Navigational Mind
00 — The ‘seat’ of memory
Before 1953, memory was not associated with any one area of the brain. Lashley spent decades looking for the trace and though this is a long story and lots of debate, the idea was sort of that memory is everywhere and nowhere. But epilepsy and surgeons removing the medial temporal lobes of patients brought scientists to focus on the hippocampus. As part of this, Henry Moalison, a 27-year-old man with intractable epilepsy, had his hippocampi revoved and then his seizures improved but he could not form any new memories, or could not attach anything new to the memory he already had.
Lashley (1950), In search of the engram — the view H.M. overturned https://scholar.google.com/scholar?q=Lashley+1950+In+search+of+the+engram
Scoville & Milner (1957) — Loss of recent memory after bilateral hippocampal lesions. The paper that gave memory a location. Free full text. https://pmc.ncbi.nlm.nih.gov/articles/PMC497229/
Milner (1962) — the mirror-drawing task. The most important experiment on the page. H.M. traced a star in a mirror, improved across days, and each day denied ever having done it. Skill and recollection came apart in one afternoon's data — memory was not one thing. https://scholar.google.com/scholar?q=Milner+1962+mirror+drawing+H.M.+hippocampal+lesions+motor+skill
Milner, Corkin & Teuber (1968) — 14-year follow-up; Corkin takes over the work https://scholar.google.com/scholar?q=Milner+Corkin+Teuber+1968+further+analysis+hippocampal+amnesic+syndrome
Corkin (2002) — What's new with the amnesic patient H.M.? Best single summary. https://scholar.google.com/scholar?q=Corkin+2002+what%27s+new+with+the+amnesic+patient+H.M.
Corkin (2013),Permanent Present Tense — often recommended to non-scientists. https://scholar.google.com/scholar?q=Corkin+Permanent+Present+Tense+amnesic+patient+H.M.
Squire (2009) — The legacy of patient H.M. for neuroscience. Written after his 2008 death, when his name could finally bepublished. https://scholar.google.com/scholar?q=Squire+2009+legacy+of+patient+H.M.+for+neuroscience
Annese et al. (2014) — postmortem histological reconstruction of H.M.'s brain. Revised what Scoville had actually removed. https://scholar.google.com/scholar?q=Annese+2014+postmortem+examination+patient+H.M.+brain+3D+reconstruction
Milner & Klein (2016) — Milner revisiting the 1957 paper nearly sixty years on https://www.mcgill.ca/denise-klein-lab/sites/denise-klein-lab//files/230.full_.pdf
Brenda Milner interviews — she tells this story on camera extremely well; possibly your opening video for the whole site https://www.youtube.com/results?search_query=Brenda+Milner+interview+H.M.+memory+neuroscience
Note: The operation was experimental, performed on a patient in distress, and it destroyed a man's capacity to build a life in exchange for knowledge the rest of us now have. Scoville came to regard it as a mistake. H.M. spent fifty-five years being tested by researchers he met, each time, for the first time.
01 — The cognitive map
A cognitive map is often defined as a type of mental representation used by an individual to order their personal store of information about their everyday or metaphorical spatial environment, and the relationship of its component parts. The concept was introduced by Edward Tolman in 1948. He tried to explain the behavior of rats that appeared to learn the spatial layout of a maze, and subsequently the concept was applied to other animals, including humans The term was later generalized by some researchers, especially in the field of operations research, to refer to a kind of semantic network representing an individual's personal knowledge or schemas.
What is happening with the navigataional approach is that the map is becoming a ‘mapping’ and there are not stored internal representations but rahter different actions of communication at many scales, from the way we communicate with our own bodies as those bodies to the ways we communicate with all we encounter.
Tolman (1948), Cognitive maps in rats and men — never only about space https://scholar.google.com/scholar?q=Tolman+1948+Cognitive+maps+in+rats+and+men
O'Keefe & Dostrovsky (1971) — the first place cell https://scholar.google.com/scholar?q=O%27Keefe+Dostrovsky+1971+hippocampus+spatial+map
O'Keefe & Nadel (1978), The Hippocampus as a Cognitive Maphttps://scholar.google.com/scholar?q=O%27Keefe+Nadel+The+Hippocampus+as+a+Cognitive+Map+1978
Hafting et al. (2005) — grid cells https://scholar.google.com/scholar?q=Hafting+2005+Microstructure+spatial+map+entorhinal+cortex
Discovering the grid cells — Kavli Institute NTNU https://www.ntnu.edu/kavli/discovering-grid-cells
Nobel Prize 2014 — O'Keefe and the Mosers, incl. lectures https://www.nobelprize.org/prizes/medicine/2014/summary/
Grid cells, independent fields and phase precession (PNAS 2012) https://www.pnas.org/doi/10.1073/pnas.1109599109
Wilson lab / Picower — cognitive maps assembling over days of sleep https://biology.mit.edu/category/news-briefs/picower-institute-for-learning-and-memory/
02 — Seeing cells fire
Grid cell recordings — grey trajectory, red spike dots https://www.youtube.com/results?search_query=grid+cell+firing+rat+entorhinal+cortex+Moser
Place cell ensemble, Wilson lab MIT https://www.youtube.com/results?search_query=place+cell+recording+Matt+Wilson+hippocampus+video
"My Running Rat" — real entorhinal spikes scored into music https://blog.medisin.ntnu.no/en/rock-and-roll-rats-in-a-new-art-video/
Human grid cells — BrainFacts primer with video https://www.sfn.org/sitecore/content/home/brainfacts2/brain-anatomy-and-function/anatomy/2013/human-grid-cells
On the Grid — the history in one read https://www.brainfacts.org/brain-anatomy-and-function/cells-and-circuits/2017/on-the-grid-033117
03 — Concepts and structure
If the grid code represents relations, it should appear wherever a task has map-like structure — including tasks with nothing spatial in them.
