chapter 7: getting around spatial learning by helen larzleer, scott mcgrath and valerie stinson

Chapter 7: Getting Chapter 7: Getting AroundAround

Spatial Learning bySpatial Learning by

Helen Larzleer,Helen Larzleer,

Scott McGrath and Scott McGrath and

Valerie StinsonValerie Stinson

OutlineOutline

7.1 Mechanisms for Spatial 7.1 Mechanisms for Spatial OrientationOrientation

7.2 How is Spatial Information 7.2 How is Spatial Information Integrated?Integrated?

7.3 Do Animals Have Cognitive Maps?7.3 Do Animals Have Cognitive Maps? 7.4 Acquiring Spatial Knowledge: The 7.4 Acquiring Spatial Knowledge: The

Conditions for LearningConditions for Learning Current ResearchCurrent Research DiscussionDiscussion

Dead ReckoningDead Reckoning

Dead reckoningDead reckoning: an internal sense of : an internal sense of the direction and distance of the goal the direction and distance of the goal from the animals current position.from the animals current position.

Path integrationPath integration: continuously : continuously integrating (in the mathematical sense) integrating (in the mathematical sense) information about the animals changes information about the animals changes in distance and direction to keep track in distance and direction to keep track of its location with respect to the of its location with respect to the predicted location of the nest.predicted location of the nest.


Dead reckoning allows Dead reckoning allows egocentricegocentric spatial location – animal is localizing spatial location – animal is localizing things in the environment with things in the environment with respect to itselfrespect to itself

AllocentricAllocentric or or geocentricgeocentric mechanisms locate the animal with mechanisms locate the animal with respect to some external cues respect to some external cues (landmarks, the sun, the earths (landmarks, the sun, the earths magnetic field).magnetic field).


Fig 7.2Fig 7.2


Disadvantages: 1) if the animal is Disadvantages: 1) if the animal is slowly “blown off course” path slowly “blown off course” path integration does not compensate. 2) integration does not compensate. 2) path integration accumulates errorpath integration accumulates error

To compensate: the farther an ant has To compensate: the farther an ant has traveled from the nest, the wider its traveled from the nest, the wider its spiraling loops when it returns. Ant spiraling loops when it returns. Ant turns left and right equally often, so turns left and right equally often, so errors tend to cancel out. errors tend to cancel out.

Ants make less large turns -> these Ants make less large turns -> these produce the greatest errors.produce the greatest errors.

BeaconsBeacons

Beacons (proximal cues)Beacons (proximal cues)

BeaconsBeacons

Beacons are cues close to a goal, Beacons are cues close to a goal, whereas landmarks (distal cues) whereas landmarks (distal cues) do not have to bedo not have to be

Animals can use both proximal Animals can use both proximal and distal cues, as illustrated by and distal cues, as illustrated by Morris (1981Morris (1981))

BeaconsBeacons

Figure 7.4Figure 7.4 Morris water Morris water

mazemaze

BeaconsBeacons

Distal and proximal cues are utilized by Distal and proximal cues are utilized by different areas of the brain, with distal different areas of the brain, with distal cues being more complicated to employcues being more complicated to employ

Evidence? Hippocampal lesion in rats Evidence? Hippocampal lesion in rats stops the use of distal cues, but stops the use of distal cues, but proximal cues are still utilizedproximal cues are still utilized

Why not only use proximal cues? Why not only use proximal cues? Because you must be within range of Because you must be within range of your target to use themyour target to use them

LandmarksLandmarks

Landmarks (Distal cues)Landmarks (Distal cues)

LandmarksLandmarks

Classic Classic demonstration demonstration by Tinbergen by Tinbergen and Kruyt and Kruyt (1938/1952) in (1938/1952) in digger waspsdigger wasps

Figure 7.5Figure 7.5

LandmarksLandmarks

Further research demonstrated Further research demonstrated that digger wasps prefer large, that digger wasps prefer large, nearby and 3-d objects as nearby and 3-d objects as landmarks, which makes sense as landmarks, which makes sense as these are easiest to seethese are easiest to see

This finding has been replicated This finding has been replicated with blue jays and honeybeeswith blue jays and honeybees

LandmarksLandmarks

Two distinguishable landmarks Two distinguishable landmarks are needed to specify a single are needed to specify a single positionposition

Question: how is information Question: how is information processed? Is there a continuum processed? Is there a continuum between competition and perfect between competition and perfect averaging?averaging?

