Aging Weakens Memory for Schema-Deviant Objects and Decouples Gaze Sampling from Retrieval Decisions
Abstract
1. Introduction
2. Materials and Methods
2.1. Participants
2.2. Experimental Materials
2.3. Design and Procedures
2.4. Acquisition of Eye Movement Data
2.5. Analysis of Subjective Ratings
2.6. Analysis of Memory Performance
2.7. Analysis of Eye Movement Data
2.8. Representational Similarity Analysis
2.9. Robustness Analysis of RSA
2.10. Transparency and Reproducibility
3. Results
3.1. Age Differences in Subjective Ratings
3.2. Age Differences in True Memory Performance
3.3. Age Differences in False Memory Performance
3.4. Age Differences in Response Time
3.5. Age Differences in Eye Movement
3.6. Age Differences in Representational Similarity
4. Discussion
Supplementary Materials
Author Contributions
Funding
Institutional Review Board Statement
Informed Consent Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
- Bartlett, F.C. Remembering: A Study in Experimental and Social Psychology; Cambridge University Press: Cambridge, UK, 1995. [Google Scholar]
- Schacter, D.L.; Addis, D.R. The cognitive neuroscience of constructive memory: Remembering the past and imagining the future. Philos. Trans. R. Soc. B Biol. Sci. 2007, 362, 773–786. [Google Scholar] [CrossRef]
- Schacter, D.L.; Thakral, P.P. Constructive memory and conscious experience. J. Cogn. Neurosci. 2024, 36, 1567–1577. [Google Scholar] [CrossRef] [PubMed]
- Ghosh, V.E.; Gilboa, A. What is a memory schema? A historical perspective on current neuroscience literature. Neuropsychologia 2014, 53, 104–114. [Google Scholar] [CrossRef] [PubMed]
- Sekeres, M.J.; Schomaker, J.; Nadel, L.; Tse, D. To update or to create? The influence of novelty and prior knowledge on memory networks. Philos. Trans. B 2024, 379, 20230238. [Google Scholar] [CrossRef] [PubMed]
- Bransford, J.D.; Johnson, M.K. Contextual prerequisites for understanding: Some investigations of comprehension and recall. J. Verbal Learn. Verbal Behav. 1972, 11, 717–726. [Google Scholar] [CrossRef]
- Gilboa, A.; Marlatte, H. Neurobiology of schemas and schema-mediated memory. Trends Cogn. Sci. 2017, 21, 618–631. [Google Scholar] [CrossRef]
- Bein, O.; Reggev, N.; Tompary, A. Working with schemas, predicting with schemas. J. Neurosci. 2018, 38, 1608–1610. [Google Scholar] [CrossRef]
- Brewer, W.F.; Treyens, J.C. Role of schemata in memory for places. Cogn. Psychol. 1981, 13, 207–230. [Google Scholar] [CrossRef]
- Moscovitch, D.A.; Moscovitch, M.; Sheldon, S. Neurocognitive model of schema-congruent and-incongruent learning in clinical disorders: Application to social anxiety and beyond. Perspect. Psychol. Sci. 2023, 18, 1412–1435. [Google Scholar] [CrossRef]
- Roediger, H.L., III; Abel, M. The double-edged sword of memory retrieval. Nat. Rev. Psychol. 2022, 1, 708–720. [Google Scholar] [CrossRef]
- Chen, X.; Varghese, L.; Jagust, W.J. A double-edged sword: The role of prior knowledge in memory aging. Front. Aging Neurosci. 2022, 14, 874767. [Google Scholar] [CrossRef] [PubMed]
- Wylie, L.E.; Patihis, L.; McCuller, L.L.; Davis, D.; Brank, E.M.; Loftus, E.F.; Bornstein, B.H. Misinformation effect in older versus younger adults: A meta-analysis and review. In The Elderly Eyewitness in Court; Taylor and Francis: Abingdon, UK, 2014; pp. 38–66. [Google Scholar]
- Schacter, D.L.; Koutstaal, W.; Norman, K.A. False memories and aging. Trends Cogn. Sci. 1997, 1, 229–236. [Google Scholar] [CrossRef] [PubMed]
- Colombel, F.; Tessoulin, M.; Gilet, A.-L.; Corson, Y. False memories and normal aging: Links between inhibitory capacities and monitoring processes. Psychol. Aging 2016, 31, 239. [Google Scholar] [CrossRef] [PubMed]