Constantinescu, O'Reilly & Behrens (2016), Science — Organizing conceptual knowledge in humans with a gridlike code. The hinge paper: a 2D bird-morph concept space, six-fold entorhinal signal. https://scholar.google.com/scholar?q=Constantinescu+O%27Reilly+Behrens+2016+conceptual+knowledge+gridlike+code
Behrens et al. (2018), Neuron — What is a cognitive map? Organizing knowledge for flexible behavior. Best entry point to this whole section. https://www.cell.com/neuron/fulltext/S0896-6273(18)30856-0 Free preprint: https://www.biorxiv.org/content/10.1101/365593v1
Garvert, Dolan & Behrens (2017), eLife — a map of abstract relational knowledge https://scholar.google.com/scholar?q=Garvert+Dolan+Behrens+2017+abstract+relational+knowledge+hippocampal+entorhinal
Whittington et al. (2020), Cell — the Tolman-Eichenbaum Machine https://scholar.google.com/scholar?q=Whittington+2020+Tolman-Eichenbaum+Machine+unifying+space+relational+memory
Baram et al. (2021), Neuron — abstracting and generalizing reward-space structure https://scholar.google.com/scholar?q=Baram+Behrens+2021+entorhinal+ventromedial+prefrontal+structure+reward+space
Whittington, McCaffary, Bakermans & Behrens — How to build a cognitive maphttps://scholar.google.com/scholar?q=Whittington+Behrens+How+to+build+a+cognitive+map
El-Gaby et al. (2024) — a cellular basis for mapping behavioural structure https://scholar.google.com/scholar?q=El-Gaby+Behrens+2024+cellular+basis+mapping+behavioural+structure
Bakermans et al. (2025), Nat Neurosci — constructing future behaviour via composition & replay https://scholar.google.com/scholar?q=Bakermans+Behrens+2025+constructing+future+behaviour+composition+replay
Behrens Lab, Sainsbury Wellcome Centre — plain-language write-ups https://www.sainsburywellcome.org/web/groups/behrens-lab
Full publication list (live) — Oxford NDCN https://www.ndcn.ox.ac.uk/team/timothy-behrens
04 — Cognitive spaces: Doeller lab at Max Planck (where i did my master thesis) and others
The human-imaging side, pushed into theory: the hippocampal formation supplies a geometric format that any set of dimensions can be mapped into.
Doeller, Barry & Burgess (2010), Nature — evidence for grid cells in a human memory network. The hexadirectional fMRI method everything after depends on. https://scholar.google.com/scholar?q=Doeller+Barry+Burgess+2010+grid+cells+human+memory+network
Bellmund et al. (2016), eLife — grid-cell representations in mental simulation https://scholar.google.com/scholar?q=Bellmund+Doeller+2016+grid-cell+representations+mental+simulation
Deuker et al. (2016), eLife — an event map of memory space in the hippocampus https://scholar.google.com/scholar?q=Deuker+Bellmund+Doeller+2016+event+map+of+memory+space+hippocampus
Nau et al. (2018), Nat Neurosci — hexadirectional coding of visual space https://scholar.google.com/scholar?q=Nau+Bellmund+Doeller+2018+hexadirectional+coding+visual+space+entorhinal
Bellmund, Gärdenfors, Moser & Doeller (2018), Science — Navigating cognition: spatial codes for human thinking. The manifesto for this section. https://www.science.org/doi/10.1126/science.aat6766 Lab summary: https://doellerlab.com/cognitive-space/
Nau, Julian & Doeller (2018) — how the navigation system shapes visual experience https://scholar.google.com/scholar?q=Nau+Julian+Doeller+2018+navigation+system+shapes+visual+experience
Theves, Fernández & Doeller (2019), Curr Biol — distances in multidimensional feature space https://scholar.google.com/scholar?q=Theves+Doeller+2019+hippocampus+distances+multidimensional+feature+space
Theves et al. (2020) — the hippocampus maps concept space, not feature space https://scholar.google.com/scholar?q=Theves+Doeller+hippocampus+maps+concept+space+not+feature+space
Bottini & Doeller (2020), TiCS — knowledge across reference frames https://scholar.google.com/scholar?q=Bottini+Doeller+2020+knowledge+across+reference+frames+cognitive+maps+image+spaces
Doeller Lab (MPI CBS Leipzig / Kavli NTNU) — full publications https://doellerlab.com/
SOCIAL SPACE People navigate social networks same way they navigate transportation networks: by studying maps
MULTI-SENSORY (Olfactory & Visual): Distinct brain regions map olfactory and visual spaces
05 — Sound, smell, faces, status
The dimensions usually called "non-spatial."
Aronov, Nevers & Tank (2017), Nature — Mapping of a non-spatial dimension by the hippocampal–entorhinal circuit. The auditory one, and the cleanest demonstration going: rats move a sound's pitch with a joystick, cells fire at specific frequencies. https://scholar.google.com/scholar?q=Aronov+Nevers+Tank+2017+mapping+non-spatial+dimension+hippocampal+entorhinal
Eichenbaum (2014) — time cells in the hippocampus https://scholar.google.com/scholar?q=Eichenbaum+2014+time+cells+in+the+hippocampus
Tavares et al. (2015), Neuron — a map for social navigation in the human brain https://scholar.google.com/scholar?q=Tavares+Schiller+2015+map+for+social+navigation+human+brain
Park et al. (2021), Nat Neurosci — inferences on a social hierarchy use a grid-like code https://scholar.google.com/scholar?q=Park+Boorman+2021+multidimensional+social+hierarchy+grid-like+code
Schafer & Schiller (2018), Neuron — navigating social space (review) https://scholar.google.com/scholar?q=Schafer+Schiller+2018+navigating+social+space
Bao et al. (2019), Neuron — grid-like codes for a two-dimensional odour space https://scholar.google.com/scholar?q=Bao+2019+grid-like+olfactory+navigation+two-dimensional+odor+space
Viganò & Piazza (2020) — navigating a novel semantic space https://scholar.google.com/scholar?q=Vigano+Piazza+2020+distance+direction+codes+novel+semantic+space
Auditory object representation in the bat hippocampus: https://www.sciencedirect.com/science/article/abs/pii/S0960982225009534
Peer et al. (2021), TiCS — cognitive maps and cognitive graphs (useful corrective) https://scholar.google.com/scholar?q=Peer+Brunec+Newcombe+Epstein+2021+cognitive+maps+and+cognitive+graphs
Note: the hexadirectional fMRI signature is an indirect, population-level proxy for grid coding, not a recording of grid cells. Productive, and contested (like everything else).
