Spatial Navigation: The Art and Science of Wayfinding

On the 5th of May, Dora -a writer and geographer interested in creative digital mapping, and Amy Young -an artsy neuroscientist fascinated by neural correlates of space and place, hosted an online workshop exploring the neuroscience of spatial navigation. Their interactive discussion featured our cellular toolkit, the systems in place that encode our wayfinding ability, how words, emotions and memories anchor our inner experience to places, how different spatial habits have developed alongside cultural identities, and what cognitive changes can occur when we map more ‘playfully.’ 

Next, Dora walked us through her collaborative, geospatial poetry map It uses What3Words, a mapping software that generates 3 word codes for 3x3m grid squares that would ordinarily be 16 digit coordinate numbers. Those are as impossible to remember as they are to communicate! Poems on Dora’s website use the 3 word codes as a departure point for sharing individual inner experiences of place as well as geolocating them onto an interactive map. In order to tag the poems to the map, they must include somewhere the 3 words that describe their coordinates. This becomes a creative challenge, and was especially so due to the time-limited workshop. Moreover, given the new significance that 3m distances has in our lives during today, the Waywriting map is especially poignant for sharing our experiences of a deeper connection to our landscapes and exploring other’s. She is accepting submissions! 

We were also delighted to be joined by Susan from feelSpace, a group who developed a “vibrating compass belt” originally from the neurobiopsychological workgroup of the University of Osnabrück. feelSpace is an “artificial sensory organ”, continuously indicating north to its wearer. The participants who wore the belt for an extended period of time were excited by the experience of experiencing the north for a few weeks, and the results were very striking. The belt is not only applicable in research but is also truly useful in everyday life. Susan recounted the truly profound effects it had on her and her sense of place, even after it was removed.

Thank you to all who joined us for this fantastic online lecture and workshop. Below you will find a more detailed review of the research that went into the talk.


the workshop

spatial navigation techniques

As participants entered the Zoom meeting, they were invited to  draw maps of their last outing (figure 1). Many people are in lockdown around the world, so imagining the last limited trip held some poignancy. This kicked off a discussion of different spatial coding techniques in our brains, which reflect how we position ourselves in relation to our surroundings and navigate them. Those who visualised themselves walking through the space whilst drawing demonstrated an egocentric navigation technique, which is a self-oriented point of view, and relies on visual cues, much like the street view version of Google maps. This type of navigation ability is characterized by activity in the caudate nucleus (Bohbot et al, 2004, Hartley et al, 2004) and is the most common method of navigation.

Figure 1: a map drawn by Tatiana Lupashina during the workshop. Tatiana is in lockdown in Montenegro and depicted her most recent outing to the beach.

However, those who pictured a top-down map of their environment when recalling their journey, picturing all the features along the way in relation to each other, exhibited allocentric spatial coding, which is mostly done in the hippocampus. There is no correct method – the reality is, we use these methods in parallel (figure 2). However, the processes that form the basis of our cognitive maps occur largely in the hippocampus (Bohbot et al, 2014).

Figure 2: Allocentric vs egocentric navigation. Image credit Kozhevnikov lab, University of Singapore

One particularly difficult method of egocentric navigation is path integration, defined as the capacity to use cues generated by an animals’ movements to calculate the updated position by monitoring trajectory in relation to a start location (Gallistel, 1990, Whishaw and Wallace, 2003). Participants in the workshop were invited to test their path integration abilities using a simple method from Science Direct.

Figure 3: Nainoa Thompson. Polynesian Master Navigator. Image credit: Hilaire Picault, 2017.

