1. Chapter 7: Getting Around Spatial Learning by Helen Larzleer, Scott McGrath and Valerie Stinson

2. Outline 7.1 Mechanisms for Spatial Orientation 7.1 Mechanisms for Spatial Orientation 7.2 How is Spatial Information Integrated? 7.2 How is Spatial Information Integrated? 7.3 Do Animals Have Cognitive Maps? 7.3 Do Animals Have Cognitive Maps? 7.4 Acquiring Spatial Knowledge: The Conditions for Learning 7.4 Acquiring Spatial Knowledge: The Conditions for Learning Current Research Current Research Discussion Discussion

3. Dead Reckoning Dead reckoning: an internal sense of the direction and distance of the goal from the animals current position. Dead reckoning: an internal sense of the direction and distance of the goal from the animals current position. Path integration: continuously integrating (in the mathematical sense) information about the animals changes in distance and direction to keep track of its location with respect to the predicted location of the nest. Path integration: continuously integrating (in the mathematical sense) information about the animals changes in distance and direction to keep track of its location with respect to the predicted location of the nest.

4. Dead Reckoning Dead reckoning allows egocentric spatial location – animal is localizing things in the environment with respect to itself Dead reckoning allows egocentric spatial location – animal is localizing things in the environment with respect to itself Allocentric or geocentric mechanisms locate the animal with respect to some external cues (landmarks, the sun, the earths magnetic field). Allocentric or geocentric mechanisms locate the animal with respect to some external cues (landmarks, the sun, the earths magnetic field).

5. Dead Reckoning Fig 7.2 Fig 7.2

6. Dead Reckoning Disadvantages: 1) if the animal is slowly “blown off course” path integration does not compensate. 2) path integration accumulates error Disadvantages: 1) if the animal is slowly “blown off course” path integration does not compensate. 2) path integration accumulates error To compensate: the farther an ant has traveled from the nest, the wider its spiraling loops when it returns. Ant turns left and right equally often, so errors tend to cancel out. To compensate: the farther an ant has traveled from the nest, the wider its spiraling loops when it returns. Ant turns left and right equally often, so errors tend to cancel out. Ants make less large turns -> these produce the greatest errors. Ants make less large turns -> these produce the greatest errors.

7. Beacons Beacons (proximal cues) Beacons (proximal cues)

8. Beacons Beacons are cues close to a goal, whereas landmarks (distal cues) do not have to be Beacons are cues close to a goal, whereas landmarks (distal cues) do not have to be Animals can use both proximal and distal cues, as illustrated by Morris (1981) Animals can use both proximal and distal cues, as illustrated by Morris (1981)

9. Beacons Figure 7.4 Figure 7.4 Morris water maze Morris water maze

10. Beacons Distal and proximal cues are utilized by different areas of the brain, with distal cues being more complicated to employ Distal and proximal cues are utilized by different areas of the brain, with distal cues being more complicated to employ Evidence? Hippocampal lesion in rats stops the use of distal cues, but proximal cues are still utilized Evidence? Hippocampal lesion in rats stops the use of distal cues, but proximal cues are still utilized Why not only use proximal cues? Because you must be within range of your target to use them Why not only use proximal cues? Because you must be within range of your target to use them

11. Landmarks Landmarks (Distal cues) Landmarks (Distal cues)

12. Landmarks Classic demonstration by Tinbergen and Kruyt (1938/1952) in digger wasps Classic demonstration by Tinbergen and Kruyt (1938/1952) in digger wasps Figure 7.5 Figure 7.5

13. Landmarks Further research demonstrated that digger wasps prefer large, nearby and 3-d objects as landmarks, which makes sense as these are easiest to see Further research demonstrated that digger wasps prefer large, nearby and 3-d objects as landmarks, which makes sense as these are easiest to see This finding has been replicated with blue jays and honeybees This finding has been replicated with blue jays and honeybees

14. Landmarks Two distinguishable landmarks are needed to specify a single position Two distinguishable landmarks are needed to specify a single position Question: how is information processed? Is there a continuum between competition and perfect averaging? Question: how is information processed? Is there a continuum between competition and perfect averaging?