06 — When the map is lost
Clive Wearing, The Man with the Seven Second Memoryhttps://www.youtube.com/results?search_query=Clive+Wearing+documentary+seven+second+memory
Oliver Sacks, "The Abyss" https://scholar.google.com/scholar?q=Sacks+The+Abyss+Clive+Wearing+musical+amnesia
Scoville & Milner (1957) — the H.M. paper https://scholar.google.com/scholar?q=Scoville+Milner+1957+loss+of+recent+memory+bilateral+hippocampal+lesions
Henry Molaison — archive footage and Suzanne Corkin interviews https://www.youtube.com/results?search_query=Henry+Molaison+H.M.+patient+documentary+Suzanne+Corkin
Maguire et al. (2006) — London taxi drivers vs. bus drivers https://scholar.google.com/scholar?q=Maguire+2006+London+taxi+drivers+bus+drivers+hippocampi
07 — Imagining is continuous with remembering?
Amnesic patients couldn't remember their past. Then someone asked them to imagine lying on a white sandy beach, and that was also a struggle and most couldn't do that either, so there is a link here. Memory isn't a recording you play back; it's a construction, and imagination runs on similar ‘habits’ and ‘machinery’.
Tulving (1985), Memory and consciousness — mental time travel; patient K.C., blank in both directions https://scholar.google.com/scholar?q=Tulving+1985+Memory+and+consciousness+autonoetic+mental+time+travel
Hassabis, Kumaran, Vann & Maguire (2007), PNAS — Patients with hippocampal amnesia cannot imagine new experiences. The central result. Imagined scenes lacked spatial coherence — fragments without a holding environment. Free full text. https://pmc.ncbi.nlm.nih.gov/articles/PMC1773058/
Hassabis & Maguire (2007), TiCS — deconstructing episodic memory with construction https://scholar.google.com/scholar?q=Hassabis+Maguire+2007+deconstructing+episodic+memory+with+construction
Addis, Wong & Schacter (2007) — remembering the past and imagining the future: common and distinct neural substrates https://scholar.google.com/scholar?q=Addis+Wong+Schacter+2007+remembering+past+imagining+future+neural+substrates
Schacter & Addis (2007) — the constructive episodic simulation hypothesis. The best account of why memory would be built this way, and why its flexibility is also its error source. https://scholar.google.com/scholar?q=Schacter+Addis+2007+constructive+memory+remembering+past+imagining+future
Schacter, Addis & Buckner (2007), Nat Rev Neurosci — Remembering the past to imagine the future: the prospective brain. The readable summary. https://scholar.google.com/scholar?q=Schacter+Addis+Buckner+2007+prospective+brain
Buckner & Carroll (2007), TiCS — Self-projection and the brain. Remembering, imagining, navigating and perspective-taking as one operation aimed different directions. Links to 05. https://scholar.google.com/scholar?q=Buckner+Carroll+2007+self-projection+and+the+brain
Squire et al. (2010) vs. Maguire & Hassabis (2011) — the exchange. Squire's group found amnesic patients who could imagine future events. Essential balance; link the exchange, not one side. https://www.pnas.org/doi/full/10.1073/pnas.1019643108
Race, Keane & Verfaellie (2011) — careful follow-up addressing the narrative-demands objection https://scholar.google.com/scholar?q=Race+Keane+Verfaellie+2011+episodic+memory+episodic+future+thinking+medial+temporal
Zeidman & Maguire (2016), Nat Rev Neurosci — anterior hippocampus: perception, imagination, episodic memory https://scholar.google.com/scholar?q=Zeidman+Maguire+2016+anterior+hippocampus+perception+imagination+episodic+memory
Addis (2020) — Mental time travel? A neurocognitive model of event simulation. The mature version, and a provocation about whether "time travel" describes either thing well.https://scholar.google.com/scholar?q=Addis+2020+mental+time+travel+neurocognitive+model+event+simulation
08 — Bodily action
Towards stronger claims: that remembering and imagining aren't just neurally similar but enacted — carried out by eyes, motor systems and place-cell sequences doing versions of what they do when you actually move and look.
