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	<title>Michael Halassa &#8211; Michael Halassa | Science</title>
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	<title>Michael Halassa &#8211; Michael Halassa | Science</title>
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		<title>The Quiet War Between Abstraction and Detail</title>
		<link>https://michaelhalassa.net/the-quiet-war-between-abstraction-and-detail/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Mon, 16 Mar 2026 22:31:52 +0000</pubDate>
				<category><![CDATA[Cognitive flexibility]]></category>
		<category><![CDATA[Cognitive Processing]]></category>
		<category><![CDATA[Computational neuroscience]]></category>
		<category><![CDATA[Halassa Lab]]></category>
		<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neural circuits]]></category>
		<category><![CDATA[NeuroAI]]></category>
		<category><![CDATA[brain]]></category>
		<category><![CDATA[cognitive flexibility]]></category>
		<category><![CDATA[Cognitive Science]]></category>
		<category><![CDATA[neurons]]></category>
		<category><![CDATA[RNN]]></category>
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					<description><![CDATA[https://michaelhalassa.substack.com/p/the-quiet-war-between-abstraction Every time you walk into a new restaurant, your brain solves an invisible problem: Which parts of this experience are specific to this place (the menu, the layout, the staff) and which parts are general rules you can transfer (how to order, where to sit, when to pay)? Extract too much shared structure and you’ll confidently [&#8230;]]]></description>
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<p>Every time you walk into a new restaurant, your brain solves an invisible problem: Which parts of this experience are specific to <em>this place</em> (the menu, the layout, the staff) and which parts are general rules you can transfer (how to order, where to sit, when to pay)? Extract too much shared structure and you’ll confidently walk to the wrong counter. Protect too many specific details and you’ll fumble through every new restaurant like it’s your first.</p>
<p>This balance, between learning patterns that transfer and preserving details that matter, appears fundamental to how we navigate a world where some things repeat (traffic patterns, social norms, the physics of thrown objects) and some things are unique (this particular intersection floods, this friend needs space when upset, this specific mushroom will kill you).</p>
<p>The capacity for structural learning, extracting regularities that apply across situations, may be one of the defining features of human cognition. When a child learns that adding “ed” creates past tense, they’ve discovered a rule that applies to thousands of verbs they’ve never encountered. When you recognize that a new coworker uses the same conflict-avoidant communication style as your sibling, you’ve extracted a pattern that predicts future interactions. The alternative, learning every situation as a unique instance requiring its own solution, would be computationally intractable.</p>
<p>But structural learning creates a fundamental tension. The same cognitive machinery that lets you rapidly transfer knowledge to new situations may overwrite what you learned before. Learn French after Spanish and you might start mixing verb conjugations. Update your mental model of how your boss communicates and you might misremember what they actually said last month. The brain must somehow balance the benefits of generalization against the risk of interference.</p>
<h2 class="header-anchor-post">A Task That Makes Strategies Visible</h2>
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<p>&nbsp;</p>
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<p>A new paper in <a href="https://www.nature.com/articles/s41562-025-02318-y" rel="noopener" target="_blank">Nature Human Behaviour</a> by Eleanor Holton, Chris Summerfield, and colleagues has developed an elegant way to observe these competing strategies in action. The design is deceptively simple: participants learned to map fictional plants to locations on a circular planet, with separate locations for summer and winter. The key structural feature is that within each set of plants, there was a consistent angular rule. Winter locations were always a fixed offset from summer locations (say, 120 degrees clockwise).</p>