LandmarksLandmarks

Gerbils use winner takes all, Gerbils use winner takes all, while pigeons use a form of while pigeons use a form of averagingaveraging


LandmarksLandmarks

Pigeons compute vector and Pigeons compute vector and directional information separately, in directional information separately, in different modules, and then combine different modules, and then combine output of these modulesoutput of these modules

Pigeons ->Pigeons ->

LandmarksLandmarks

Fig 7.7Fig 7.7

LandmarksLandmarks

Bees trained in 2 featureless huts Bees trained in 2 featureless huts demonstrate ability to distinguish demonstrate ability to distinguish between landmarks…practical between landmarks…practical application evident (must know application evident (must know which landmark is which)which landmark is which)

Bee ->Bee ->

LandmarksLandmarks

Bees use template matching, in Bees use template matching, in which they move in their current which they move in their current environment to reduce the environment to reduce the discrepancy with a stored discrepancy with a stored templatetemplate

Fig.7.8Fig.7.8

RoutesRoutes

““Learning a route” can refer to a Learning a route” can refer to a mechanism of egocentric orientation in mechanism of egocentric orientation in which an animal records the movements it which an animal records the movements it makes in traveling between two places. makes in traveling between two places. Usually referred to Usually referred to response learningresponse learning vs. vs. place learningplace learning. So, ‘run straight for a . So, ‘run straight for a certain distance then go left’ vs. ‘go to certain distance then go left’ vs. ‘go to goal box’goal box’

Lorenz’s (1952) water shrewsLorenz’s (1952) water shrews

RoutesRoutes

Advantages: can travel exceedingly fast Advantages: can travel exceedingly fast without wasting a minute on orientationwithout wasting a minute on orientation

Disadvantages: change in the Disadvantages: change in the environment is not detected immediately.environment is not detected immediately.

If an animal has learned a route based on If an animal has learned a route based on landmarks, altering the features of the landmarks, altering the features of the route should disorient itroute should disorient it

Water shrew ->Water shrew ->

Environmental ShapeEnvironmental Shape


Animals will use geometric information Animals will use geometric information about their environment as a spatial cue, about their environment as a spatial cue, such as the geometric relationship between such as the geometric relationship between objects or the shape of an enclosed spaceobjects or the shape of an enclosed space

Closely related to landmarks - the Closely related to landmarks - the arrangement of landmarks and the arrangement of landmarks and the geometric relationship between them form geometric relationship between them form the environmental shape in natural settingsthe environmental shape in natural settings

StudiesStudies

Cheng (1986): Rats trained to find food hidden in the Cheng (1986): Rats trained to find food hidden in the corner of a rectangular box will dig almost as often in corner of a rectangular box will dig almost as often in the diagonal corner (geometrically identical), even in the diagonal corner (geometrically identical), even in the presence of secondary cues such as colored or the presence of secondary cues such as colored or patterned walls and scents.patterned walls and scents.

Hermer & Spelke (1994, 1996): College students and Hermer & Spelke (1994, 1996): College students and 20 month old children tested in featureless room, 20 month old children tested in featureless room, made diagonal errors. When one wall colored blue, made diagonal errors. When one wall colored blue, college students stopped making diagonal errors, but college students stopped making diagonal errors, but babies did not improve.babies did not improve.


Implications: Both rats and people appear to Implications: Both rats and people appear to have a separate cognitive module to process have a separate cognitive module to process environmental shape. Throughout environmental shape. Throughout development, people overcome the reliance on development, people overcome the reliance on environmental shape and can make better use environmental shape and can make better use of other available cues. of other available cues.

Rozin (1976) refers to this change in cue Rozin (1976) refers to this change in cue dependency throughout development as dependency throughout development as “increasing accessibility of modular “increasing accessibility of modular processing”.processing”.