Laeng & Teodorescu (2002) — eye scanpaths during visual imagery reenact those of perception, including in complete darkness https://scholar.google.com/scholar?q=Laeng+Teodorescu+2002+eye+scanpaths+visual+imagery+reenact+perception
Johansson & Johansson (2014), Psych Science — Look here, eye movements play a functional role in memory retrieval. The upgrade from correlation to function: force gaze to the wrong region during recall and memory gets worse. https://journals.sagepub.com/doi/abs/10.1177/0956797613498260
Wynn, Shen & Ryan (2019) — gaze reinstatement review, including where effects are inconsistent https://scholar.google.com/scholar?q=Wynn+Shen+Ryan+2019+eye+movements+reinstate+spatiotemporal+mnemonic+content
Ferreira, Apel & Henderson (2008) — Taking a new look at looking at nothing. People stare at the empty place where a thing used to be. https://scholar.google.com/scholar?q=Ferreira+Apel+Henderson+2008+taking+a+new+look+at+looking+at+nothing
Kent & Lamberts (2008), TiCS — retrieval as mental simulation https://scholar.google.com/scholar?q=Kent+Lamberts+2008+encoding-retrieval+relationship+retrieval+as+mental+simulation
Nyberg et al. (2001) — reactivation of motor areas during memory for actions https://scholar.google.com/scholar?q=Nyberg+2001+reactivation+of+motor+brain+areas+explicit+memory+for+actions
Jeannerod (2001) — neural simulation of action. Imagined movement takes about as long as real movement and obeys the same biomechanical constraints. https://scholar.google.com/scholar?q=Jeannerod+2001+neural+simulation+of+action+motor+cognition
Johnson & Redish (2007) — place cells sweep down each path in turn while a rat pauses at a junction. As close to watching an animal imagine as the literature gets. https://scholar.google.com/scholar?q=Johnson+Redish+2007+CA3+encode+paths+forward+decision+point
Pfeiffer & Foster (2013), Nature — place-cell sequences depict future paths to remembered goals https://scholar.google.com/scholar?q=Pfeiffer+Foster+2013+place-cell+sequences+future+paths+remembered+goals
Ólafsdóttir, Bush & Barry (2018) — replay in memory and planning (best rodent-side entry) https://scholar.google.com/scholar?q=Olafsdottir+Bush+Barry+2018+role+of+hippocampal+replay+in+memory+and+planning
Bellmund et al. (2016) — grid-cell representations in mental simulation (also in section 04; belongs in both) https://scholar.google.com/scholar?q=Bellmund+Doeller+2016+grid-cell+representations+mental+simulation
Barsalou (2008) — grounded cognition. The framework these findings get read through, and the handoff into section 11. https://scholar.google.com/scholar?q=Barsalou+2008+grounded+cognition+annual+review+psychology
Note: evidence is strong for shared ‘machinery’ — overlapping networks, re-enacted scanpaths, motor reactivation, sequences running both directions. Remebering and imagining are not the same but have operational coherence, is how I would put it.
09 —Selective attention; the worlds choose and the worlds chosen for us and the worlds we miss
The original Selective Attention Test https://www.theinvisiblegorilla.com/IGvideos.html
Simons & Chabris (1999), Gorillas in our midsthttps://scholar.google.com/scholar?q=Simons+Chabris+1999+gorillas+in+our+midst+inattentional+blindness
Simons on the sequel — expecting the gorilla doesn't help https://www.smithsonianmag.com/science-nature/but-did-you-see-the-gorilla-the-problem-with-inattentional-blindness-17339778/
Daniel Simons' full demo archive https://www.youtube.com/c/DanielSimons/videos
Drew, Võ & Wolfe (2013) — radiologists miss a gorilla on the CT scan https://scholar.google.com/scholar?q=Drew+Vo+Wolfe+2013+invisible+gorilla+strikes+again+expert+observers
Craik et al. (1996) — divided attention at encoding vs. retrieval https://scholar.google.com/scholar?q=Craik+1996+divided+attention+encoding+retrieval+processes
Uncapher & Wagner (2009) — attention and the subsequent memory effect https://scholar.google.com/scholar?q=Uncapher+Wagner+2009+divided+attention+subsequent+memory
10 — Attention writing to the hippocampus
Aly & Turk-Browne (2016), PNAS — Attention promotes episodic encoding by stabilizing hippocampal representations. The key paper. Free full text. https://pmc.ncbi.nlm.nih.gov/articles/PMC4743819/
Poskanzer et al. (2025) — encoding vs. retrieval states in the hippocampus https://www.ncbi.nlm.nih.gov/pmc/articles/PMC12373456/
Muzzio et al. (2009) — attention stabilizes place representations (rodent single-unit version; ties this section straight back to 01) https://scholar.google.com/scholar?q=Muzzio+2009+attention+stability+representations+dorsal+hippocampus
Tonegawa lab / Picower — selectively remembering new places; locus coeruleus and attention https://tonegawalab.mit.edu/news/in-the-news
Picower Institute — cognition & motivation research hubhttps://picower.mit.edu/research/cognition-and-motivation
MIT News (2017) — the circuit necessary for memory formation https://news.mit.edu/2017/neuroscientists-identify-brain-circuit-necessary-memory-formation-0406
11 — Rereading the map
If one system maps rooms, concepts, pitches and people, either "space" was the wrong word, or thinking was always more like moving than we assumed.
Hiott (2025), Topoi — Radical Embodied Relation at any Scale, from Remembering to Navigating. Open access. Recent hippocampal findings put old notions of representation in question; thinking and remembering can be assessed by the same process as navigating and wayfinding. https://link.springer.com/article/10.1007/s11245-025-10256-7
Hiott, A. (2027). Holding paradox: A navigational approach to mind and consciousness & other gifts from the hippocampus. Iff Books, Simon & Schuster.
