Part 2 - Cognitive Maps Introduction

Summary

The video script discusses the influential work of psychologist Edward Tolman, who introduced the concept of cognitive maps in the 1940s. It highlights his ideas on multiple scales of mapping, different types of learning, latent learning, and the importance of expectation. The speaker, Lucy, connects Tolman's theories to subsequent research, including the discovery of place cells by John O'Keefe, which led to a deeper understanding of spatial memory and the brain's predictive capabilities. The script also touches on the relationship between Tolman's scientific ideas and his political beliefs, emphasizing the ongoing relevance of his work in cognitive science.

Takeaways

• 🎓 Tolman's work on cognitive maps was foundational, coining the term and starting the field in the 1940s.
• 🌟 The concept of cognitive maps includes multiple scales, from narrow strip maps to broad comprehensive maps.
• 🧠 Tolman argued for different kinds of learning, each supporting different behavioral strategies.
• 🐭 Tolman demonstrated latent learning, showing animals can learn through curiosity and exploration without external rewards.
• 🔄 Tolman's ideas were ahead of their time, predicting modern concepts like the predictive brain and the role of the hippocampus in spatial memory.
• 📚 O'Keefe and Nadell's work built upon Tolman's, with the discovery of place cells being a key development.
• 🏆 Nadell's academic journey included influential figures like Donald Hebb and Brenda Milner, and significant contributions to neuroscience.
• 🏛️ The historical context of Tolman's work is highlighted, including his political activism and the influence of his environment on his scientific ideas.
• 🧭 The importance of exploration in map-making was emphasized, with the suggestion that the reward for an animal is knowledge of the environment.
• 🔮 Predictions made by O'Keefe and Nadell about the necessity of direction and distance information in cognitive maps were later confirmed by discoveries of head direction cells and grid cells.
• 🔄 The script reflects on the evolution of ideas from Tolman's initial concepts to current neuroscience research, showing a continuum of scientific thought.

Q & A

Who is the speaker being introduced in the script?

The speaker being introduced is Lynn Nadel.

What is the significance of the location where the event is taking place?

The event is taking place in Tolman Hall and the Faculty Club, which is significant because Tolman led a revolt on the campus, and it's a poignant location for discussing his work on cognitive maps.

What is the connection between Lynn Nadel's early life and his academic career?

Lynn Nadel grew up in New York City, did his degrees at McGill, and worked with notable figures like Donald Hebb and Brenda Milner. He then spent time in London, contributing to his academic development.

What was the main focus of Lynn Nadel's book-length manifesto on the hippocampus?

The main focus was that the hippocampus functions as a cognitive map, addressing two significant problems in Neuroscience: how the brain makes a map and what the hippocampus actually does.

What are the four key concepts that Lynn Nadel discussed from Tolman's work?

The four key concepts are: 1) Cognitive maps come in multiple scales, 2) There are different kinds of learning, 3) Latent learning can occur without reward, and 4) Animals have expectations and the brain acts as a predictor.

What was the place versus response controversy mentioned in the script?

The controversy was about whether there were different kinds of learning, such as place learning and response learning. Tolman argued for different kinds of learning supporting different behavioral strategies, while Frank Westell's solution suggested they weren't fundamentally different.

What was the significance of Tolman's latent learning experiments?

Tolman's latent learning experiments showed that animals could learn about their environment through curiosity and exploration without the need for external rewards, challenging the prevailing view that learning required reinforcement.

What was the 'Tinkle Paw' experiment mentioned in the script?

The 'Tinkle Paw' experiment demonstrated that animals have expectations. In the experiment, a monkey was trained to expect a reward (bananas), and when it was given lettuce instead, it showed clear signs of disappointment, indicating an expectation of bananas.

How did Lynn Nadel's work with O'Keefe build upon Tolman's ideas?

Nadel and O'Keefe built upon Tolman's ideas by developing a theory of cognitive maps that included the discovery of place cells by O'Keefe, and they emphasized the importance of exploration and the generation of expectations in animal behavior.

What predictions did Nadel and O'Keefe make in their book about the hippocampus?

They predicted that if the hippocampus was a cognitive map, it would need cells that conveyed information about direction and distance. This prediction was later supported by the discovery of head direction cells and grid cells.

What is the significance of the discovery of grid cells in relation to cognitive maps?

The discovery of grid cells, which fire in a hexagonal pattern across an environment, provided a mechanism for the hippocampus to create a spatial map, supporting the idea that the hippocampus functions as a cognitive map.

