that 33 guide everyday behaviour. 34 The term ‘schema’ refers to complex knowledge structures within memory that 35 connect overlapping elements of information. Schemas are believed to support the 36 retention of information derived from a multitude of individual episodic memories, and 37 to help us generalize from previous experiences and predict the outcomes of future 38 events (e.g., Ghosh & Gilboa, 2014; van Kesteren, Ruiter, Fernández, & Henson, 39 2012). It has long been known that memory schemas are beneficial for the 40 remembering of schema congruent versus incongruent information (Bartlett, 1932; 41 Bransford & Johnson, 1972). This effect has been attributed to both enhanced deep 42 encoding (Staresina, Gray, & Davachi, 2009) and increases in schema-consistent 43 guesses (Rojahn & Pettigrew, 1992; Sakamoto & Love, 2004). During encoding, 44 schema congruency can accelerate the onset of memory formation processes 45 (Packard et al., 2017). During retrieval, the automatic activation of related information 46 in response to a retrieval cue enhances retrieval success (van Kesteren et al., 2012). 47 Thus, schemas contribute to memory performance by influencing processing during 48 both encoding and retrieval. 49 50 The development of knowledge structures such as schemas relies heavily on 51 consolidation, the process by which memories stabilize over time (Nadel & 52 Moscovitch, 1997; Squire & Alvarez, 1995). Consolidated schemas provide us with a 53 framework to predict outcomes according to our knowledge of events that fit the 54 existing schema. Models of reinforcement learning regard the drive to optimally 55 predict outcomes of the world as one of the main sources of learning (cf. Schultz, 56 Dayan, & Montague, 1997). Erroneous predictions due to inconsistent information 57 (i.e., unexpected outcomes) may be one source of such learning (Sutton & Barto, 58 1998). Similar mechanisms have been postulated in the long-term memory literature: 59 predictive coding models of memory are based on the idea that learning results from 60 divergences between experience-based predictions and observed outcomes 61 (Henson & Gagnepain, 2010; cf. Krawczyk, Fernández, Pedreira, & Boccia, 2017). 62 63 For an organism to be able to make predictions based on schematized memories, a 64 consolidated schema needs to exist to guide those predictions. Findings consistent 65 with this view have recently been reported by Richards and colleagues (2014), who 66 used a water-maze task in which mice had to learn to swim to platforms that 67 clustered according to a location schema. The authors found that a 30vs. 1-day 68 consolidation period augmented the formation of schemas: mice were better at 69 finding schema-consistent platforms with the longer compared to the shorter delay 70 after learning a location schema. Furthermore, consolidated schema memories 71 increased sensitivity to information that was inconsistent with the schema: mice 72 initially displayed higher errors, but more learning of this information was evident in a 73 later final test. That is, the authors found stronger updating of consolidated schemas 74 in the face of inconsistent information. Moreover, the authors hypothesized that the 75 area responsible for the updating of schemas would be the medial prefrontal cortex 76 (MPFC). Consistently, they found that pharmacological inhibition of MPFC impaired 77 updating of the schema. In line with this finding by Richards et al., human fMRI and 78 patient work has also postulated a role of (ventro-) medial PFC in schematic 79 memories (e.g., Brod & Shing, 2018; Ghosh, Moscovitch, Melo Colella, & Gilboa, 8