Hiott — Waymaking: a nested approach to cognition inspired by cognitive and computational hippocampal modelshttps://scholar.google.com/scholar?q=Hiott+Waymaking+nested+approach+cognition+hippocampal+models
Hiott — Navigability: a common orientation for the study of cognitionhttps://scholar.google.com/scholar?q=Hiott+Navigability+common+orientation+study+of+cognition
Andrea Hiott — some of the work and research index (papers, books, interview archive) but needs major updating https://www.andreahiott.com/my-work-and-research
Gärdenfors (2000), Conceptual Spaces: The Geometry of Thought — the philosophy that predicted the neuroscience, then co-wrote itself into it (see Bellmund 2018 in section 04) https://scholar.google.com/scholar?q=Gardenfors+Conceptual+Spaces+The+Geometry+of+Thought
Tolman (1948) again — end where it began https://scholar.google.com/scholar?q=Tolman+1948+Cognitive+maps+in+rats+and+men
Some conversations about some of this
Hippocampus Love — full video playlist. The single best link here; start with this rather than any one episode. https://www.youtube.com/playlist?list=PLV2AfcoqBukwqHsSTZVBBe0rBc5ZzpVWB
Lynn Nadel, part 1 — Among the Superheroes. McGill, the invasion of Prague, UCL, and the work that became the cognitive map. Show notes link to Penfield, Milner, Hebb, H.M., O'Keefe's Nobel lecture — a bibliography in itself. https://lovephilosophy.substack.com/p/among-the-superheroes
Lynn Nadel — the representation debate. Pairs directly with section 11: whether the hippocampus represents anything at all. Published alongside her Topoi paper. https://loveandphilosophy.com/beyond-dichotomy-podcast/lynn-nadel-representations
Same conversation, audio + slides https://www.buzzsprout.com/2229994/episodes/17920060-hippocampus-love-the-neural-representation-debate-cognitive-maps-with-lynn-nadel-bonus-episode
Brad Love — BrainGPT, models, and the hippocampus. Where sections 03–04 meet the question of what neuroscience is actually accumulating. https://lovephilosophy.substack.com/p/braingpt-neuroscience-and-loving-ce9
Start here — the navigational approach in brief. Fastest way in for a cold visitor. https://lovephilosophy.substack.com/p/quick-intro-navigational-approach
Navigational Mind — roots and branches. The genealogy, with the caveat built in: all navigability is partial, because a map is made from a position and for a reason. https://lovephilosophy.substack.com/p/navigational-mind-a-look-into-the
Full episode feed: https://podbay.fm/p/love-and-philosophy Apple: https://podcasts.apple.com/my/podcast/love-philosophy/id1701577110
Cognition in Motion, with Alexander Beiner — her being questioned rather than questioning https://lovephilosophy.substack.com/p/cognition-in-motion-with-andrea-hiott
Love, Phenomenology and Waymaking (Intelligent Teams) — shorter, non-specialist framing https://intelligentteams.substack.com/p/love-phenomenology-and-waymaking
Great video about Tolman and latent learning and some distinctions between learning and behaving in ways that do not require learning (like because the body is tired or hungry): https://youtu.be/jiyreiHp56g?si=ll7xL8uSeH6E-ZrI
Tim Behrens on cognitive maps — Brain Inspired. Section 03 from the source. https://player.fm/series/brain-inspired/bi-024-tim-behrens-cognitive-maps
Kim Stachenfeld — Brain Inspired. Co-author on the 2018 review; the predictive-map account. https://braininspired.co/podcast/193/
Brain Inspired full archive — 200+ episodes; more guests from sections 03–05 in there https://braininspired.co/ (Brain Inspired in general is a great resource if you want to go into the weeds of neuroscience but also in a way that can be doable without a degree in this stuff)
https://brainsciencepodcast.com/ Brain Science Podcast with Ginger Campbell is also awesome and an easier slide into these subjects.
The Mosers — lectures and interviews https://www.youtube.com/results?search_query=May-Britt+Moser+OR+Edvard+Moser+lecture+grid+cells
Demis Hassabis on the memory/imagination work — first author on the 2007 paper in section 07, before DeepMind. The useful interviews are the ones about the hippocampus, not about AI. https://www.youtube.com/results?search_query=Demis+Hassabis+interview+memory+imagination+hippocampus
Daniel Schacter lectures on constructive memory — the other half of section 07 https://www.youtube.com/results?search_query=Daniel+Schacter+lecture+constructive+memory+imagining+future
1. Foundational constructs (classic cognitive science & theory)
These define the ideas the neuroscience later tried to localize. Mostly books/older papers; could not always find links.
Bartlett, F. C. (1932).Remembering: A Study in Experimental and Social Psychology. Cambridge University Press. — Introduced "schema" as an active, reconstructive structure: memory is rebuilt to fit expectations rather than replayed verbatim.
Piaget, J. (1926/1952).The Origins of Intelligence in Children. — Schemas as assimilation/accommodation structures through which knowledge is built and updated during development.
Schank, R. C., & Abelson, R. P. (1977).Scripts, Plans, Goals and Understanding. Lawrence Erlbaum. — The origin of "script": stereotyped event sequences (the famous restaurant script) used to understand language and predict how situations unfold.
Miller, G. A. (1956). "The Magical Number Seven, Plus or Minus Two." Psychological Review 63:81–97. — The original "chunking" idea: we compress information into a handful of packaged units to beat capacity limits. Later generalized to action.
Tolman, E. C. (1948). "Cognitive maps in rats and men." Psychological Review 55:189–208. https://doi.org/10.1037/h0061626 — Proposed that animals build internal "maps" of relationships, not just stimulus–response chains — the seed of the modern cognitive-map program.
O'Keefe, J., & Nadel, L. (1978).The Hippocampus as a Cognitive Map. Clarendon Press. — Tied Tolman's map to the hippocampus after the discovery of place cells; the anatomical anchor for everything downstream.
Rumelhart, D. E., Hinton, G. E., & Williams, R. J. (1986). "Learning representations by back-propagating errors." Nature 323:533–536. — The connectionist backdrop in which schemas emerge from distributed networks rather than living as discrete symbols.
2. Rodent studies — schemas and rapid systems consolidation
causal, cellular-level evidence.
Tse, D., et al. (Morris lab) (2007). "Schemas and memory consolidation." Science 316:76–82. https://www.science.org/doi/abs/10.1126/science.1135935 — Landmark study: once rats had a flavor–place schema, brand-new pairings learned in a single trial became neocortical and hippocampus-independent within ~48 hours, showing schemas can hugely accelerate consolidation. (Follow-up: Tse et al., 2011, Science showed matching immediate-early-gene activity in medial prefrontal cortex.)