- Dennis, N.A.; Overman, A.A.; Carpenter, C.M.; Gerver, C.R. Understanding associative false memories in aging using multivariate analyses. Aging Neuropsychol. Cogn. 2022, 29, 500–525. [Google Scholar] [CrossRef]
- Popoviciu, A.; Richmond, L.L. A case for characterizing declarative memory commission errors in healthy aging. Front. Cogn. 2025, 3, 1505492. [Google Scholar] [CrossRef]
- McClelland, J.L.; McNaughton, B.L.; O’Reilly, R.C. Why there are complementary learning systems in the hippocampus and neocortex: Insights from the successes and failures of connectionist models of learning and memory. Psychol. Rev. 1995, 102, 419. [Google Scholar] [CrossRef]
- McClelland, J.L.; McNaughton, B.L.; Lampinen, A.K. Integration of new information in memory: New insights from a complementary learning systems perspective. Philos. Trans. R. Soc. B 2020, 375, 20190637. [Google Scholar] [CrossRef]
- Van Kesteren, M.T.; Ruiter, D.J.; Fernández, G.; Henson, R.N. How schema and novelty augment memory formation. Trends Neurosci. 2012, 35, 211–219. [Google Scholar] [CrossRef]
- Van Kesteren, M.T.; Beul, S.F.; Takashima, A.; Henson, R.N.; Ruiter, D.J.; Fernández, G. Differential roles for medial prefrontal and medial temporal cortices in schema-dependent encoding: From congruent to incongruent. Neuropsychologia 2013, 51, 2352–2359. [Google Scholar] [CrossRef]
- Hasher, L.; Campbell, K.L. Inhibitory theory: Assumptions, findings, and relevance to interventions. In The Cambridge Handbook of Cognitive Aging; Cambridge University Press: Cambridge, UK, 2020. [Google Scholar]
- Hasher, L.; Zacks, R.T. Working memory, comprehension, and aging: A review and a new view. Psychol. Learn. Motiv. 1988, 22, 193–225. [Google Scholar]
- Amer, T.; Wynn, J.S.; Hasher, L. Cluttered memory representations shape cognition in old age. Trends Cogn. Sci. 2022, 26, 255–267. [Google Scholar] [CrossRef] [PubMed]
- Campbell, K.L.; Lustig, C.; Hasher, L. Aging and inhibition: Introduction to the special issue. Psychol. Aging 2020, 35, 605. [Google Scholar] [CrossRef] [PubMed]
- Amer, T.; Giovanello, K.S.; Nichol, D.R.; Hasher, L.; Grady, C.L. Neural correlates of enhanced memory for meaningful associations with age. Cereb. Cortex 2019, 29, 4568–4579. [Google Scholar] [CrossRef] [PubMed]
- Amer, T.; Campbell, K.L.; Hasher, L. Cognitive control as a double-edged sword. Trends Cogn. Sci. 2016, 20, 905–915. [Google Scholar] [CrossRef]
- Friedman, N.P.; Robbins, T.W. The role of prefrontal cortex in cognitive control and executive function. Neuropsychopharmacology 2022, 47, 72–89. [Google Scholar] [CrossRef]
- Van der Linden, M.; Meulemans, T.; Marczewski, P.; Collette, F. The relationships between episodic memory, working memory, and executive functions: The contribution of the prefrontal cortex. Psychol. Belg. 2000, 40, 275–297. [Google Scholar] [CrossRef]
- Wynn, J.S.; Ryan, J.D.; Moscovitch, M. Effects of prior knowledge on active vision and memory in younger and older adults. J. Exp. Psychol. Gen. 2020, 149, 518. [Google Scholar] [CrossRef]
- Damiano, C.; Walther, D.B. Distinct roles of eye movements during memory encoding and retrieval. Cognition 2019, 184, 119–129. [Google Scholar] [CrossRef]
- Hannula, D.E.; Baym, C.L.; Warren, D.E.; Cohen, N.J. The eyes know: Eye movements as a veridical index of memory. Psychol. Sci. 2012, 23, 278–287. [Google Scholar] [CrossRef]
- Shing, Y.L.; Brod, G.; Greve, A. Prediction error and memory across the lifespan. Neurosci. Biobehav. Rev. 2023, 155, 105462. [Google Scholar] [CrossRef]