Outlines

00:00

🎓 Introduction to Tolman's Cognitive Map Legacy

The speaker begins by setting the context of the event, celebrating Tolman's contributions to cognitive mapping. They highlight Tolman's leadership during the nonsigners Faculty movement and the significance of the event's location. The introduction continues with a warm welcome for the first speaker, Lynn Nadel, emphasizing his academic journey and contributions to cognitive science. Nadel's work on memory development in Down syndrome and the Multiple Trace Theory of human memory consolidation are acknowledged. The speaker also touches upon the importance of curiosity and exploration in learning, as championed by Tolman, and the idea that cognitive maps can exist at different scales.

05:01

🧠 Tolman's Theories on Learning and Cognitive Maps

This paragraph delves into Tolman's theories on learning, emphasizing the distinction between different types of learning, such as place learning and response learning. It discusses how Frank Westell's solution to these different learning types was not entirely in line with Tolman's view, which suggested that different learning types have distinct properties and are supported by different brain systems. The paragraph also covers Tolman's concept of latent learning, which challenged the notion that reward is necessary for learning, instead arguing that curiosity and exploration can be sufficient motivators. Lastly, it touches on Tolman's ideas about expectation and how animals anticipate outcomes, which was a precursor to modern concepts of the predictive brain.

10:03

📚 Building on Tolman: O'Keefe and Nadel's Cognitive Map Book

The speaker discusses the foundational work of O'Keefe and Nadel on cognitive maps, which was heavily influenced by Tolman's ideas. They outline the key concepts from their book, including the role of cognitive maps as spatial systems, the existence of multiple memory systems supporting different behavioral strategies, and the importance of exploration in map formation. The speaker also highlights how cognitive maps enable flexible behavior by providing a rich information base. Additionally, they mention the predictive nature of the hippocampus and how it relates to Tolman's ideas about expectation and anticipation.

15:05

🚀 Advancing Cognitive Map Research

In the final paragraph, the speaker concludes their introduction by emphasizing the predictions made in O'Keefe and Nadel's book about the necessity of direction and distance information for cognitive maps to function effectively. They note the subsequent discovery of head direction cells and grid cells, which supported these predictions. The speaker then transitions the discussion to the next speaker, David Foster, who will delve into the role of the hippocampus in generating predictions, thus continuing the exploration of cognitive map research beyond the foundational work of Tolman.

Keywords

💡Cognitive Maps

Cognitive maps refer to the mental representations that individuals create of their environment, which help them navigate and understand spatial relationships. In the video, cognitive maps are central to the discussion on how the brain processes spatial information. The concept is rooted in Tolman's work, which suggested that these maps come in multiple scales, from narrow strip maps to broad comprehensive maps, indicating different levels of detail and complexity.

💡Place Learning

Place learning is a type of learning that involves understanding the environment based on spatial cues rather than response to specific stimuli. The video discusses Tolman's view that place learning is distinct from response learning, suggesting different kinds of learning that support different behavioral strategies. This concept is linked to the discovery of place cells in the brain, which are activated when an animal is in a specific location.

💡Response Learning

Response learning is a behavioral process where an organism learns to make specific responses to particular stimuli. In the video, it is contrasted with place learning. Tolman argued that there were different kinds of learning with different properties, and response learning was one of them, which could be dependent on different brain systems than place learning.

💡Latent Learning

Latent learning is the process by which learning occurs without an immediate observable outcome. The video mentions Tolman's experiments that demonstrated animals could learn about their environment in the absence of reward or external motivation, suggesting that curiosity and exploration are sufficient motivators for learning.

💡Hippocampus

The hippocampus is a region of the brain that plays a major role in memory and spatial navigation. In the video, it is highlighted as the area of the brain where cognitive maps are thought to be formed and where place cells are located. The hippocampus is also discussed in relation to its sensitivity to stress and its role in generating predictions about the environment.

💡Place Cells

Place cells are neurons in the hippocampus that become active when an animal is in a particular location. The video discusses the discovery of place cells by O'Keefe as a foundational moment in understanding how cognitive maps are represented in the brain.

💡Grid Cells

Grid cells are neurons that fire when an animal is in a specific location on a hexagonal grid, providing a coordinate system for spatial navigation. The video anticipates the discovery of grid cells as part of the cognitive map system, which was later confirmed by later research.

💡Head Direction Cells

Head direction cells are neurons that fire based on the direction an animal is facing. The video mentions that the prediction of such cells was made in the book by O'Keefe and Nadell, and their discovery in the mid-1980s confirmed another aspect of the cognitive map theory.