Some Polynesian sailors show an incredible capacity for this ‘dead reckoning’; master navigator Nainoa Thompson (figure 3) has sailed double-hulled, traditional voyaging canoes around Micronesia, Polynesia, Canada and the US, all using this ancient wayfinding technique, as well as celestial navigation. His advice, ‘just don’t forget, that’s not an option. Forgetting means you’re lost.’ This indicates how memory-building is to being a good navigator. Constructing scenes in the mind is crucial for recalling the past, imagining the future and getting from one to the other.  Luckily, we have a spatial navigation toolkit which enables us to do this. It contains a list of players that is by no means complete as new discoveries are always being made. In this workshop we touched on just a few of its stars, starting with a discovery that kicked it off.

the toolkit

place cells

One of the reasons that place and memory are so tied is because of these cells. Discovered by John O’Keefe and Jonathan Dostrovsky in 1971 in the rat hippocampus, these neurons have since been found in many more mammals including humans. When an animal freely moves in space a single place cell increases its action potential/ firing probability in response to a place field. When an animal moves to another area altogether, the cells get remapped and the same cell might be used to code a spot in this new area.

This forms an important component of our spatial cognition, but what’s more, these cells are thought to be key in episodic memory. Most memories took place somewhere, and although we don’t know exactly how memories are coded- it’s not like-for-like – we do know that place cells have a role in episodic memory because it’s hard to remember an event without the place it happened in. This might remind some of the “memory palace” or loci method, the mnemonic technique which has been used for centuries. It’s easier to remember things if you visualize them somewhere in space.

head direction cells

Head direction cells (HDC) are nerve cells that increase their firing rates above baseline levels when an animal’s head points in a specific direction (figure 4). They are found in many mammals (first rats), and supposed in all, although it’s quite hard to test in humans. They are found in many locations in the brain, although it is thought that the cortical head direction cells process information about the environment, while the subcortical (hippocampal) ones process information about angular head movements.

Figure 4: Head direction cell tuning-curve polar plots for recorded neurons. Image credit: Page et al, 2018.

Although based on vestibular system and internal inertia, HDC firing is not only determined by sensory features. When an animal first arrives at a new location, the alignment of HDC/system seems arbitrary. Over the first few minutes of exploration, the animal learns to associate the landmarks in the environment with directions. When the animal comes back into the same environment at a later time, if the head direction system is misaligned, the learned associations serve to realign it.

boundary vector cells

O’Keefe & Burgess found if you doubled the size of the room the rat was moving in, the place cell still fired in that location. There was correspondence. The existence of boundary vector cells (BVC) were predicted from the place cell experiments, and extensive models were made about where they should be and what they look like (figure 5). So in what one can only imagine as an incredibly satisfying moment boundary vector cells were discovered in the rat subiculum by Colin Lever in 2009. They are found in the hippocampal units: subiculum, presubiculum and entorhinal cortex (in the MEC, BVC comprise 10% cell population). BVCs tell the place cells to reset, that a threshold has been crossed. They are very strong anchoring cues. Translated into behaviour, perhaps it is no coincidence that when people are lost in the forest they are often found along borders to things such as fences or forests.

Figure 5: The BVC model. A BVC responds maximally when a boundary is perceived at a preferred distance and allocentric direction from the animal, regardless of the animal’s heading direction. A, The receptive field of a BVC tuned to respond to a barrier at a short distance east-northeast from the animal. B, BVCs tuned to respond to barriers farther from the animal will have broader receptive fields. C, The firing field (firing rate as a function of the animal’s location; top) for a BVC with a receptive field tuned to respond to a boundary at a short distance to the east (bottom). D, Predicted firing fields in different environments for the BVC shown in C. Insertion of a barrier causes a doubling of the field (bottom right panel). Image credit: Lever et al, 2009.

Boundary cells also strongly interact with our memory.  This is well illustrated by a study by Adrian Horner called Walking Through Doors, participants were asked to go through VR doors and remember two objects. It was easier to remember two objects in the same room, than those either side of a door. The place has been remapped. Spatial boundaries represent event boundaries. Our continuous life experience gets broken up in our memory, in theory partially because of crossing boundaries.

grid cells

The story was not yet complete; place, internal direction, and borders were beginning to be understood, but how was distance coded? As the medial entorhinal cortex (MEC) is the largest input to the hippocampus, it was hypothesized to be of relevance. Single cell recording in the rat MEC showed this to be the case as Edvard & May-Britt Moser discovered in 2005.