Landmarks and environmental shapeLandmarks and environmental shapeStudies:Studies:

Cartwright & Collett (1983): In studies in which Cartwright & Collett (1983): In studies in which 2 landmarks are separated or compressed, bees 2 landmarks are separated or compressed, bees will search in the middle of the moved will search in the middle of the moved landmarks, a distance proportional to the landmarks, a distance proportional to the amount the landmarks are moved. These results amount the landmarks are moved. These results are consistent with snapshot or template are consistent with snapshot or template matching model.matching model.


Spetch et al (1997): Pigeons and people Spetch et al (1997): Pigeons and people were “trained” to search in the middle of 4 were “trained” to search in the middle of 4 identical landmarks, and then the landmarks identical landmarks, and then the landmarks were separated. People still searched in the were separated. People still searched in the middle, while pigeons chose one or two middle, while pigeons chose one or two landmarks and maintained the same landmarks and maintained the same distance to them.distance to them.

In other research with Clark’s nutcrackers, In other research with Clark’s nutcrackers, the birds searched in the middle (Kamil & the birds searched in the middle (Kamil & Jones 1997).Jones 1997).


The Sun CompassThe Sun Compass


Using the sun for direction is Using the sun for direction is inherently complex. Why?inherently complex. Why?


The sun is constantly moving relative to The sun is constantly moving relative to the earth, so it is useless as a landmark.the earth, so it is useless as a landmark.

The movement of the sun through the sky The movement of the sun through the sky is different depending on both location on is different depending on both location on the earths surface and time of year. the earths surface and time of year.

So how do they do it?So how do they do it?


The ability to use the sun for The ability to use the sun for directional information requires two directional information requires two things: an things: an ephemeris functionephemeris function and an and an internal clock or time sense.internal clock or time sense.

Ephemeris function:Ephemeris function: “A stored “A stored representation of how the sun, or some representation of how the sun, or some correlate of it, moves across the sky at correlate of it, moves across the sky at the current location and season.”the current location and season.”


The compass direction of the sun The compass direction of the sun relative to the north is called the sun’s relative to the north is called the sun’s azimuth.azimuth.



StudiesStudies Wehner & Lanfranconi (1981): Desert ants Wehner & Lanfranconi (1981): Desert ants

get lost when they can’t see landmarks or get lost when they can’t see landmarks or the sun, but when they can see the proper the sun, but when they can see the proper position of the sun for the time of day they position of the sun for the time of day they head straight home.head straight home.

Papi & Wallraff (1992): When homing Papi & Wallraff (1992): When homing pigeons have their internal clocks either pigeons have their internal clocks either pushed forward or back by about 3 hours, pushed forward or back by about 3 hours, they will head off in the wrong direction they will head off in the wrong direction (about 45 degrees) when released.(about 45 degrees) when released.


Dyer (1987): Bees trained to find feeder based on sun Dyer (1987): Bees trained to find feeder based on sun compass and landmarks, and then the landmarks are compass and landmarks, and then the landmarks are moved. If released on a cloudy day, bees will use moved. If released on a cloudy day, bees will use landmark cues alone to find the new location of the landmark cues alone to find the new location of the feeder. When back at the hive, their dance reflects feeder. When back at the hive, their dance reflects the previous compass direction of the original feeder. the previous compass direction of the original feeder. However when the sun comes out, they immediately However when the sun comes out, they immediately change their dance to reflect the actual new compass change their dance to reflect the actual new compass direction of the feeder.direction of the feeder.

Dyer & Dickinson (1994):Young bees raised with their Dyer & Dickinson (1994):Young bees raised with their only experience in the sun being in late afternoon are only experience in the sun being in late afternoon are then released in the morning of a cloudy day. Their then released in the morning of a cloudy day. Their dances after returning to the hive are reversed from dances after returning to the hive are reversed from the afternoon, however the dance did not change the afternoon, however the dance did not change gradually but very abruptly at noon they changed 180 gradually but very abruptly at noon they changed 180 degrees.degrees.

Combining OutputsCombining Outputs

Information Integration: When do Information Integration: When do I use what?I use what?

Combining OutputsCombining Outputs

Modularity of spatial processing Modularity of spatial processing evidenced by apparently stupid evidenced by apparently stupid behaviour by animals i.e. ant behaviour by animals i.e. ant runs past nest, gerbil ignores runs past nest, gerbil ignores cries of babies, shrew jumps over cries of babies, shrew jumps over nonexistent stonenonexistent stone

Why?Why?