💡Multiple Memory Systems

Multiple memory systems refer to the idea that different types of memory are stored and processed by distinct neural systems in the brain. The video builds on Tolman's idea that there is more than one kind of learning, suggesting that different memory systems support various behavioral strategies.

💡Exploration

Exploration in the video is discussed as a key motivator for the creation of cognitive maps. It is suggested that animals explore their environment out of curiosity, and this exploration is what leads to the formation of cognitive maps, independent of immediate rewards such as food or safety.

💡Flexible Behavior

Flexible behavior refers to the ability to adapt actions based on changing circumstances or goals. The video argues that cognitive maps enable flexible behavior because they provide a rich information base that allows for various possible actions, rather than just a single, fixed response.

Highlights

Introduction to Tolman's celebration and his influence on cognitive mapping.

Lynn Nadel's introduction as the speaker and his background in cognitive science.

Tolman's concept of cognitive maps and their significance in neuroscience.

The idea that cognitive maps exist in multiple scales.

Tolman's view on different kinds of learning and their implications.

The historical controversy over place versus response learning.

Tolman's demonstration of latent learning and the role of curiosity in animal behavior.

The concept of expectation and how animals predict outcomes.

The connection between Tolman's scientific ideas and his political beliefs.

The discovery of place cells by O'Keefe and its impact on cognitive mapping theory.

The four key concepts in Nadel and O'Keefe's book about cognitive maps.

The role of exploration in creating cognitive maps.

The prediction of cells that convey direction and distance information.

The discovery of head direction cells and grid cells supporting the cognitive map theory.

The importance of cognitive maps in generating predictions and expectations.

The influence of stress and emotion on cognitive mapping and behavior.

Nadel's closing remarks and introduction of the next speaker, David Foster.

Transcripts

[Music]