- Ramey, M.M.; Yonelinas, A.P.; Henderson, J.M. How schema knowledge influences memory in older adults: Filling in the gaps, or leading memory astray? Cognition 2024, 250, 105826. [Google Scholar] [CrossRef]
- Yu, J.; Li, J.; Huang, X. The Beijing version of the Montreal Cognitive Assessment as a brief screening tool for mild cognitive impairment: A community-based study. BMC Psychiatry 2012, 12, 156. [Google Scholar] [CrossRef]
- Yeung, P.Y.; Wong, L.L.; Chan, C.C.; Yung, C.Y.; Leung, L.J.; Tam, Y.Y.; Tang, L.N.; Li, H.S.; Lau, M.L. Montreal cognitive assessment—Single cutoff achieves screening purpose. Neuropsychiatr. Dis. Treat. 2020, 16, 2681–2687. [Google Scholar] [CrossRef] [PubMed]
- Connell, L.; Keane, M.T. A model of plausibility. Cogn. Sci. 2006, 30, 95–120. [Google Scholar] [CrossRef] [PubMed]
- Murphy, G.L.; Wisniewski, E.J. Familiarity and plausibility in conceptual combination: Reply to Gagné and Spalding (2006). J. Exp. Psychol. Learn. Mem. Cogn. 2006, 32, 1431–1442. [Google Scholar] [CrossRef]
- Ramm, B.J.; Halford, G.S. Novelty and processing demands in conceptual combination. Aust. J. Psychol. 2012, 64, 199–208. [Google Scholar] [CrossRef]
- Pastukhov, A. eyelinkReader: Import Gaze Data for EyeLink Eye Tracker, R Package Version 1.0.0 Computer Software; The Comprehensive R Archive Network (CRAN): Vienna, Austria, 2022. Available online: https://CRAN.R-project.org/package=eyelinkReader (accessed on 3 December 2025).
- Andersson, R.; Larsson, L.; Holmqvist, K.; Stridh, M.; Nyström, M. One algorithm to rule them all? An evaluation and discussion of ten eye movement event-detection algorithms. Behav. Res. Methods 2017, 49, 616–637. [Google Scholar] [CrossRef]
- Holmqvist, K.; Nyström, M.; Andersson, R.; Dewhurst, R.; Jarodzka, H.; Van de Weijer, J. Eye Tracking: A Comprehensive Guide to Methods and Measures; Oxford University Press: Oxford, UK, 2011. [Google Scholar]
- Chamberlain, L. Eye tracking methodology; theory and practice. Qual. Mark. Res. Int. J. 2007, 10, 217–220. [Google Scholar] [CrossRef]
- Ryan, J.D.; Althoff, R.R.; Whitlow, S.; Cohen, N.J. Amnesia is a deficit in relational memory. Psychol. Sci. 2000, 11, 454–461. [Google Scholar] [CrossRef]
- Hannula, D.E.; Ranganath, C. The eyes have it: Hippocampal activity predicts expression of memory in eye movements. Neuron 2009, 63, 592–599. [Google Scholar] [CrossRef]
- Wynn, J.S.; Buchsbaum, B.R.; Ryan, J.D. Encoding and retrieval eye movements mediate age differences in pattern completion. Cognition 2021, 214, 104746. [Google Scholar] [CrossRef]
- Heisz, J.J.; Ryan, J.D. The effects of prior exposure on face processing in younger and older adults. Front. Aging Neurosci. 2011, 3, 15. [Google Scholar] [CrossRef] [PubMed]
- Klever, L.; Islam, J.; Võ, M.L.-H.; Billino, J. Aging attenuates the memory advantage for unexpected objects in real-world scenes. Heliyon 2023, 9, e20241. [Google Scholar] [CrossRef] [PubMed]
- Brod, G.; Shing, Y.L. A boon and a bane: Comparing the effects of prior knowledge on memory across the lifespan. Dev. Psychol. 2019, 55, 1326. [Google Scholar] [CrossRef] [PubMed]
- Brod, G.; Shing, Y.L. Are there age-related differences in the effects of prior knowledge on learning? Insights gained from the memory congruency effect. Mind Brain Educ. 2022, 16, 89–98. [Google Scholar] [CrossRef]
- Cangelosi, M.; Rinaldi, L.; Dijkstra, T.; Palladino, P.; Cavallini, E. Older adults produce more verbal false memories than younger adults: Is it semantics or executive functioning? Aging Clin. Exp. Res. 2025, 37, 87. [Google Scholar] [CrossRef]