Figure 6: A) Nissl stained rat MEC. B) rhythmic triangular grid firing patterns of grid cells in the MEC. Image credit: Hafting et al, 2005

Grid cells store information about location, distance, and direction by rhythmically firing in a triangular grid pattern, hence the name (figure 6). There are many layers of grid cells, the top layers firing nodes are at about 30cm distance, the bottom layers are about 10m. Which seems excessive until you consider behaviours that only require low resolution understanding about the place one is in, such as fleeing. The brain is exceptionally efficient.

Figure 7: Mosers and John O’Keefe won the nobel prize in 2014 for their outstanding contributions to understanding our navigation. In a wonderful moment for art and science May-Britt wore a dress to the ceremony embroidered with grid cells. Designed and created by fashion house Matthew Hubble.

HDC are also in the MEC as will as grid cells. During development HDC and BVC show adult-like firing fields (day 16- 1yr humans). HDC and BVC are important for initial learning of the environment, but not grid cells. Is this why we are so bad at judging distance when we are young?

Figure 8:  Image credit: 2014 nobel prize committee.

How do these systems tie together? One of the ways hypothesized that these cells and regions communicate with each other is by synchronizing the cell population firing to certain rhythmic frequencies. Populations of hippocampal place cells have been found to fire, as in let off their electrical action potentials at theta frequency (6–9 Hz) which is thought to be responsible for some of the coding of place. As the animal advances to the spot where the place cell fires most rapidly, it’s firing coincides with an earlier and earlier point on the population theta wave firing.  This is called theta phase precession (O’Keefe & Reece, 1993).

The interaction between the hippocampus and MEC of rats and humans are shown in figure 8. We owe a lot to the model animals scientists have learnt from over the years as recording a single cell’s activities is exceptionally difficult, and is not experimentally feasible in humans. Research shows for example, that there is a cognitive replay, or consolidation, of the cells firing patterns forged in the day, again during sleep at 10-20x the original speed (Buzsáki, 1998). This strengthens the connection, and cements the learning experience. This “replay” also happens before an animal makes a decision/ does an activity (Roux et al, 2017) and is called “cognitive preplay”. It is thought to play a role in decision making.

putting our tools to use

Looking at our spatial navigation toolkit, it is easy to see why some cities are so disorientating. The spatial neurons discussed above rely on paths, edges, nodes and landmarks. Our encoding processes become confused by their repetitive features, sheer scale, and complex hidden transport systems. It is often beyond our ability to imagine the city in full, so we can’t position ourselves in space, and it is therefore ‘Illegible’ (Lynch, 1960). Environmental psychologist Negin Minaei investigated if these tools further impede our ability to make cognitive maps. She found that Londoners who used these apps were terrible at drawing their city from memory (2014). London is a particularly vicious example, more people get lost there than any other metropole (Nokia, 2008). Efforts to understand London are further obstructed by the ‘Goldilocks’ tube map that plays to our efficient – lazy – brains, giving us just enough information to get around, but straightening lines and distorting distances so that it doesn’t actually represent the city. 

As such, we depend on web-based directional tools like Google Maps and Citymapper. However, by not making spatial decisions for ourselves, we aren’t building a cognitive map efficiently. We aren’t looking around and using the head direction cells that orient us. Further, the speed cells that fire in theta oscillations in our entorhinal cortex are activated by our bodily movement. That means, when we ride the tube, these oscillations drop off and no hexagonal grid patterns are formed in our grid cells, meaning that we aren’t navigating the space (Winter et al, 2015). 

We investigated if this is actually a problem. After all, what’s so wrong with being a passenger when we can still get from A to B? Even healthy adults tend to use automated egocentric response of the caudate nucleus, rather than more active allocentric spatial technique. Is that because we lose grey matter as we get older so favour the hippocampal method less, or because we are using it less, grey matter declines? We are not yet sure. Additionally, we cannot be certain that smartphones have a negative effect on our hippocampal grey matter as we are decades away from meaningful data on that topic. What we do know, however, is that the old adage goes, “use it or lose it”. And if losing it means letting hippocampal grey matter that could prove protective in cognitive impairment, waste away out of inactivity, we suggest erring on the side of “using”. 

