<?xml version="1.0" encoding="UTF-8"?><?xml-stylesheet type="text/xsl" href="static/style.xsl"?><OAI-PMH xmlns="http://www.openarchives.org/OAI/2.0/" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xsi:schemaLocation="http://www.openarchives.org/OAI/2.0/ http://www.openarchives.org/OAI/2.0/OAI-PMH.xsd"><responseDate>2026-09-19T12:17:39Z</responseDate><request verb="GetRecord" identifier="oai:dspace.mit.edu:1721.1/138514" metadataPrefix="dim">https://dspace.mit.edu/server/oai/request</request><GetRecord><record><header><identifier>oai:dspace.mit.edu:1721.1/138514</identifier><datestamp>2026-06-16T18:55:04Z</datestamp><setSpec>com_1721.1_7582</setSpec><setSpec>com_1721.1_7581</setSpec><setSpec>col_1721.1_131022</setSpec></header><metadata><dim:dim xmlns:dim="http://www.dspace.org/xmlns/dspace/dim" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:doc="http://www.lyncode.com/xoai" xsi:schemaLocation="http://www.dspace.org/xmlns/dspace/dim http://www.dspace.org/schema/dim.xsd">
   <dim:field mdschema="dc" element="contributor" qualifier="advisor">Matthew A. Wilson.</dim:field>
   <dim:field mdschema="dc" element="contributor" qualifier="author" lang="en_US">Ma, Tzuhsuan.</dim:field>
   <dim:field mdschema="dc" element="contributor" qualifier="other" lang="en_US">Massachusetts Institute of Technology. Department of Brain and Cognitive Sciences.</dim:field>
   <dim:field mdschema="dc" element="contributor" qualifier="department" lang="en_US">Massachusetts Institute of Technology. Department of Brain and Cognitive Sciences</dim:field>
   <dim:field mdschema="dc" element="date" qualifier="accessioned">2021-12-17T17:04:27Z</dim:field>
   <dim:field mdschema="dc" element="date" qualifier="available">2021-12-17T17:04:27Z</dim:field>
   <dim:field mdschema="dc" element="date" qualifier="copyright" lang="en_US">2020</dim:field>
   <dim:field mdschema="dc" element="date" qualifier="issued" lang="en_US">2020</dim:field>
   <dim:field mdschema="dc" element="identifier" qualifier="uri">https://hdl.handle.net/1721.1/138514</dim:field>
   <dim:field mdschema="dc" element="identifier" qualifier="oclc" lang="en_US">1280701988</dim:field>
   <dim:field mdschema="dc" element="description" lang="en_US">Thesis: Ph. D., Massachusetts Institute of Technology, Department of Brain and Cognitive Sciences, February, 2020</dim:field>
   <dim:field mdschema="dc" element="description" lang="en_US">Manuscript.</dim:field>
   <dim:field mdschema="dc" element="description" lang="en_US">Includes bibliographical references (pages 227-238).</dim:field>
   <dim:field mdschema="dc" element="description" qualifier="abstract" lang="en_US">This work utilizes theoretical approaches to answer the question: which functions grid and place cells perform that directly lead to their own emergence? To answer such a question, an approach that goes beyond a simple modelling is necessary given the fact that there could be circuit solutions other than grid or place cells that better perform these functions. With this reasoning, I adopted a systematic guideline that aims for an optimization principle attempting to find the optimal solution for performing the hypothesized functions while reproducing the correct phenomenology. Within the optimization principle framework, I applied both recurrent neural network (RNN) training and coding-theoretic approaches to set up appropriate optimization problems for testing a given function hypotheses. The descriptive function hypotheses: 1) Grid cells exist for having a high-capacity and robust path-integrating code and 2) Place cells exist for having a sequentially-learnable and highly-separable path-integrating code were adopted. The non-converging performance in training an RNN to perform a hard navigation task suggests that the attractor dynamics forbids a network to simultaneously possess online learnability and high coding capacity. Because of this dynamical constraint in learning, a grid cell circuit has to be hardwired through some developmental process and cannot be easily modified by an experience-based synaptic rule without compromising its capacity. On the contrary, a place cell circuit being able to continually learn a novel environment inevitably have a mere linear capacity. These results imply that the functional separation of grid and place cell systems observed in the brain could be a result of an unavoidable dynamical constraint from their underlying RNNs. Lastly, a fundamental principle called the tuning-learnability correspondence was uncovered in pursuit of a sequentially learnable neural implementation for place cells. It explains that the seemingly incidental existence of conjunctive tuning property is in fact caused by a necessary metastable attractor dynamics for having sequential learnability rather than by another functional need attached to a particular tuning property. In addition, from the unique property of metastable attractor dynamics, I also predicted that the biased place field propensity recently observed in CA1 sub-region should originate from CA3 due to an inevitable biased activation in the RNN as a side effect of such a dynamical property. In sum, both this principle and the subsequent prediction thus provide a new perspective that contradicts the conventional wisdom which often assumed that a certain nonspatial tuning property exists for performing a relevant task.</dim:field>