- Devitt, A.L.; Schacter, D.L. False memories with age: Neural and cognitive underpinnings. Neuropsychologia 2016, 91, 346–359. [Google Scholar] [CrossRef]
- Rayner, K. Eye movements in reading and information processing: 20 years of research. Psychol. Bull. 1998, 124, 372. [Google Scholar] [CrossRef]
- Orquin, J.L.; Loose, S.M. Attention and choice: A review on eye movements in decision making. Acta Psychol. 2013, 144, 190–206. [Google Scholar] [CrossRef]
- Krajbich, I.; Armel, C.; Rangel, A. Visual fixations and the computation and comparison of value in simple choice. Nat. Neurosci. 2010, 13, 1292–1298. [Google Scholar] [CrossRef]
- Cavanagh, J.F.; Wiecki, T.V.; Kochar, A.; Frank, M.J. Eye tracking and pupillometry are indicators of dissociable latent decision processes. J. Exp. Psychol. Gen. 2014, 143, 1476. [Google Scholar] [CrossRef]
- Li, Z.-W.; Ma, W.J. An uncertainty-based model of the effects of fixation on choice. PLoS Comput. Biol. 2021, 17, e1009190. [Google Scholar] [CrossRef] [PubMed]
- Kim, A.J.; Senior, J.; Chu, S.; Mather, M. Aging impairs reactive attentional control but not proactive distractor inhibition. J. Exp. Psychol. Gen. 2024, 153, 1938. [Google Scholar] [CrossRef] [PubMed]
- Braver, T.S. The variable nature of cognitive control: A dual mechanisms framework. Trends Cogn. Sci. 2012, 16, 106–113. [Google Scholar] [CrossRef] [PubMed]
- Just, M.A.; Carpenter, P.A. A theory of reading: From eye fixations to comprehension. Psychol. Rev. 1980, 87, 329. [Google Scholar] [CrossRef]
- Korkki, S.M.; Richter, F.R.; Gellersen, H.M.; Simons, J.S. Reduced memory precision in older age is associated with functional and structural differences in the angular gyrus. Neurobiol. Aging 2023, 129, 109–120. [Google Scholar] [CrossRef]
- Zheng, L.; Gao, Z.; Xiao, X.; Ye, Z.; Chen, C.; Xue, G. Reduced fidelity of neural representation underlies episodic memory decline in normal aging. Cereb. Cortex 2018, 28, 2283–2296. [Google Scholar] [CrossRef]
- Webb, C.E.; Dennis, N.A. Differentiating true and false schematic memories in older adults. J. Gerontol. Ser. B 2019, 74, 1111–1120. [Google Scholar] [CrossRef]
- Wedel, M.; Pieters, R.; van der Lans, R. Modeling eye movements during decision making: A review. Psychometrika 2023, 88, 697–729. [Google Scholar] [CrossRef]
- Walshe, R.C.; Nuthmann, A. A computational dual-process model of fixation-duration control in natural scene viewing. Comput. Brain Behav. 2021, 4, 463–484. [Google Scholar] [CrossRef]
- Fisher, G. A multiattribute attentional drift diffusion model. Organ. Behav. Hum. Decis. Process. 2021, 165, 167–182. [Google Scholar] [CrossRef]
- Molter, F.; Thomas, A.W.; Huettel, S.A.; Heekeren, H.R.; Mohr, P.N. Gaze-dependent evidence accumulation predicts multi-alternative risky choice behaviour. PLoS Comput. Biol. 2022, 18, e1010283. [Google Scholar] [CrossRef] [PubMed]
- Braem, S.; Bugg, J.M.; Schmidt, J.R.; Crump, M.J.; Weissman, D.H.; Notebaert, W.; Egner, T. Measuring adaptive control in conflict tasks. Trends Cogn. Sci. 2019, 23, 769–783. [Google Scholar] [CrossRef] [PubMed]
- Gajewski, P.D.; Falkenstein, M.; Thönes, S.; Wascher, E. Stroop task performance across the lifespan: High cognitive reserve in older age is associated with enhanced proactive and reactive interference control. NeuroImage 2020, 207, 116430. [Google Scholar] [CrossRef]
- Peng, Y.; Zhu, Q.; Wang, B.; Ren, J. A cross-sectional study on interference control: Age affects reactive control but not proactive control. PeerJ 2020, 8, e8365. [Google Scholar] [CrossRef]