McKenzie, S., et al. (Eichenbaum lab) (2014). "Hippocampal Representation of Related and Opposing Memories Develop within Distinct, Hierarchically Organized Neural Schemas." Neuron 83:202–215. https://www.cell.com/neuron/fulltext/S0896-6273(14)00405-X — Hippocampal ensembles organize related events hierarchically by shared dimensions and keep opposing contexts separate — a neural instantiation of a relational schema. (See also the McKenzie & Eichenbaum, 2011 review.)
McKenzie, S., et al. (2013). "Learning Causes Reorganization of Neuronal Firing Patterns to Represent Related Experiences within a Hippocampal Schema." Journal of Neuroscience 33:10243–10256. https://www.jneurosci.org/content/33/25/10243 — New goal memories are first assimilated into existing goal-related firing patterns, then gradually differentiate — consolidation as assimilation-then-separation.
Zhou, J., et al. (Schoenbaum lab) (2019). "Complementary task structure representations in hippocampus and orbitofrontal cortex during an odor sequence task." Current Biology 29:3402–3409. https://doi.org/10.1016/j.cub.2019.08.040 — Hippocampus and OFC carry complementary representations of abstract task structure (schema), extending schema coding beyond space.
Wang, S.-H., & Morris, R. G. M. (2010). "Hippocampal–neocortical interactions in memory formation, consolidation, and reconsolidation." Annual Review of Psychology 61:49–79. https://www.researchgate.net/publication/26645633 — Authoritative review situating the schema/rapid-consolidation findings within hippocampal–neocortical dialogue.
3. Human neuroimaging — schemas, congruency, and event scripts
van Kesteren, M. T. R., Ruiter, D. J., Fernández, G., & Henson, R. N. (2012). "How schema and novelty augment memory formation." Trends in Neurosciences 35:211–219. — The SLIMM model (Schema-Linked Interactions between Medial prefrontal and Medial temporal regions): mPFC detects congruency with existing schemas and modulates hippocampal encoding accordingly. (Locate by title/DOI 10.1016/j.tins.2012.02.001.)
Baldassano, C., Hasson, U., & Norman, K. A. (2018). "Representation of Real-World Event Schemas during Narrative Perception." Journal of Neuroscience 38:9689–9699. https://www.jneurosci.org/content/38/45/9689 — The key "script" imaging study: mPFC, posterior medial cortex, and superior frontal gyrus carried restaurant-vs-airport script patterns that generalized across stories and modalities; mPFC uniquely tracked event order.
Baldassano, C., et al. (2017). "Discovering Event Structure in Continuous Narrative Perception and Memory." Neuron 95:709–721. https://doi.org/10.1016/j.neuron.2017.06.041 — Introduced the Hidden Markov Model method for finding event boundaries in continuous experience — the methodological backbone of the script work.
Masís-Obando, R., Norman, K. A., & Baldassano, C. (2022). "Schema representations in distinct brain networks support narrative memory during encoding and retrieval." eLife 11:e70445. https://elifesciences.org/articles/70445 — Dissociated story-specific vs. general schematic representations across cortical and hippocampal regions during memory, not just perception.
Song, H., et al. (2023). "Top-down attention shifts behavioral and neural event boundaries in narratives with overlapping event scripts." bioRxiv.https://www.biorxiv.org/content/10.1101/2023.08.08.552465 — Tackles the realistic case where multiple scripts overlap (e.g., a birthday at a restaurant), showing attention reshapes where event boundaries fall.
Greve, A., Cooper, E., Tibon, R., & Henson, R. N. (2019). "Knowledge Is Power: Prior Knowledge Aids Memory for Both Congruent and Incongruent Events, but in Different Ways." Journal of Experimental Psychology: General.https://pmc.ncbi.nlm.nih.gov/articles/PMC6390882/ — Confirmed SLIMM's predicted U-shaped memory curve: both highly schema-congruent and highly incongruent events are remembered better than neutral ones, via different mechanisms.
Sommer, T., et al. (2022). "The Assimilation of Novel Information into Schemata and Its Efficient Consolidation." Journal of Neuroscience 42:5916. https://www.jneurosci.org/content/42/30/5916 — Human evidence on vmPFC–hippocampal coupling when new facts can be related to prior knowledge, testing competing schema-consolidation accounts.
Study on self-generated learning (2024). "Recognizing ideas generated in a creative task: the roles of the hippocampus and medial prefrontal cortex." Cerebral Cortex 34:bhae219. https://academic.oup.com/cercor/article/34/5/bhae219/7682115 — Applies the schema/SLIMM framework to how prior knowledge scaffolds novel idea generation.
Post-encoding connectivity & durable memory (2023). "Effects of schema on the relationship between post-encoding brain connectivity and subsequent durable memory." https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10229577/ — Shows schema-consistency reconfigures which post-encoding networks (vmPFC–hippocampal vs. sensory–vmPFC) predict lasting memory.
Spatial schemas & one-shot navigation (2018). "Learned Spatial Schemas and Prospective Hippocampal Activity Support Navigation After One-Shot Learning." Frontiers in Human Neuroscience 12:486. https://www.frontiersin.org/articles/10.3389/fnhum.2018.00486/full — Human analogue of Tse: a learned spatial schema supports rapid, one-shot integration of new locations with reduced hippocampal dependence.
4. Theoretical frameworks and reviews
Ghosh, V. E., & Gilboa, A. (2014). "What is a memory schema? A historical perspective on current neuroscience literature." Neuropsychologia 53:104–114. https://www.sciencedirect.com/science/article/abs/pii/S0028393213003990 — Attempts to give "schema" a rigorous, componential definition, cutting through inconsistent usage across the field.