   <dim:field mdschema="dc" element="description" qualifier="statementofresponsibility" lang="en_US">by Tzuhsuan Ma.</dim:field>
   <dim:field mdschema="dc" element="description" qualifier="degree" lang="en_US">Ph.D.</dim:field>
   <dim:field mdschema="dc" element="description" qualifier="collection" lang="en_US">Ph. D. Massachusetts Institute of Technology, Department of Brain and Cognitive Sciences</dim:field>
   <dim:field mdschema="dc" element="format" qualifier="extent" lang="en_US">238 pages</dim:field>
   <dim:field mdschema="dc" element="language" qualifier="iso" lang="en_US">eng</dim:field>
   <dim:field mdschema="dc" element="publisher" lang="en_US">Massachusetts Institute of Technology</dim:field>
   <dim:field mdschema="dc" element="rights" lang="en_US">MIT theses may be protected by copyright. Please reuse MIT thesis content according to the MIT Libraries Permissions Policy, which is available through the URL provided.</dim:field>
   <dim:field mdschema="dc" element="rights" qualifier="uri" lang="en_US">http://dspace.mit.edu/handle/1721.1/7582</dim:field>
   <dim:field mdschema="dc" element="subject" lang="en_US">Brain and Cognitive Sciences.</dim:field>
   <dim:field mdschema="dc" element="title" lang="en_US">Towards a theory for the emergence of grid and place cell codes</dim:field>
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   <dim:field mdschema="mit" element="thesis" qualifier="degree" lang="en_US">Doctoral</dim:field>
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   	&lt;Title>Towards a theory for the emergence of grid and place cell codes&lt;/Title>
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   	&lt;PublicationDate>2020&lt;/PublicationDate>
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        	&lt;DisplayName>Ma, Tzuhsuan.&lt;/DisplayName>
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    &lt;Keyword>Brain and Cognitive Sciences.&lt;/Keyword>
   	&lt;Abstract>This work utilizes theoretical approaches to answer the question: which functions grid and place cells perform that directly lead to their own emergence? To answer such a question, an approach that goes beyond a simple modelling is necessary given the fact that there could be circuit solutions other than grid or place cells that better perform these functions. With this reasoning, I adopted a systematic guideline that aims for an optimization principle attempting to find the optimal solution for performing the hypothesized functions while reproducing the correct phenomenology. Within the optimization principle framework, I applied both recurrent neural network (RNN) training and coding-theoretic approaches to set up appropriate optimization problems for testing a given function hypotheses. The descriptive function hypotheses: 1) Grid cells exist for having a high-capacity and robust path-integrating code and 2) Place cells exist for having a sequentially-learnable and highly-separable path-integrating code were adopted. The non-converging performance in training an RNN to perform a hard navigation task suggests that the attractor dynamics forbids a network to simultaneously possess online learnability and high coding capacity. Because of this dynamical constraint in learning, a grid cell circuit has to be hardwired through some developmental process and cannot be easily modified by an experience-based synaptic rule without compromising its capacity. On the contrary, a place cell circuit being able to continually learn a novel environment inevitably have a mere linear capacity. These results imply that the functional separation of grid and place cell systems observed in the brain could be a result of an unavoidable dynamical constraint from their underlying RNNs. Lastly, a fundamental principle called the tuning-learnability correspondence was uncovered in pursuit of a sequentially learnable neural implementation for place cells. It explains that the seemingly incidental existence of conjunctive tuning property is in fact caused by a necessary metastable attractor dynamics for having sequential learnability rather than by another functional need attached to a particular tuning property. In addition, from the unique property of metastable attractor dynamics, I also predicted that the biased place field propensity recently observed in CA1 sub-region should originate from CA3 due to an inevitable biased activation in the RNN as a side effect of such a dynamical property. In sum, both this principle and the subsequent prediction thus provide a new perspective that contradicts the conventional wisdom which often assumed that a certain nonspatial tuning property exists for performing a relevant task.&lt;/Abstract>
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