We have so far looked at how nurture/culture/society affects people’s wayfinding abilities or appetites, yet there is much research into the genetic effects. One notable contender are the APOE gene variants. APOE is a protein involved in fat metabolism, and humans have 4 gene variants that make this protein. Veronique Bohbot at Mcgill University in Montreal found people with APOE 2 gene variant more likely to use allocentric (spatial) strategy. This is interesting because the APOE 2 gene variant is widely accepted to confer some sort of protection against mild cognitive impairment (MCI) and Alzheimer’s disease (AD), whereas variant 4 is meant to be a biomarker, a risk. Something like 12% of people with 2 copies of APOE 4 get AD, it’s the strongest gene biomarker for AD we know.

Bohbot says that ‘the future of our species depends on our ability to transcend our robot behaviour,’ meaning our lazy, efficient tendencies from the automated caudate nucleus. How can we become active navigators again?

creatively mapping to improve our wayfinding ability

One of our participants, Johnny, who grew up in London shared that he didn’t roam further than a mile from his home until he was 10. Now, he cycles across the city, discovering new routes daily. That adventure, freedom and confidence improve human cognitive mapping abilities is ubiquitous. When Stanely Milgram asked Parisians to draw maps of the city, 40 years before Minaei asked Londoners, they drew maps ‘rich in symbolic imagery,’ that were ‘usually pretty accurate’ and all included secret places, indicating that their saw their city as ‘rich, variegated and inexhaustible in its offerings,’ and could hold onto this magic more clearly in their mind’s eye (1992:88). 

Re-enchanting the city has been, historically, a subversive practice, as the ‘disorientation maps’ of the Situationist International and Debord’s theory of the ‘Derive’ demonstrate (1958). Dropping the usual motivations for movement, letting oneself be drawn more playfully by the immediate attractions of the terrain constitutes ‘psychogeography.’ Many contemporary, digital counter-mapping apps can facilitate this. Representing these experiences on new, relational maps of the city uncover hidden narratives, routes, networks and perceptions (figure 9). 

Figure 9: Contemporary interpretation of Situationist International map. Image credit: Ailish Walker, Dublin, 2014.

Creative, personalised maps can be useful for planners in creating inclusive, user-friendly, and safer cities. Minaei showed in a study in 2018 that people from different ethnic backgrounds understand, learn and recall cities differently. For example, Chinese participants saw a more colourful and visual city, and reflected this attitude in their maps. Urban designers should consider these differences to ensure multicultural and global cities are navigable to all people with different cognitive and map-learning abilities. Self-assurance and spatial memory are interconnected; a city that is readable for its diverse residence will mean that their populus find their place in space easier. As a side note, uncovering these hidden and personalised ways of seeing the city can make for more inclusive, user-friendly landscapes. Participatory GIS and or other digital techniques of mapping the city from the ground-up, like the SMARTSTREETS initiative from UCL’s Bartlett School of Planning, help to situate city planners and will be a crucial tool for the healthy, post-COVID city.

the place of words

Place-cells are so called because they let us attach meaning to spatial locations. Once a ‘space’ is imbued with meaning, it becomes a ‘place.’ This significance helps us recall it – as the memory palaces indicate – but we need to describe it too. Imagine telling a story without the location it took place in? John O’Keeffe, who discovered place cells, also asserted that our spatial cognition systems are a deep structure for language. We are creatures of narrative, and words likely evolved because of our need to build memories and share our impressions of the spatial attributes of our surroundings, which would have been essential for survival. This related to the work of philosopher AJ Ayer, who introduced the idea that unless we can name something, we cannot conceive of it – there is a direct relationship between our vocabularies and our imaginations (1936). 