- Ebaid, D.; Crewther, S.G. The contribution of oculomotor functions to rates of visual information processing in younger and older adults. Sci. Rep. 2020, 10, 10129. [Google Scholar] [CrossRef]
- Chamberlain, J.D.; Bowman, C.R.; Dennis, N.A. Age-related differences in encoding-retrieval similarity and their relationship to false memory. Neurobiol. Aging 2022, 113, 15–27. [Google Scholar] [CrossRef]
- O’Reilly, R.C.; Norman, K.A. Hippocampal and neocortical contributions to memory: Advances in the complementary learning systems framework. Trends Cogn. Sci. 2002, 6, 505–510. [Google Scholar] [CrossRef]
- Owsley, C. Vision and aging. Annu. Rev. Vis. Sci. 2016, 2, 255–271. [Google Scholar] [CrossRef]
- Savitz, D.A.; Wellenius, G.A. Can cross-sectional studies contribute to causal inference? It depends. Am. J. Epidemiol. 2023, 192, 514–516. [Google Scholar] [CrossRef]
- Almidani, L.; Mihailovic, A.; Varadaraj, V.; Swenor, B.K.; Ramulu, P.Y. Longitudinal changes in visual acuity and contrast sensitivity and incident dementia. Am. J. Ophthalmol. 2024, 266, 227–234. [Google Scholar] [CrossRef] [PubMed]
- Dhara, G.; Kumar, R.K. A survey on visual saliency detection approaches and attention models. Multimed. Tools Appl. 2025, 84, 44183–44225. [Google Scholar] [CrossRef]
- Endemann, R.; Kamp, S.-M. Examining the role of stimulus complexity in item and associative memory. Mem. Cogn. 2025, 53, 628–644. [Google Scholar] [CrossRef] [PubMed]
- Kyle-Davidson, C.; Solis, O.; Robinson, S.; Tan, R.T.W.; Evans, K.K. Scene complexity and the detail trace of human long-term visual memory. Vis. Res. 2025, 227, 108525. [Google Scholar] [CrossRef]
- Beason-Held, L.L.; Shafer, A.T.; Goh, J.O.; Landman, B.A.; Davatzikos, C.; Viscomi, B.; Ash, J.; Kitner-Triolo, M.; Ferrucci, L.; Resnick, S.M. Hippocampal activation and connectivity in the aging brain. Brain Imaging Behav. 2021, 15, 711–726. [Google Scholar] [CrossRef]
- Nyberg, L.; Andersson, M.; Lundquist, A.; Salami, A.; Wåhlin, A. Frontal contribution to hippocampal hyperactivity during memory encoding in aging. Front. Mol. Neurosci. 2019, 12, 229. [Google Scholar] [CrossRef]
- Andersson, P.; Samrani, G.; Andersson, M.; Persson, J. Hippocampal subfield volumes contribute to working memory interference control in aging: Evidence from longitudinal associations over 5 years. Neuroimage Rep. 2023, 3, 100189. [Google Scholar] [CrossRef]
- Li, Z.; He, H.; Chen, Y.; Guan, Q. Effects of engagement, persistence and adherence on cognitive training outcomes in older adults with and without cognitive impairment: A systematic review and meta-analysis of randomised controlled trials. Age Ageing 2024, 53, afad247. [Google Scholar] [CrossRef]






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Xu, H.-Z.; Huan, S.-Y. Aging Weakens Memory for Schema-Deviant Objects and Decouples Gaze Sampling from Retrieval Decisions. Brain Sci. 2026, 16, 289. https://doi.org/10.3390/brainsci16030289
Xu H-Z, Huan S-Y. Aging Weakens Memory for Schema-Deviant Objects and Decouples Gaze Sampling from Retrieval Decisions. Brain Sciences. 2026; 16(3):289. https://doi.org/10.3390/brainsci16030289
Chicago/Turabian StyleXu, Hong-Zhou, and Sheng-Yin Huan. 2026. "Aging Weakens Memory for Schema-Deviant Objects and Decouples Gaze Sampling from Retrieval Decisions" Brain Sciences 16, no. 3: 289. https://doi.org/10.3390/brainsci16030289
APA StyleXu, H.-Z., & Huan, S.-Y. (2026). Aging Weakens Memory for Schema-Deviant Objects and Decouples Gaze Sampling from Retrieval Decisions. Brain Sciences, 16(3), 289. https://doi.org/10.3390/brainsci16030289