Rules for CombiningRules for Combining

If all cues will lead to the same If all cues will lead to the same goal, why not just use one goal, why not just use one instead of processing them all?instead of processing them all?

Evolution? New modules Evolution? New modules developed that are more flexible, developed that are more flexible, instead of modifying old modules.instead of modifying old modules.


Fig 7.14Fig 7.14

What is a Cognitive Map?What is a Cognitive Map?

Representations have three essential parts: a Representations have three essential parts: a represented system (eg distance and represented system (eg distance and direction) a representing system (eg an direction) a representing system (eg an animals nervous system), and rules for animals nervous system), and rules for correspondence between them.correspondence between them.

So, do animals have cognitive So, do animals have cognitive representations of the world in mind when representations of the world in mind when they travel?they travel?

What is a Cognitive Map?What is a Cognitive Map?


Cognitive MapsCognitive Maps

E.C. Tolman (1948) – introduced the E.C. Tolman (1948) – introduced the cognitive map.cognitive map. Claimed rats in Claimed rats in mazes learn about places.mazes learn about places.

Rat ->Rat ->

Cognitive MapsCognitive Maps

O’Keefe and Nadel – properties of cognitive O’Keefe and Nadel – properties of cognitive maps and how they might be acquired. maps and how they might be acquired. Acquisition and use of the cognitive map is Acquisition and use of the cognitive map is supported by the supported by the locale systemlocale system, a cognitive , a cognitive module located in the hippocampus of module located in the hippocampus of vertebrates. Contrasts the vertebrates. Contrasts the taxon systemtaxon system which supports route learning.which supports route learning.

Gallistel – definition of cognitive map is Gallistel – definition of cognitive map is considerably looser: any orientation based considerably looser: any orientation based on implicitly computing distances and on implicitly computing distances and directions rather than responding to a directions rather than responding to a beacon is evidence of a cognitive map.beacon is evidence of a cognitive map.

The local view hypothesisThe local view hypothesis(Leonard & McNaughton 1990), (McNaughton, Knierim & Wilson (Leonard & McNaughton 1990), (McNaughton, Knierim & Wilson

1995):1995):

There are no cognitive mapsThere are no cognitive maps

The animal has a set of memories of local views of the The animal has a set of memories of local views of the environment which are associatively linked to each environment which are associatively linked to each other by the memories of how to get from one to other by the memories of how to get from one to another.another.

Supported by neural net models & functions of certain Supported by neural net models & functions of certain populations of cells used in spatial learningpopulations of cells used in spatial learning

Behaviorally difficult to distinguish from cognitive mapsBehaviorally difficult to distinguish from cognitive maps

Alternatives to the cognitive mapAlternatives to the cognitive map

Alternatives to the cognitive mapAlternatives to the cognitive map

Bennett (1996):Bennett (1996): There is no good evidence for cognitive mapsThere is no good evidence for cognitive maps

Research in this area should be abandoned and Research in this area should be abandoned and instead focused on operational discussions of how instead focused on operational discussions of how animals get aroundanimals get around

Researchers don’t always mean the same thing by Researchers don’t always mean the same thing by ‘cognitive map’‘cognitive map’

The only agreed upon behavioral test for cognitive The only agreed upon behavioral test for cognitive mapping is an animal’s ability to take a novel route mapping is an animal’s ability to take a novel route without dead reckoning or generalization of local without dead reckoning or generalization of local viewsviews

Do bees have cognitive Do bees have cognitive maps?maps?

Experiment: Gould’s beesExperiment: Gould’s bees

Do bees have cognitive maps?Do bees have cognitive maps?

But…is there a problem with this But…is there a problem with this interpretation?interpretation?


When bees fly up to orient When bees fly up to orient themselves, can they get a visual themselves, can they get a visual representation of the situation representation of the situation and move to feeder A in order to and move to feeder A in order to reduce the discrepancy between reduce the discrepancy between their stored template and current their stored template and current situation?situation?

How can we know?How can we know?