What makes members of Indigenous communities so good at spatial cognition is that they have developed what anthropologists call ‘memory-scapes’ – mental maps encompassing both physical and cultural worlds, in which environmental features are imbued with meaning to make them more legible. Toponyms exemplify these linguistic navigational aids. An etymological map of Welsh place-names demonstrates how these could have, historically, assisted survival by describing what was to be found in certain places. Claudio Aporta’s research with the Inuit communities of Igloolik in the Canadian Arctic also demonstrates their detailed linguistic inventory of visual cues. Everything is named, from ice features to ocean currents, drift patterns in the snow made by primary winds, and the term for a ‘good navigator’ is ‘aangaittuq,’ meaning ‘attentive’ (2016). Another example would be the Aboriginal Songlines: tracks of country extending thousands of miles are described in ancient songs that also tell the story of the land’s creation. These lines are mapped in the mind, embodied by one who walks them, and often painted in detail by memory from a bird’s-eye perspective. Figure 10 shows the waterholes and topographic lines of sand dunes in Balgo Territory.

Figure 9: Tjumpo Tjapanangka’s Kangaroo Dreaming, 2014.

Colonial forces have re-named toponyms and obstructed aural storytelling traditions as well as language continuity. Losing such  intergenerational vectors of communication can impede navigational skills with negative impacts on community wellbeing. And, in Western, post-exclusion societies, we have been moved off the land for so long that the ability to roam freely is beyond living memory, we are ‘lost’ because we are ‘illiterate in the language of the Earth itself,’ or simply don’t stop to read it anymore (Solnit, 2010:10). 

If, as we have seen, story-based knowledge facilitates navigational capacities, it follows that we should curate banks of personal, descriptive markers as reference points for exploring shared spaces. There are existing efforts to do so. For instance, INDIGITAL is an Aboriginal community VR app that allows users an immersive experience of the ancestral stories geolocated to landmarks they walk past. Many geospatial poetry anthologies also exist, such as the Places of Poetry map built by two poets from the University of Exeter last year, and the Disappearing in Australia. The Disappearing specifically tags work to ‘lost’ places, invisible histories and threatened non-human natures in rapidly developing urban places, using language of loss, land, culture and memory. Whilst seemingly bleak, it is important to recognise the similarities between this nostalgic, backward facing practice and the navigational technique of ‘dead reckoning’ mentioned earlier. Species and landscapes are swallowed up behind us by our so-called progress, as in thick sea fog. In order to navigate without losing sight of our desired destination (that is, the environmentally sound shared future we want), we have to note what we have left behind in order to find our current bearings, speed and direction, and reorientate ourselves accordingly. When it comes to memory, silence has dangerous implications. 

Dora has created a collaborative, geospatial poetry map, which uses What3Words, a mapping software that generates 3 word codes for these 3x3m grid squares instead of 16 digit coordinate numbers that are as impossible to remember as they are to communicate. In order to tag the poems to the map, they must include somewhere the three words that describe their coordinates. In this way, it becomes a creative challenge, though it is forgiving in that one can toggle around the 3m squares in your chosen location until three workable words are found. Given the new significance that 3m distances have in our lives today, the Waywriting map is especially poignant for sharing our experiences of a deeper connection to our landscapes and exploring other’s. 

Thank you to all who made it through the workshop and/or this text. It felt super special to be able to work together and share our passions, and thanks to you for the opportunity and interest. Feel free to contact us for any questions. Check out EDGE’s social media or sign up for our newsletter for future workshops on Creativity in Neuroscience.

By Dora and Amy Young




Aporta, C. 2016. ‘Markers in space and time: reflections on the nature of place names as events in the Inuit approach to the territory.’ In: Marking the Land: Hunter-gatherer creation of meaning in their environment. William Lois and Robert Whallon, eds. Chapter 4, London: Routledge. See also, Aporta’s PhD. thesis, ‘Chapter 5: Old Routes, New Trails: Contemporary Inuit Travel and Orienting in Igloolik, Nunavut,’ University of Alberta, 2003.

Ayer, A. J. 1936. Language, Truth and Logic. London: Gollancz.

