Memory for order in monkeys (Macaca mulatta) Pubblico

Templer, Victoria Lynn (2014)

Permanent URL: https://etd.library.emory.edu/concern/etds/k930bx24d?locale=it
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Abstract

One important aspect of episodic memory is the ability to remember the unique order of events that comprise personal experience. For example, remembering that you first went snorkeling, then visited a volcano, and finally went sailing, is essential to accurately retelling your vacation story. Mnemonic representation of order is also necessary when actions are taken in routinized habit sequences, such as executing a cooking recipe. The distinction between episodic and habit memory is fundamental to theories of human memory, but it is difficult to test in nonhuman animals. Cognitive tests suitable for nonverbal species have been developed that model the temporal order aspect of consecutively experienced unique events and the order of routinized responses in a serial order task. This dissertation includes four papers that explain attempts to characterize memory for order of events and order of responses in rhesus monkeys (Macaca mulatta). Paper 1 is a review article about the study of episodic memory in nonhuman animals. Paper 2 presents evidence that monkeys use a temporal order mechanism, rather than knowledge of ordinal position, or relative recency, to remember the order in which trial-unique images were seen and touched. Monkeys better encoded order when intervening images occurred, but not when unfilled intervals of equivalent duration occurred, indicating that the occurrence of intervening events may serve to mark the passage of time, thereby increasing the subjective separation of events. Paper 3 provides evidence that when fixed orders are repeatedly executed in a serial order task, knowledge of ordinal position, rather than image-image associations, maintains mental representation of order. Errors during list execution were consistently prospective, indicating use of a prospective rather than retrospective code to update position in the list. Paper 4 reviews evidence that suggests the study of order memory along analogous cognitive behavioral manipulations across paradigms is a promising avenue for understanding the types of memory systems existing in monkeys. Together, these studies provide characterizations of cognitive representations underlying memory for ordered stimuli. Ultimately these findings help make small steps towards answering questions about whether the types of memory systems nonhuman primates possess overlap with those that define common human memory taxonomies.

Table of Contents

Table of Contents

1.General introduction………………...……………………….……......1

1.1. Episodic memory…………………………………….…...............3
1.2. Aims………………….…………………………………….................5
1.3. Approach……………..……………………………………..............8

Introduction to Paper 1……………………………………...….........10

Paper 1: Episodic memory in nonhuman animals………….....11
3.1. Abstract………………………………………………...……….........12
3.2. Introduction………………………………………….…................13
3.3. Moving beyond anthropocentrism and

phenomenology………………………………………........................15
3.4. Neurobiological approaches………………………….………......17
3.5. Cognitive behavioral approaches……………….…….…….....19
3.6. What is mentally represented in episodic memory?......21
3.7. Memory is for the future……………………………................21
3.8. Recall and recognition………………………………..………........22
3.9. Convergence of cognition and neurobiology…...…….…...23
3.10. Gaps and future studies……………………………………….......23
3.11. Conclusions………………………..…………………………............26

4. Introduction to Paper 2………………………………………............28

Paper 2: Cognitive mechanisms of memory for order in

rhesus monkeys (Macaca mulatta)……………………….………..…...29
5.1. Abstract……………………………………………….…….................30
5.2. Introduction……………...……………………………..…...............31
5.3. Materials and methods………………………………..……...........32
5.4. Experiment 1-Training on non-adjacent images and

transfer to adjacent images…………………………………….……......34
5.5. Experiment 2- Evaluating the influence of

memory strength……………………………………………….…..............38
5.6. Experiment 3- Evaluating the influence of list position..42
5.7. Experiment 4- Temporal order: intervening images

and intervening time…..……………………………….….………...........46
5.8. Discussion…………………………………………..….……...............51

6.Introduction to Paper 3…………………………………………...........56

Paper 3: Memory for ordinal position in monkeys………..…......57
7.1. Abstract……………………………………………….…....................58
7.2. Introduction…………………………………………...…..........……...59
7.3. Material and methods……………………………………...............61
7.4. Experiment 1- Image-to-image association vs.

ordinal position……………………………………………….....…….……......64
7.5. Experiment 2- Prospective vs. retrospective coding…...70
7.6. Experiment 3- Evaluating the possibility of tracking

responses made………………….…………………….........................75
7.7. Experiment 4- Evaluating the possibility of list-linking....79
7.8. Discussion…………………………………………..……….................85

8.Introduction to Paper 4……………………………………….….........96

Paper 4: Mental representation of order in non-human

primates…………………………………………………….……..............…..97
9.1. Abstract……………………………….………………...…........….....98
9.2. Three types of ordinal memories: common features

and differences..……………………………………………...................99
9.3. Evidence for common coding…………………...................102
9.4. Differences in the contents of memory during ordinal

judgments…….………………………………...…….…......................106
9.5. Implications for characterizations of memory types....108
9.6. Conclusions……………………………………...….……...............109

10.Conclusions and future directions………………….…….........111

11. References……………………………………………………...............116

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