Gilboa, A., & Marlatte, H. (2017). "Neurobiology of Schemas and Schema-Mediated Memory." Trends in Cognitive Sciences 21:618–631. https://www.sciencedirect.com/science/article/abs/pii/S1364661317300864 — The major synthesis; positions vmPFC, hippocampus, angular gyrus and posterior cortex as the schema network and details their interactions.
Sekeres, M. J., Winocur, G., & Moscovitch, M. (2017/2018). "Details, gist and schema: hippocampal–neocortical interactions underlying recent and remote episodic and spatial memory." Current Opinion in Behavioral Sciences / related Trace Transformation work. https://www.sciencedirect.com/science/article/abs/pii/S2352154616302777 — Trace Transformation Theory: memories shift from detailed to gist-like/schematic over time, with the entorhinal cortex bridging anterior hippocampus and mPFC.
Kesner, R. P., & Rolls, E. T. / "Revisiting a unified theory" (2018). "Coordinating what we've learned about memory consolidation: Revisiting a unified theory." Neuroscience & Biobehavioral Reviews.https://www.sciencedirect.com/science/article/abs/pii/S0149763418302021 — Integrates schema findings (Tse) and engram findings (Kitamura/Tonegawa) into a unified account of cellular + systems consolidation.
Preston, A. R., & Eichenbaum, H. (2013). "Interplay of hippocampus and prefrontal cortex in memory." Current Biology 23:R764–R773. https://doi.org/10.1016/j.cub.2013.05.041 — Widely cited framework for how hippocampus and PFC divide labor in encoding, schema use, and retrieval.
McClelland, McNaughton & O'Reilly (1995) / Kumaran, Hassabis & McClelland (2016). Complementary Learning Systems theory. Psychological Review 102:419–457; Trends in Cognitive Sciences 20:512–534. — The backdrop: a fast hippocampal learner and a slow neocortical generalizer must be separate to avoid catastrophic interference — the reason schemas exist at all.
5. Entorhinal–hippocampal cognitive maps & structural knowledge
most specific to the hippocampal–entorhinal area — schema as structural code in a spatial-like format.
Constantinescu, A. O., O'Reilly, J. X., & Behrens, T. E. J. (2016). "Organizing conceptual knowledge in humans with a gridlike code." Science 352:1464–1468. https://pmc.ncbi.nlm.nih.gov/articles/PMC5248972/ — When people navigated an abstract 2D concept space, entorhinal cortex and vmPFC showed the same hexagonal grid signal as in spatial navigation — evidence that grid codes organize non-spatial knowledge.
Whittington, J. C. R., et al. (Behrens lab) (2020). "The Tolman-Eichenbaum Machine: Unifying Space and Relational Memory through Generalization in the Hippocampal Formation." Cell 183:1249–1263. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC7707106/ — A model in which medial entorhinal cells encode reusable structure and hippocampal cells bind it to sensory content, reproducing grid/place cells and generalizing schemas across environments.
Behrens, T. E. J., et al. (2018). "What Is a Cognitive Map? Organizing Knowledge for Flexible Behavior." Neuron 100:490–509. https://www.researchgate.net/publication/328509715 — The conceptual manifesto arguing the hippocampal–entorhinal map encodes relational structure of any domain, enabling inference and generalization.
Bellmund, J. L. S., Gärdenfors, P., Moser, E. I., & Doeller, C. F. (2018). "Navigating cognition: Spatial codes for human thinking." Science 362:eaat6766. https://doi.org/10.1126/science.aat6766 — Proposes that hippocampal–entorhinal geometric codes map "cognitive spaces," letting us reason about concepts as if navigating them.
Peer, M., Brunec, I. K., Newcombe, N. S., & Epstein, R. A. (2021). "Structuring Knowledge with Cognitive Maps and Cognitive Graphs." Trends in Cognitive Sciences 25:37–54. https://www.cell.com/trends/cognitive-sciences/abstract/S1364-6613(20)30250-3 — Distinguishes continuous "map" codes from graph-like relational codes as two formats the brain uses to structure knowledge.
Hernández-Frausto, M., et al. (2024). "Entorhinal cortex–hippocampal circuit connectivity in health and disease." Frontiers in Human Neuroscience 18:1448791. https://www.frontiersin.org/journals/human-neuroscience/articles/10.3389/fnhum.2024.1448791/full — Up-to-date review of the EC–hippocampal circuitry (including its breakdown in Alzheimer's) that underlies memory and map formation.
Bellmund, J. L. S., et al. "Mapping sequence structure in the human lateral entorhinal cortex." bioRxiv.https://www.biorxiv.org/content/10.1101/458133 — Human lateral entorhinal cortex represents the temporal/sequence structure of events, extending map coding into time.
Working-memory & the EC–hippocampal circuit. "Successful working memory linked to theta connectivity patterns in the hippocampal-entorhinal circuit." bioRxiv.https://www.biorxiv.org/content/10.1101/2022.09.08.507081 — Theta-band EC–hippocampal coupling supports holding information online, linking the circuit to active maintenance.
Eichenbaum, H. (2017). "On the Integration of Space, Time, and Memory." Neuron 95:1007–1018. https://www.sciencedirect.com/science/article/pii/S0896627317305603 — Argues the hippocampus maps memories across space, time, and abstract relations via one generalized organizing mechanism.
6. Automatization — habits, model-based vs. model-free control, action chunking
The procedural face of the same efficiency principle.
Daw, N. D., Niv, Y., & Dayan, P. (2005). "Uncertainty-based competition between prefrontal and dorsolateral striatal systems for behavioral control." Nature Neuroscience 8:1704–1711. https://www.nature.com/articles/nn1560 — The formal bridge: a flexible model-based (prefrontal) controller and a cheap model-free (dorsolateral striatal) controller, arbitrated by which is more accurate given uncertainty. Model-based control ≈ using a cognitive map.