Bohbot, V.D., Iaria, G. and Petrides, M., 2004. Hippocampal function and spatial memory: evidence from functional neuroimaging in healthy participants and performance of patients with medial temporal lobe resections. Neuropsychology, 18(3), p.418.

Bond, M. 2020. ‘Chapter 9: City Sense.’ In: Wayfinding: the Art and Science of How we Find and Lose Our Way. London: Picador, p192.

Buzsáki, G., 1998. Memory consolidation during sleep: a neurophysiological perspective. Journal of sleep research, 7(S1), pp.17-23.

Cornell, E. and Heth, C. D. 2006. ‘Home range and the development of children’s wayfinding,’ Advances in Child Development and Behaviour, 34, pp173-206.  

Debord, G. 1958. The Theory of the Derive. England: Atlantic Books (1997).

Hafting, T., Fyhn, M., Molden, S., Moser, M.B. and Moser, E.I., 2005. Microstructure of a spatial map in the entorhinal cortex. Nature, 436(7052), pp.801-806.

Hartley, T., Trinkler, I. and Burgess, N., 2004. Geometric determinants of human spatial memory. Cognition94(1), pp.39-75.

Lever, C., Burton, S., Jeewajee, A., O’Keefe, J. and Burgess, N., 2009. Boundary vector cells in the subiculum of the hippocampal formation. Journal of Neuroscience, 29(31), pp.9771-9777.

Lynch, K. 1960. The Image of the City. MIT Press, p4. 

MacFarlane, R. 2015. Landmarks. London: Hamish Hamilton, pp19-20.

Milgram, S. 1992. ‘Psychological Maps of Paris.’ The Individual in a Social World: Essays and Experiments, 2nd Ed, McGraw-Hill, p88.

Minaei, N. 2014. ‘Do modes of transportation and GPS affect cognitive maps of Londoners?’ Transportation Research Part A 70, pp162-180.

Minaei, N. 2018. ‘Inhabitants’ Cognitive Maps Represent Ethnic-Based Variations in Learning and Recalling London.’ Faculty of Engineering, University of Windsor, Canada, Papers Session 43 – Values, Norms and Beliefs, p1051. 

Nokia Press release, 2008. ‘Lost in the City.’ October. Available here:

O’Keeffe, J., Nadel, L. 1978. The Hippocampus as a Cognitive Map. England: OUP, pp391-410.

O’Keefe, J. and Recce, M.L., 1993. Phase relationship between hippocampal place units and the EEG theta rhythm. Hippocampus, 3(3), pp.317-330.

Page, H.J., Wilson, J.J. and Jeffery, K.J., 2018. A dual-axis rotation rule for updating the head direction cell reference frame during movement in three dimensions. Journal of neurophysiology, 119(1), pp.192-208.

Roux, L., Hu, B., Eichler, R., Stark, E. and Buzsáki, G., 2017. Sharp wave ripples during learning stabilize the hippocampal spatial map. Nature neuroscience, 20(6), p.845.

Sonit, R. 2009. A Field Guide to Getting Lost. England: Canongate, p.10.

Winter, S.S., Mehlman, M.L., Clark, B.J. and Taube, J.S., 2015. Passive transport disrupts grid signals in the parahippocampal cortex. Current Biology, 25(19), pp.2493-2502.

One thought on “Spatial Navigation: The Art and Science of Wayfinding”

  1. thanks a lot 🙂


    Would love to use it for our impro dance in public space but I just understand a little bit from the first to parts.. it would be so wonderful if through an easy language it would be accessible/ understanable for non-scientist but e.g. for me as researching dancer (who is a psychologist as well) and influences public spaces through dance and healing interventions <3







    Gesendet: Samstag, 30. Mai 2020 um 18:21 Uhr Von: "EDGE" <> An: Betreff: [New post] Spatial Navigation: The Art and Science of Wayfinding

    edge-neuroart posted: " On the 5th of May, Amy -an artsy neuroscientist fascinated by neural correlates of space and place, and Dora Young – a writer and geographer interested in creative digital mapping, hosted an online workshop exploring the neuroscience of spatial navig"

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