15. Landmarks Gerbils use winner takes all, while pigeons use a form of averaging Gerbils use winner takes all, while pigeons use a form of averaging Figure 7.6 Figure 7.6

16. Landmarks Pigeons compute vector and directional information separately, in different modules, and then combine output of these modules Pigeons compute vector and directional information separately, in different modules, and then combine output of these modules Pigeons -> Pigeons ->

17. Landmarks Fig 7.7 Fig 7.7

18. Landmarks Bees trained in 2 featureless huts demonstrate ability to distinguish between landmarks…practical application evident (must know which landmark is which) Bees trained in 2 featureless huts demonstrate ability to distinguish between landmarks…practical application evident (must know which landmark is which) Bee -> Bee ->

19. Landmarks Bees use template matching, in which they move in their current environment to reduce the discrepancy with a stored template Bees use template matching, in which they move in their current environment to reduce the discrepancy with a stored template Fig.7.8 Fig.7.8

20. Routes “Learning a route” can refer to a mechanism of egocentric orientation in which an animal records the movements it makes in traveling between two places. Usually referred to response learning vs. place learning. So, ‘run straight for a certain distance then go left’ vs. ‘go to goal box’ “Learning a route” can refer to a mechanism of egocentric orientation in which an animal records the movements it makes in traveling between two places. Usually referred to response learning vs. place learning. So, ‘run straight for a certain distance then go left’ vs. ‘go to goal box’ Lorenz’s (1952) water shrews Lorenz’s (1952) water shrews

21. Routes Advantages: can travel exceedingly fast without wasting a minute on orientation Advantages: can travel exceedingly fast without wasting a minute on orientation Disadvantages: change in the environment is not detected immediately. Disadvantages: change in the environment is not detected immediately. If an animal has learned a route based on landmarks, altering the features of the route should disorient it If an animal has learned a route based on landmarks, altering the features of the route should disorient it Water shrew -> Water shrew ->

22. Environmental Shape

23. Animals will use geometric information about their environment as a spatial cue, such as the geometric relationship between objects or the shape of an enclosed space Animals will use geometric information about their environment as a spatial cue, such as the geometric relationship between objects or the shape of an enclosed space Closely related to landmarks - the arrangement of landmarks and the geometric relationship between them form the environmental shape in natural settings Closely related to landmarks - the arrangement of landmarks and the geometric relationship between them form the environmental shape in natural settings

24. Studies Studies Cheng (1986): Rats trained to find food hidden in the corner of a rectangular box will dig almost as often in the diagonal corner (geometrically identical), even in the presence of secondary cues such as colored or patterned walls and scents. Cheng (1986): Rats trained to find food hidden in the corner of a rectangular box will dig almost as often in the diagonal corner (geometrically identical), even in the presence of secondary cues such as colored or patterned walls and scents. Hermer & Spelke (1994, 1996): College students and 20 month old children tested in featureless room, made diagonal errors. When one wall colored blue, college students stopped making diagonal errors, but babies did not improve. Hermer & Spelke (1994, 1996): College students and 20 month old children tested in featureless room, made diagonal errors. When one wall colored blue, college students stopped making diagonal errors, but babies did not improve. Environmental Shape

25. Figure 7.9 Figure 7.9

26. Implications: Both rats and people appear to have a separate cognitive module to process environmental shape. Throughout development, people overcome the reliance on environmental shape and can make better use of other available cues. Implications: Both rats and people appear to have a separate cognitive module to process environmental shape. Throughout development, people overcome the reliance on environmental shape and can make better use of other available cues. Rozin (1976) refers to this change in cue dependency throughout development as “increasing accessibility of modular processing”. Rozin (1976) refers to this change in cue dependency throughout development as “increasing accessibility of modular processing”. Environmental Shape

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

28. Spetch et al (1997): Pigeons and people were “trained” to search in the middle of 4 identical landmarks, and then the landmarks were separated. People still searched in the middle, while pigeons chose one or two landmarks and maintained the same distance to them. Spetch et al (1997): Pigeons and people were “trained” to search in the middle of 4 identical landmarks, and then the landmarks were separated. People still searched in the middle, while pigeons chose one or two landmarks and maintained the same distance to them. In other research with Clark’s nutcrackers, the birds searched in the middle (Kamil & Jones 1997). In other research with Clark’s nutcrackers, the birds searched in the middle (Kamil & Jones 1997). Environmental Shape

29. Figure 7.10 Figure 7.10

30. The Sun Compass

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

32. The Sun Compass The sun is constantly moving relative to the earth, so it is useless as a landmark. The sun is constantly moving relative to the earth, so it is useless as a landmark. The movement of the sun through the sky is different depending on both location on the earths surface and time of year. The movement of the sun through the sky is different depending on both location on the earths surface and time of year. So how do they do it?