Dyer replicated study, but feeder Dyer replicated study, but feeder B was placed in a quarryB was placed in a quarry

Bees trained at A, but placed at Bees trained at A, but placed at B, could not orient themselves, B, could not orient themselves, and flew in same direction they and flew in same direction they would as if they were leaving the would as if they were leaving the hivehive

Do Rats Have Cognitive Do Rats Have Cognitive Maps?Maps? Morris swim task – rats experienced in the Morris swim task – rats experienced in the

swimming task rapidly approached the swimming task rapidly approached the hidden platform. However – rats typically hidden platform. However – rats typically swim all over the tank in early training – swim all over the tank in early training – no location, view or route is completely no location, view or route is completely novel?novel?

When rats experience of different routes When rats experience of different routes has been restricted they are sometimes has been restricted they are sometimes unable to navigate successfully.unable to navigate successfully.

Keith & McVety (1988) Keith & McVety (1988)

Do Rats Have Cognitive Maps?Do Rats Have Cognitive Maps?

If an animal’s travels are controlled by If an animal’s travels are controlled by reference to a single overall allocentric reference to a single overall allocentric representation of space, it should not representation of space, it should not matter if information about different parts matter if information about different parts of a journey is obtained in different ways.of a journey is obtained in different ways.

Do Rats Have Cognitive Maps?Do Rats Have Cognitive Maps?

Disadvantages with rat research: Disadvantages with rat research: almost all done in labs and makes almost all done in labs and makes little or no reference to what the little or no reference to what the animals might be doing in nature.animals might be doing in nature.

Do other animals have Do other animals have cognitive maps?cognitive maps?

Do other animals have cognitive maps?Do other animals have cognitive maps?

Studies:Studies: Menzel (1978): Chimps were shown the locations of Menzel (1978): Chimps were shown the locations of

multiple items hidden in various locations and then multiple items hidden in various locations and then allowed to retrieve them. Their routes did appear to allowed to retrieve them. Their routes did appear to be efficient and novel in many cases, but the results be efficient and novel in many cases, but the results were not significantly different from chance.were not significantly different from chance.

Fig. 7.17 aFig. 7.17 a


Shettleworth & Krebs (1982): Marsh Shettleworth & Krebs (1982): Marsh tits take novel routes to retrieve tits take novel routes to retrieve stored seeds.stored seeds.


Sherry (1984): Black-capped chickadees remember Sherry (1984): Black-capped chickadees remember not only the locations of stored seeds but their not only the locations of stored seeds but their relative value.relative value.

Cramer & Gallistel (1996): In research with vervet Cramer & Gallistel (1996): In research with vervet monkeys, the monkeys appear to be planning ahead monkeys, the monkeys appear to be planning ahead to take the most efficient route depending on whether to take the most efficient route depending on whether or not they are returning to the starting point.or not they are returning to the starting point.

Fig. 7.17bFig. 7.17b


Clutton-Brock & Harvey (1977): Clutton-Brock & Harvey (1977): Larger brain size is correlated with Larger brain size is correlated with fruit-eating species as opposed to fruit-eating species as opposed to leaf-eating species of primates.leaf-eating species of primates.

ExplorationExploration

What is the purpose of What is the purpose of exploration?exploration?


Rats allowed to get reward on one table, Rats allowed to get reward on one table, removed for a period, then placed on another removed for a period, then placed on another table and allowed free travel to get back to table and allowed free travel to get back to reward tablereward table

If only allowed to explore one track, rats do If only allowed to explore one track, rats do not move to reward table above chance…not move to reward table above chance…seeing the connection does not indicate that it seeing the connection does not indicate that it can be travelledcan be travelled

If allowed to explore all tracks, rats perform If allowed to explore all tracks, rats perform well above chancewell above chance

If allowed to explore two tracks, performance If allowed to explore two tracks, performance is intermediateis intermediate

Latent Inhibition, Perceptual Latent Inhibition, Perceptual Learning, and Cognitive Learning, and Cognitive

MappingMapping O’Keefe and Nadel: place learning is O’Keefe and Nadel: place learning is

subserved by a special learning and subserved by a special learning and memory system, the local system, memory system, the local system, whereas response leaning, route whereas response leaning, route learning and classical conditioning are learning and classical conditioning are part of the taxon system. part of the taxon system.