Dolan, R. J., & Dayan, P. (2013). "Goals and habits in the brain." Neuron 80:312–325. https://doi.org/10.1016/j.neuron.2013.09.007 — Definitive review of the goal-directed/habitual (model-based/model-free) distinction and the classic "driving to your old house" illustration of automatized control.
A tutorial-style primer (2020). "Model-based decision making and model-free learning." Current Biology 30:R860–R865. https://www.sciencedirect.com/science/article/pii/S0960982220309039 — Accessible walkthrough of outcome-devaluation logic and the dorsomedial-vs-dorsolateral striatal dissociation.
"Model-based learning protects against forming habits" (2015).Cognitive, Affective, & Behavioral Neuroscience 15:523–536. https://link.springer.com/article/10.3758/s13415-015-0347-6 — In humans, stronger model-based control is associated with reduced slipping into rigid stimulus–response habits.
Cortico-striatal compartments model (2023). "Distinct cortico-striatal compartments drive competition between adaptive and automatized behavior." https://www.ncbi.nlm.nih.gov/pmc/articles/PMC10030038/ — Computational model of how dorsomedial (goal-directed) and dorsolateral (habitual) striatum compete, and how weakened prefrontal control tips behavior toward rigidity.
Model-based control in psychiatry (2017). "Model-Based Control in Dimensional Psychiatry." Biological Psychiatry.https://www.sciencedirect.com/science/article/abs/pii/S0006322317314816 — Frames compulsion across disorders as an imbalance favoring model-free/habitual over model-based control — the pathological edge of automatization.
Parkinson's & habit control (2021). "Impaired Formation and Expression of Goal-Directed and Habitual Control in Parkinson's Disease." https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8574955/ — Dopamine loss in sensorimotor striatum disrupts the normal goal-directed↔habitual balance, linking the framework to disease.
Graybiel, A. M. (1998). "The basal ganglia and chunking of action repertoires." Neurobiology of Learning and Memory 70:119–136. https://pubmed.ncbi.nlm.nih.gov/9753592/ — Proposes the striatum "chunks" action sequences into performance units, explicitly generalizing Miller's information-chunking idea to action control.
Smith, K. S., & Graybiel, A. M. — "task bracketing." QnAs / review: https://www.pnas.org/doi/10.1073/pnas.1315012110 — As behavior becomes habitual, sensorimotor striatal neurons fire mainly at the start and end of a routine, "bracketing" it as one packaged chunk.
Graybiel, A. M., & Grafton, S. T. (2015). "The Striatum: Where Skills and Habits Meet." Cold Spring Harbor Perspectives in Biology.https://pmc.ncbi.nlm.nih.gov/articles/PMC4526748/ — Reviews how skills and habits share striatal machinery, including the intrastriatal reorganization behind task-bracketing.
Martiros, N., Burgess, A. A., & Graybiel, A. M. (2018). "Inversely Active Striatal Projection Neurons and Interneurons Selectively Delimit Useful Behavioral Sequences." Current Biology. Explainer: https://mcgovern.mit.edu/2018/02/08/distinctive-brain-pattern-helps-habits-form/ — Identified specific striatal neurons that mark the beginning and end of chunked, learned routines.
Sensory compression & chunking (2025). "Sensory Compression as a Unifying Principle for Action Chunking and Time Coding in the Brain." bioRxiv.https://www.biorxiv.org/content/10.1101/2025.09.05.674585 — Recent proposal that compression links stereotyped action chunks and elapsed-time coding in the same dorsolateral-striatal circuits.
7. Perception and prediction — "how we see and move through the world"
The perceptual face: schemas as priors in a predictive brain.
Bar, M. (2007). "The proactive brain: using analogies and associations to generate predictions." Trends in Cognitive Sciences 11:280–289. https://doi.org/10.1016/j.tics.2007.05.005 — Argues the brain is fundamentally predictive: it uses stored associations/schemas to anticipate incoming input rather than passively receiving it.
Clark, A. (2013). "Whatever next? Predictive brains, situated agents, and the future of cognitive science." Behavioral and Brain Sciences 36:181–204. (DOI 10.1017/S0140525X12000477.) — The influential statement of predictive processing: perception is the brain testing top-down predictions (priors/schemas) against sensory evidence, mostly processing the mismatch.
Friston, K. (2010). "The free-energy principle: a unified brain theory?" Nature Reviews Neuroscience 11:127–138. (DOI 10.1038/nrn2787.) — The mathematical parent framework (active inference): both perception and action work to minimize prediction error — action makes predicted sensory states come true, which is one way to understand automatized, skilled movement.
Schemas/scripts (Themes 1–5) and habits/automatization (Themes 6–7) are ways we assess how the brain extracts regularities, caches them as reusable structure, and runs that structure automatically to avoid recomputing from scratch.
Intro Texts From Class and Misc:
Intro to Affordances: https://ixdf.org/literature/book/the-encyclopedia-of-human-computer-interaction-2nd-ed/affordances#423_theoretical_roots-2
AFFORDANCE and Design by Don Norman: https://jnd.org/affordances-and-design/
J.J. Gibson’s The Theory of Affordances (the original text)
Ch 8 of Ecological Approach to Visual Perception https://monoskop.org/images/c/c6/Gibson_James_J_1977_1979_The_Theory_of_Affordances.pdf
Even more clarification if you really want it
https://graphicsinterface.org/wp-content/uploads/gi2000-24.pdf
https://media.pluto.psy.uconn.edu/MC.pdf
A very cool and influential but maybe not so easy paper from some friends in Amsterdam about Landscape and Affordances
Hidden Brain episode about mental mapping (Thanks Amanda)
Thought Process by JR in New YorkerDiaries, Talks, Creations: Here is playlist of messy attempts and conversations and experimenting towards this approach and articulating it across the usual bounds
Here are about 90 more talks on the subjects here form different philosophical POVs.