33. The Sun Compass The ability to use the sun for directional information requires two things: an ephemeris function and an internal clock or time sense. The ability to use the sun for directional information requires two things: an ephemeris function and an internal clock or time sense. Ephemeris function: “A stored representation of how the sun, or some correlate of it, moves across the sky at the current location and season.” Ephemeris function: “A stored representation of how the sun, or some correlate of it, moves across the sky at the current location and season.”

34. The Sun Compass The compass direction of the sun relative to the north is called the sun’s azimuth. The compass direction of the sun relative to the north is called the sun’s azimuth. Figure 7.11 Figure 7.11

35. The Sun Compass Studies Wehner & Lanfranconi (1981): Desert ants get lost when they can’t see landmarks or the sun, but when they can see the proper position of the sun for the time of day they head straight home. Wehner & Lanfranconi (1981): Desert ants get lost when they can’t see landmarks or the sun, but when they can see the proper position of the sun for the time of day they head straight home. Papi & Wallraff (1992): When homing pigeons have their internal clocks either pushed forward or back by about 3 hours, they will head off in the wrong direction (about 45 degrees) when released. Papi & Wallraff (1992): When homing pigeons have their internal clocks either pushed forward or back by about 3 hours, they will head off in the wrong direction (about 45 degrees) when released.

36. The Sun Compass Dyer (1987): Bees trained to find feeder based on sun compass and landmarks, and then the landmarks are moved. If released on a cloudy day, bees will use landmark cues alone to find the new location of the feeder. When back at the hive, their dance reflects the previous compass direction of the original feeder. However when the sun comes out, they immediately change their dance to reflect the actual new compass direction of the feeder. Dyer (1987): Bees trained to find feeder based on sun compass and landmarks, and then the landmarks are moved. If released on a cloudy day, bees will use landmark cues alone to find the new location of the feeder. When back at the hive, their dance reflects the previous compass direction of the original feeder. However when the sun comes out, they immediately change their dance to reflect the actual new compass direction of the feeder. Dyer & Dickinson (1994):Young bees raised with their only experience in the sun being in late afternoon are then released in the morning of a cloudy day. Their dances after returning to the hive are reversed from the afternoon, however the dance did not change gradually but very abruptly at noon they changed 180 degrees. Dyer & Dickinson (1994):Young bees raised with their only experience in the sun being in late afternoon are then released in the morning of a cloudy day. Their dances after returning to the hive are reversed from the afternoon, however the dance did not change gradually but very abruptly at noon they changed 180 degrees.

37. Combining Outputs Information Integration: When do I use what? Information Integration: When do I use what?

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

39. Rules for Combining If all cues will lead to the same goal, why not just use one instead of processing them all? If all cues will lead to the same goal, why not just use one instead of processing them all? Evolution? New modules developed that are more flexible, instead of modifying old modules. Evolution? New modules developed that are more flexible, instead of modifying old modules.

40. Rules for Combining Figure 7.13 Figure 7.13

41. Rules for Combining Fig 7.14 Fig 7.14

42. What is a Cognitive Map? Representations have three essential parts: a represented system (eg distance and direction) a representing system (eg an animals nervous system), and rules for correspondence between them. Representations have three essential parts: a represented system (eg distance and direction) a representing system (eg an animals nervous system), and rules for correspondence between them. So, do animals have cognitive representations of the world in mind when they travel? So, do animals have cognitive representations of the world in mind when they travel?

43. What is a Cognitive Map? Figure 7.16 Figure 7.16

44. Cognitive Maps E.C. Tolman (1948) – introduced the cognitive map. Claimed rats in mazes learn about places. E.C. Tolman (1948) – introduced the cognitive map. Claimed rats in mazes learn about places. Rat -> Rat ->

45. Cognitive Maps O’Keefe and Nadel – properties of cognitive maps and how they might be acquired. Acquisition and use of the cognitive map is supported by the locale system, a cognitive module located in the hippocampus of vertebrates. Contrasts the taxon system which supports route learning. O’Keefe and Nadel – properties of cognitive maps and how they might be acquired. Acquisition and use of the cognitive map is supported by the locale system, a cognitive module located in the hippocampus of vertebrates. Contrasts the taxon system which supports route learning. Gallistel – definition of cognitive map is considerably looser: any orientation based on implicitly computing distances and directions rather than responding to a beacon is evidence of a cognitive map. Gallistel – definition of cognitive map is considerably looser: any orientation based on implicitly computing distances and directions rather than responding to a beacon is evidence of a cognitive map.