Latent Inhibition, Perceptual Learning, and Cognitive Latent Inhibition, Perceptual Learning, and Cognitive MappingMapping

Latent Inhibition, Perceptual Learning, and Cognitive Latent Inhibition, Perceptual Learning, and Cognitive MappingMapping

In associative learning, exposure to a In associative learning, exposure to a situation may retard acquisition – that is, it situation may retard acquisition – that is, it can lead to can lead to latent inhibitionlatent inhibition. In contrast, . In contrast, exploring novel items in a familiar space is exploring novel items in a familiar space is assumed to allow an animal to update its assumed to allow an animal to update its cognitive map.cognitive map.

pre-exposure enhances discrimination (ie pre-exposure enhances discrimination (ie perceptual learning occurs) when the perceptual learning occurs) when the locations to be learned are similar, while locations to be learned are similar, while latent inhibition occurs when they are very latent inhibition occurs when they are very different.different.

Learning About Redundant Cues: Learning About Redundant Cues:

Competition or Parallel ProcessingCompetition or Parallel Processing

Does overshadowing and blocking occur Does overshadowing and blocking occur in spatial learning?in spatial learning?

Rescorla-Wager model describes trade-Rescorla-Wager model describes trade-off among potential cues, but off among potential cues, but redundancy makes more sense for redundancy makes more sense for important tasks like getting home – important tasks like getting home – redundant cues could be used as backup redundant cues could be used as backup if the primary cues were unavailable. Eg if the primary cues were unavailable. Eg experience homing pigeonsexperience homing pigeons

CalibrationCalibration


The act of checking or adjusting (by The act of checking or adjusting (by comparison with a standard) the accuracy of comparison with a standard) the accuracy of a measuring instrument. a measuring instrument.

Many species of animals appear to calibrate Many species of animals appear to calibrate orientation methods against one another.orientation methods against one another.

Most commonly seen in migratory birds who Most commonly seen in migratory birds who adjust their magnetic compass against the adjust their magnetic compass against the sun compass or celestial cues.sun compass or celestial cues.

Why would they adjust a magnetic compass Why would they adjust a magnetic compass to the sun compass or celestial cues?to the sun compass or celestial cues?


Because the earth’s magnetic field changes Because the earth’s magnetic field changes in strength from one place to another (the in strength from one place to another (the sun compass and celestial cues are more sun compass and celestial cues are more accurate).accurate).


Studies:Studies: Able (1991), Able & Able (1990): Savannah Able (1991), Able & Able (1990): Savannah

sparrows raised indoors would orient based sparrows raised indoors would orient based entirely upon magnetic cues. After different entirely upon magnetic cues. After different groups of the birds were subjected to groups of the birds were subjected to different directional cues (sun, celestial, and different directional cues (sun, celestial, and artificially altered magnetic field), the birds artificially altered magnetic field), the birds adjusted (calibrated) their magnetic adjusted (calibrated) their magnetic compasses to a different extent depending compasses to a different extent depending on which cues they had been exposed to.on which cues they had been exposed to.


Fig 7.21Fig 7.21


Emlen (1970): Raised 3 groups of indigo buntings Emlen (1970): Raised 3 groups of indigo buntings indoors with no view of the sky. 2 of the groups were indoors with no view of the sky. 2 of the groups were then exposed to “the sky” in a planetarium – One then exposed to “the sky” in a planetarium – One group an accurate representation of it, the other an group an accurate representation of it, the other an altered view in which the axis of rotation centered on altered view in which the axis of rotation centered on Betelgeuse. Betelgeuse.

All 3 groups were later tested under a stationary sky. All 3 groups were later tested under a stationary sky.

The group with no experience did not use celestial The group with no experience did not use celestial cues to orientcues to orient

The group with accurate experience oriented The group with accurate experience oriented properly based on the position of the North Starproperly based on the position of the North Star

The 3rd group oriented as if Betelgeuse was the The 3rd group oriented as if Betelgeuse was the North StarNorth Star

Current ResearchCurrent Research

DiscussionDiscussion

chapter 7: getting around spatial learning by helen larzleer, scott mcgrath and valerie stinson

Documents

use of distal cues

external cues landmarks

spatial information

landmarksgerbils use

mathematical sense information

spatial knowledge

spatial orientation7

animal records