welcome welcome to tolman's um

celebration in fact we heard this

morning how uh his um leading of the The

nonsigners Faculty nonsigners were met

in this building every Friday um during

during the fight so this is particularly

poignant that we're we were able to meet

in the morning in Tolman Hall and in the

afternoon in in The Faculty Club um so

it gives me great pleasure to introduce

the afternoon session now on the

cognitive map and where it has come

since Tolman coined the phrase and and

started the whole field um started the

ball rolling in the 19 uh 1940 um

7 um our first speaker is um ly nadell

and um Lynn is a a wonderful I'm so glad

he could come and and introduce um

this this session

today Lynn knows where to go get

educated he he first the first thing um

you need to do if you want to make a big

impact in this field is grow up in New

York City but then do all your degrees

Ma at um ba ma PhD at McGill hanging out

with the like of Donald Hebb and Brenda

Milner then go to swinging London for

seven years in the 1970s but while

you're there hang out with your um your

pal joh o Keef who is discovering

hippocampal play cells write a book

length Manifesto the hippocampus is a

cognitive map which for the first time

solves two huge problems in um

Neuroscience one how the brain makes a

map and two what the hippocampus

actually does continue on interesting

academic per paginations through Daly

Toronto San Diego Irvine and finally

settle down as regions Professor

psychology and cognitive science

director of the cognition and neural

systems program University of Arizona

while serving as department chair and

currently as chair of all University

faculty don't forget to collect more

awards for your work on the development

of memory on down syndrome and to knock

out another transformational theory this

time with Morris Mo mosovich on the

multiple Trace theory of human memory

consolidations please join me in

welcoming Linda

D thank you uh and I I have to call her

Lucy I mean apparently everyone calls

her Lu now but it's always been Lucy

thank you very much Lucy for inviting me

to this this is really a very meaningful

meeting for me uh and I was happy to be

able to come at this at at this point um

I teach the history of Psych and I teach

a variety of classes and every time I

introduce the issue of Tolman I start by

talking about his politics I mean it

doesn't start with what he said about

cognitive maps for me it starts with

what an amazing person this was so for

me this this really is a a personally

rewarding moment to be able to stand

here in this Faculty Club where he led

the Revolt uh on this campus and to be

able to contribute to this so thank you

for that so I'm going to be very brief

I'm just here to to do a brief

introduction and make a few small points

uh well maybe not so small but

smallish and I'm going to re be

repeating a few of the things that that

uh that Dr dubury mentioned earlier in

the day for those of you were at the at

the morning session but it starts with

this observation uh one of the

observations that Tolman made uh this is

a quote from that famous psych review

paper in

1948 uh but the point the few points I

want to make uh really are about the

fact that actually there was more than

just the notion that there are cognitive

maps in the brain and a bunch of those

ideas came out this morning but I want

to highlight a few of them because

they're very important for the theory

that O'Keefe and I then developed

and for the sort of the way the rest of

the field perhaps uh continued so one

point that that Tolman mentioned about

when he brought up the notion of

cognitive maps and this was already

mentioned earlier today can you hear me

back there good um was the notion that

these Maps may actually come in multiple

scales right there's narrow strip Map

There's a comprehensive broad map that

idea that there are kind of different

grains of mapping is a is an important

idea that actually we now know that

there's data to support that idea I

don't know if Edward's going to say

anything about that in particular but

but this general idea has survived and

it's an important idea not just that

there are maps but there are different

kind of kinds of maps

right second thing he also wrote a very

famous paper called there's more than

one kind of learning right and he

championed the idea and this came out in

the place versus response controversy

that we heard about today he championed

the idea that there were different kinds

of learning but what happened then

historically what happened then was that

Frank Westell came up with a solution to

the problem of that there are different

kinds of learning that placees place

learning and response learning according

to wrestle solution were not actually

that different it was just that the

animal was using different kinds of cues

there was nothing fundamentally

different about them they just depended

upon different cues that was not what

Tolman was saying what Tolman said was

that there were different kinds of

learning that supported different sorts

of Behavioral strategies andp hypotheses

and that these things might actually

have different properties that was an

important point that got lost with

wrestle solution but wrestle solution I

think it was a 57 paper is that right

you might remember it was something in

the in the 50s that kind of put the

story to bed for a while so people

stopped arguing about is it Place

Learning versus response learning and

they sort of decided that doesn't matter

it's just a question of what kind of

cues they're learning now we know that's

absolutely wrong that those two kinds of

learning depend upon very different

brain systems with rather different

properties they interact of course

they're all part of the same brain they

may compete with each other they may

they may uh cooperate with each other

but they're different and they have

different properties so that was a very

important Insight on tolman's point on

tolman's part wrestle so-called solution

another point that came up this morning

so so Tolman was very interested and and

was one of the first to demonstrate this

notion of latent learning and here the

key Point here was the big debate in the

1950s about the role of reward in

learning and whether or not learning

could happen in the absence of reward

and reinforcement and the discovery in

the late 40s early 50s Not discoveries

is the wrong word but the demonstration

that Curiosity and exploration can

motivate animals to do stuff you don't

have to give them rewards you can just

give them access to something novel and

and Tolman then focused on the fact that

animals will learn about the environment

in the absence of any extrinsic

motivation in the absence of any or

sorry intrinsic motivation no they don't

have to be hungry they don't have to be

thirsty they do need to be those things

perhaps to show you that they've learned

but they don't need be they don't need

to be like that to actually learn and

and this came out this morning in some

of the experiments on Laden learning so

here again Tolman was focusing on the

role of curiosity and exploration which

most psychologists at that point were

saying was

unimportant finally there was this point

about means and Readiness the sort of

sici significant the notion but what

what what was buried in this idea was

that Tolman was trying to get to the to

the to to the concept of expectation how

do anim anim animals seem to show you

that they are expecting something when

they and the tinkle paaw experiment love

that name I always tell my students

about that it's one of the great

experiments simply because of the name

of the experimenter um that tinkle paaw

experiment showed quite profoundly that

those and it was talked about this

morning but basically chimp is trained

I'm I'm going to give you a very short