46. The local view hypothesis (Leonard & McNaughton 1990), (McNaughton, Knierim & Wilson 1995): There are no cognitive maps There are no cognitive maps The animal has a set of memories of local views of the environment which are associatively linked to each other by the memories of how to get from one to another. The animal has a set of memories of local views of the environment which are associatively linked to each other by the memories of how to get from one to another. Supported by neural net models & functions of certain populations of cells used in spatial learning Supported by neural net models & functions of certain populations of cells used in spatial learning Behaviorally difficult to distinguish from cognitive maps Behaviorally difficult to distinguish from cognitive maps Alternatives to the cognitive map

47. Bennett (1996): There is no good evidence for cognitive maps There is no good evidence for cognitive maps Research in this area should be abandoned and instead focused on operational discussions of how animals get around Research in this area should be abandoned and instead focused on operational discussions of how animals get around Researchers don’t always mean the same thing by ‘cognitive map’ Researchers don’t always mean the same thing by ‘cognitive map’ The only agreed upon behavioral test for cognitive mapping is an animal’s ability to take a novel route without dead reckoning or generalization of local views The only agreed upon behavioral test for cognitive mapping is an animal’s ability to take a novel route without dead reckoning or generalization of local views

48. Do bees have cognitive maps? Experiment: Gould’s bees Experiment: Gould’s bees

49. Do bees have cognitive maps? But…is there a problem with this interpretation? But…is there a problem with this interpretation?

50. Do bees have cognitive maps? When bees fly up to orient themselves, can they get a visual representation of the situation and move to feeder A in order to reduce the discrepancy between their stored template and current situation? When bees fly up to orient themselves, can they get a visual representation of the situation and move to feeder A in order to reduce the discrepancy between their stored template and current situation? How can we know? How can we know?

51. Do bees have cognitive maps? Dyer replicated study, but feeder B was placed in a quarry Dyer replicated study, but feeder B was placed in a quarry Bees trained at A, but placed at B, could not orient themselves, and flew in same direction they would as if they were leaving the hive Bees trained at A, but placed at B, could not orient themselves, and flew in same direction they would as if they were leaving the hive

52. Do Rats Have Cognitive Maps? Morris swim task – rats experienced in the swimming task rapidly approached the hidden platform. However – rats typically swim all over the tank in early training – no location, view or route is completely novel? Morris swim task – rats experienced in the swimming task rapidly approached the hidden platform. However – rats typically swim all over the tank in early training – no location, view or route is completely novel? When rats experience of different routes has been restricted they are sometimes unable to navigate successfully. When rats experience of different routes has been restricted they are sometimes unable to navigate successfully. Keith & McVety (1988) Keith & McVety (1988)

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

54. Do Rats Have Cognitive Maps? Disadvantages with rat research: almost all done in labs and makes little or no reference to what the animals might be doing in nature. Disadvantages with rat research: almost all done in labs and makes little or no reference to what the animals might be doing in nature.

55. Do other animals have cognitive maps?

56. Studies:  Menzel (1978): Chimps were shown the locations of multiple items hidden in various locations and then allowed to retrieve them. Their routes did appear to be efficient and novel in many cases, but the results were not significantly different from chance.  Fig. 7.17 a

57. Do other animals have cognitive maps? Shettleworth & Krebs (1982): Marsh tits take novel routes to retrieve stored seeds. Shettleworth & Krebs (1982): Marsh tits take novel routes to retrieve stored seeds.

58. Do other animals have cognitive maps? Sherry (1984): Black-capped chickadees remember not only the locations of stored seeds but their relative value. Sherry (1984): Black-capped chickadees remember not only the locations of stored seeds but their relative value. Cramer & Gallistel (1996): In research with vervet monkeys, the monkeys appear to be planning ahead to take the most efficient route depending on whether or not they are returning to the starting point. Cramer & Gallistel (1996): In research with vervet monkeys, the monkeys appear to be planning ahead to take the most efficient route depending on whether or not they are returning to the starting point. Fig. 7.17b Fig. 7.17b

59. Do other animals have cognitive maps? Clutton-Brock & Harvey (1977): Larger brain size is correlated with fruit-eating species as opposed to leaf-eating species of primates. Clutton-Brock & Harvey (1977): Larger brain size is correlated with fruit-eating species as opposed to leaf-eating species of primates.

