Abstract
Humans think about order and number using a spatial
representation, and the orientation of this representation is
influenced by learning to read and count in a particular direction.
For example, people who read and count from left to right represent
small/early items on the left side of space and large/late items on
the right side of space. Evidence that pre-verbal children organize
order and number spatially raises the possibility that humans are
predisposed to organize magnitude and order using space without
explicit language training. Nonhuman animals also represent
magnitudes and order. I tested the hypothesis that nonhuman
primates use spatial representations to code order. I reasoned that
if ordering depends on spatial representation then performance on
an ordering task would be more impaired when performed concurrently
with a spatial memory task than when performed concurrently with a
non-spatial visual matching task. Across species and tasks I found
that concurrent cognitive load impaired performance, but I did not
find that this impairment was especially large with concurrent
spatial and ordinal processing. In Manuscript 1, orangutan and
chimpanzee performance was generally impaired in all concurrent
cognitive load conditions, suggesting that spatial memory, visual
memory, and ordering are all supported by a general working memory
resource. In Manuscript 2, I tested rhesus monkeys on a similar set
of experiments in which an ordering task was embedded within a
spatial memory task and a non-spatial visual matching-to-sample
task. I again found that spatial memory was impaired under all
concurrent cognitive load conditions. Taken together, this set of
experiments suggests that there is a domain general working memory
resource supporting spatial and visual cognition in apes and
monkeys, rather than a resource supporting spatial and ordinal
processing specifically.
Table of Contents
General Introduction 1
Spatial representations of magnitude and order in humans 2
A mental number line 2
Spatial representations of order 5
Animals show patterns that suggest spatial representations across
tasks involving ordered stimuli 9
Transitive inference 10
Evidence for spatial representations in other ordering paradigms
16
Direct tests of spatial representations in chicks 19
Direct tests of space-order associations in nonhuman primates
21
The parietal lobe is implicated in both spatial and ordinal
cognition across species 24
Human imaging studies 25
Nonhuman primate single-cell recordings 27
Conclusion 28
Introduction to Thesis 30
Manuscript 1: Ordering images requires spatial working memory in
apes 33
Abstract 34
Introduction 35
General Methods 40
Subjects and Apparatus 40
Experiment 1 41
Procedure 41
Results and Discussion 49
Experiment 2: Intermixed Test 53
Procedure 53
Results and Discussion 54
Experiment 3: Ordering Embedded in Spatial Memory and Delayed MTS
58
Methods 58
Results and Discussion 66
General Discussion 70
Introduction to Manuscript 2 73
Manuscript 2: Spatial representations for order in monkeys are not
supported by a domain specific spatial working memory resource
74
Abstract 75
Introduction 76
Methods 83
Subjects 83
Apparatus 83
Procedure 84
Experiment 1: Concurrent Cognitive Load Test Ordering Two Items
90
Results and Discussion 93
Experiment 2: Concurrent Cognitive Load Test Ordering 3 Items
97
Methods 98
Results and Discussion 99
Experiment 3: Object Discrimination 102
Methods 103
Results and Discussion 108
Experiment 4: Ordering and Object Discrimination Together 114
Methods 114
Results and Discussion 116
General Discussion 120
General Discussion 125
Three species and three testing environments 127
Spatial memory as a general resource 129
Other questions this thesis does not address 131
Summary 132
References 133
About this Dissertation
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