version of it uh a monkey is trained uh

for a reward of bananas and then after

getting these bananas is suddenly given

some Lett and is pretty upset right

there was it was a better controlled

experiment than that but that's the

point showing that that animal was

expecting bananas right it wasn't just

good enough to give it some food it

actually had a specific expectation of

something so those expectations mean

that the brain is a kind of a predictor

it's predicting what is going to be

happening that of course we now you

can't open up a neuroscience book now

without seeing the predictive brain all

over the place but Tolman was one of the

first to point out how important this

was that animals are engaging in

constant predictions and expectations

being generated out of these maps that

they have built up right so those four

ideas are what leads to what

essentially before before I say two

words about how those ideas became the

basis of uh of the book that O'Keefe and

I

wrote I just wanted to again point out

and this came out this this morning that

there was in Tolman this clear

connection between science and politics

so when he talked about these narrow

strip maps and Broad comprehensive Maps

this was almost shorthand for him I

think to say narrow-minded people versus

broad-minded people right and that there

are circumstances you know brain damage

being one thing and and you know drugs

and all that other being another but

stress emotion frustration the things

that Lucy was pointing out all of those

things can actually shift the brain so

that it is you might say behaving more

on the basis of a narrow strip map than

on the basis of a comprehensive broad

map and that and we now know there's now

good data to show exactly how that

happens that stress that the hippocampus

is loaded with stress receptors and

stress can kind of selectively

downregulate the hippocampus and leave

other structures in in in charge of

behavior and those other structures are

more you might say survival oriented and

less exploration curiosity oriented

narrow your focus stay alive and so on

so again Tolman was on to something very

important here onto the differences

between the sort of brain states that

leg legitimate different forms of

behavior right so all four of those kind

of Notions including this one and one

thing I do want to point out this right

here see this by an overdose of

repetitions on the original this what

he's saying here is that when you

overtrain an animal it starts it becomes

a Hulan stimulus response animal and we

now know that that's true and we now

know the the the Neuroscience behind

that when you overtrain animals that the

striatum begins to Take Over Control of

the behavior there's a whole r of of

experiments buried in that idea so this

quote alone probably has six or seven

PhD dissertations you know buried in it

or could have probably

did all right so this was the book that

o'keef and I wrote in 78 which sort of

built on and it had a couple of key

Concepts amongst the key Concepts was

this one of course this book was was was

written based on the Discovery by o'keef

of play cells right it was that

discovery that sort of triggered you

know our thinking about this theory that

led to all of the other work and

everything that's come since then I

would say well not everything but you

know the spatial story so to speak so

the four key Concepts in the book the

first one in the title is that it's all

about cognitive maps and these are a

special kind of spatial system right

second key thing that the mapping system

was one kind of memory system

responsible for one specific kind of

information and that multiple memory

Systems Support various behaviors that's

a misprint there various behavioral

strategies so this again Builds on

tolman's idea that there was more than

one kind of learning and we really

insisted on this notion that these

different kinds of learning have

different properties and that those

different properties help us understand

what you see when an animal has damaged

to the hippocampus or when a person has

damaged to the

hippocampus third point we said very

very explicitly maps are created during

exploration so we bought completely into

woman's insistence that these cognitive

maps that what is the reward for an

animal in generating a cognitive map was

finding out about the world it was not

food it was not water it was not safety

although all of those things you might

find out about in exploring the world

but those were not the motivations in

any simple sense it was simple curiosity

to map the environment and exploration

was a key so in the book a chapter on

exploration was the first chapter in in

discussing the actual data of about the

effects of hippocampal lesions all right

and finally that Maps enable flexible

Behavior we made the argument that if

you have a map just like if you have a

map of San Francisco it doesn't tell you

you know where you are it simply tells

you if you're here and you want to get

there this is what you do right so a map

is a map sort of enables flexibility and

behavior because it's such a rich

information base rather than just if I'm

here turn left at the corner that was

going to get you to where you'd get to

if you turned left but it's not going to

tell you anything anything more than

that so those were the four ideas or

some of the main ideas conceptually in

the book I'm leaving us I mean I'm not

talking now about the physiology the

play cells these were the sort of

theoretical conceptual ideas and the

point is each one of those is one of the

ideas that here's tolman's idea and how

here's how it maps on to the things that

we said so this really we really did

build extensively on tolman's thinking

not only about the idea of a cognitive

map but about the property of cognitive

maps about the properties of Place

Learning and how that informs

Behavior so I will close I'm staying on

time I think we made a variety of

predictions in the book uh one of those

predictions was if this thing was really

a cognitive map then Not only would it

need the play cells that o'keef and dski

had described but it would also need

something that conveyed information

about direction and distance at a

minimum if it was going to function like

a map so there was built into this the

prediction that sooner or later

somebody's going to find cells that

provide direction information and those

the head Direction cells came when did

Ron discover the head Direction cells 80

the mid 80s and then the final and in

many ways the cement piece the grid

cells that uh that Edvard and my Brit M

had discovered and and so you're going

to hear about those and hear that aspect

of the story from Edvard as soon as I

get out of well not as soon as I get out

of the way because the first talk by

David Foster will be about this issue

about the hippocampus generating

predictions uh and we talked about that

in the book for the reasons I described

and but again we didn't know anything

about the mechanisms even though

actually John was I believe the first

person to notice the ripples uh and I

think there was a there's a mention in

the book about the ripples uh but we had

no idea what they were right and now we

understand that there are mechanisms in

hippocampal activity that have to do

with what's actually going on when an

animal is predicting what if I go down

this road what am I going to see and so

on and so forth so that talk now let's

finish this so David Foster will tell

you more about that particular part of

the story so with that introduction to

kind of where sort of how O'Keefe and I

built on the cognitive map story and

then lots of other really exciting

things happen and we're going to hear

about those exciting things now from

people who have actually done the work

so I'm going to get out of the way and

let you introduce

David

[Music]