60. Exploration What is the purpose of exploration? What is the purpose of exploration?

61. Exploration

62. Exploration Rats allowed to get reward on one table, removed for a period, then placed on another table and allowed free travel to get back to reward table Rats allowed to get reward on one table, removed for a period, then placed on another table and allowed free travel to get back to reward table If only allowed to explore one track, rats do not move to reward table above chance…seeing the connection does not indicate that it can be travelled If only allowed to explore one track, rats do not move to reward table above chance…seeing the connection does not indicate that it can be travelled If allowed to explore all tracks, rats perform well above chance If allowed to explore all tracks, rats perform well above chance If allowed to explore two tracks, performance is intermediate If allowed to explore two tracks, performance is intermediate

63. Latent Inhibition, Perceptual Learning, and Cognitive Mapping O’Keefe and Nadel: place learning is subserved by a special learning and memory system, the local system, whereas response leaning, route learning and classical conditioning are part of the taxon system. O’Keefe and Nadel: place learning is subserved by a special learning and memory system, the local system, whereas response leaning, route learning and classical conditioning are part of the taxon system.

64. Latent Inhibition, Perceptual Learning, and Cognitive Mapping

65. In associative learning, exposure to a situation may retard acquisition – that is, it can lead to latent inhibition. In contrast, exploring novel items in a familiar space is assumed to allow an animal to update its cognitive map. In associative learning, exposure to a situation may retard acquisition – that is, it can lead to latent inhibition. In contrast, exploring novel items in a familiar space is assumed to allow an animal to update its cognitive map. pre-exposure enhances discrimination (ie perceptual learning occurs) when the locations to be learned are similar, while latent inhibition occurs when they are very different. pre-exposure enhances discrimination (ie perceptual learning occurs) when the locations to be learned are similar, while latent inhibition occurs when they are very different.

66. Learning About Redundant Cues: Competition or Parallel Processing Does overshadowing and blocking occur in spatial learning? Does overshadowing and blocking occur in spatial learning? Rescorla-Wager model describes trade-off among potential cues, but redundancy makes more sense for important tasks like getting home – redundant cues could be used as backup if the primary cues were unavailable. Eg experience homing pigeons Rescorla-Wager model describes trade-off among potential cues, but redundancy makes more sense for important tasks like getting home – redundant cues could be used as backup if the primary cues were unavailable. Eg experience homing pigeons

67. Calibration

68. Calibration The act of checking or adjusting (by comparison with a standard) the accuracy of a measuring instrument. The act of checking or adjusting (by comparison with a standard) the accuracy of a measuring instrument. Many species of animals appear to calibrate orientation methods against one another. Many species of animals appear to calibrate orientation methods against one another. Most commonly seen in migratory birds who adjust their magnetic compass against the sun compass or celestial cues. Most commonly seen in migratory birds who adjust their magnetic compass against the sun compass or celestial cues. Why would they adjust a magnetic compass to the sun compass or celestial cues?

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

70. Calibration Studies: Able (1991), Able & Able (1990): Savannah sparrows raised indoors would orient based entirely upon magnetic cues. After different groups of the birds were subjected to different directional cues (sun, celestial, and artificially altered magnetic field), the birds adjusted (calibrated) their magnetic compasses to a different extent depending on which cues they had been exposed to. Able (1991), Able & Able (1990): Savannah sparrows raised indoors would orient based entirely upon magnetic cues. After different groups of the birds were subjected to different directional cues (sun, celestial, and artificially altered magnetic field), the birds adjusted (calibrated) their magnetic compasses to a different extent depending on which cues they had been exposed to.

71. Calibration Fig 7.21 Fig 7.21

72. Calibration Emlen (1970): Raised 3 groups of indigo buntings indoors with no view of the sky. 2 of the groups were then exposed to “the sky” in a planetarium – One group an accurate representation of it, the other an altered view in which the axis of rotation centered on Betelgeuse. Emlen (1970): Raised 3 groups of indigo buntings indoors with no view of the sky. 2 of the groups were then exposed to “the sky” in a planetarium – One group an accurate representation of it, the other an altered view in which the axis of rotation centered on 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 cues to orient The group with no experience did not use celestial cues to orient The group with accurate experience oriented properly based on the position of the North Star The group with accurate experience oriented properly based on the position of the North Star The 3rd group oriented as if Betelgeuse was the North Star The 3rd group oriented as if Betelgeuse was the North Star

73. Current Research

74. Discussion