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<picture><source srcset="https://substackcdn.com/image/fetch/$s_!CR4L!,w_424,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 424w, https://substackcdn.com/image/fetch/$s_!CR4L!,w_848,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 848w, https://substackcdn.com/image/fetch/$s_!CR4L!,w_1272,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 1272w, https://substackcdn.com/image/fetch/$s_!CR4L!,w_1456,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 1456w" type="image/webp" sizes="100vw" /><img loading="lazy" decoding="async" class="sizing-normal" src="https://substackcdn.com/image/fetch/$s_!CR4L!,w_1456,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png" sizes="100vw" srcset="https://substackcdn.com/image/fetch/$s_!CR4L!,w_424,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 424w, https://substackcdn.com/image/fetch/$s_!CR4L!,w_848,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 848w, https://substackcdn.com/image/fetch/$s_!CR4L!,w_1272,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 1272w, https://substackcdn.com/image/fetch/$s_!CR4L!,w_1456,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png 1456w" alt="https%3A%2F%2Fsubstack post media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d 736c 4181 895f" width="764" height="380" data-attrs="{&quot;src&quot;:&quot;https://substack-post-media.s3.amazonaws.com/public/images/18f7458d-736c-4181-895f-27288dc60711_764x380.png&quot;,&quot;srcNoWatermark&quot;:null,&quot;fullscreen&quot;:null,&quot;imageSize&quot;:null,&quot;height&quot;:380,&quot;width&quot;:764,&quot;resizeWidth&quot;:null,&quot;bytes&quot;:94963,&quot;alt&quot;:null,&quot;title&quot;:null,&quot;type&quot;:&quot;image/png&quot;,&quot;href&quot;:null,&quot;belowTheFold&quot;:false,&quot;topImage&quot;:false,&quot;internalRedirect&quot;:&quot;https://michaelhalassa.substack.com/i/178980901?img=https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F18f7458d-736c-4181-895f-27288dc60711_764x380.png&quot;,&quot;isProcessing&quot;:false,&quot;align&quot;:null,&quot;offset&quot;:false}" title="The Quiet War Between Abstraction and Detail 8"></picture>
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<p>&nbsp;</p>
<p>Participants first learned one set of six plants (Task A) over ten repetitions.</p>
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<picture><source srcset="https://substackcdn.com/image/fetch/$s_!EscE!,w_424,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 424w, https://substackcdn.com/image/fetch/$s_!EscE!,w_848,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 848w, https://substackcdn.com/image/fetch/$s_!EscE!,w_1272,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 1272w, https://substackcdn.com/image/fetch/$s_!EscE!,w_1456,c_limit,f_webp,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 1456w" type="image/webp" sizes="100vw" /><img loading="lazy" decoding="async" class="sizing-normal" src="https://substackcdn.com/image/fetch/$s_!EscE!,w_1456,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png" sizes="100vw" srcset="https://substackcdn.com/image/fetch/$s_!EscE!,w_424,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 424w, https://substackcdn.com/image/fetch/$s_!EscE!,w_848,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 848w, https://substackcdn.com/image/fetch/$s_!EscE!,w_1272,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 1272w, https://substackcdn.com/image/fetch/$s_!EscE!,w_1456,c_limit,f_auto,q_auto:good,fl_progressive:steep/https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png 1456w" alt="https%3A%2F%2Fsubstack post media.s3.amazonaws.com%2Fpublic%2Fimages%2F3cf15183 5eee 48d0 9bf5" width="409" height="402" data-attrs="{&quot;src&quot;:&quot;https://substack-post-media.s3.amazonaws.com/public/images/3cf15183-5eee-48d0-9bf5-2f72fe67394a_409x402.png&quot;,&quot;srcNoWatermark&quot;:null,&quot;fullscreen&quot;:null,&quot;imageSize&quot;:null,&quot;height&quot;:402,&quot;width&quot;:409,&quot;resizeWidth&quot;:null,&quot;bytes&quot;:108293,&quot;alt&quot;:null,&quot;title&quot;:null,&quot;type&quot;:&quot;image/png&quot;,&quot;href&quot;:null,&quot;belowTheFold&quot;:true,&quot;topImage&quot;:false,&quot;internalRedirect&quot;:&quot;https://michaelhalassa.substack.com/i/178980901?img=https%3A%2F%2Fsubstack-post-media.s3.amazonaws.com%2Fpublic%2Fimages%2F5fbfc8db-7cb3-4cde-893b-23dd4939a7c8_410x415.png&quot;,&quot;isProcessing&quot;:false,&quot;align&quot;:null,&quot;offset&quot;:false}" title="The Quiet War Between Abstraction and Detail 9"></picture>
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<p>&nbsp;</p>
<p>Then they encountered six entirely new plants (Task B) with their own summer-winter rule, either identical to Task A, shifted by 30 degrees, or rotated 180 degrees.</p>
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<p>&nbsp;</p>
<p>After learning Task B, they returned to the original Task A plants, but this time received feedback only for summer locations. This allowed the team to observe what rule participants spontaneously applied when retested.</p>
<p>The experimental design cleverly separates two phenomena that usually travel together. Transfer refers to how much knowledge of rule A accelerates learning of rule B. If you immediately apply the 120-degree rule to new plants, your initial performance on Task B should be good (when the rule is similar) or systematically biased (when the rule differs). Interference refers to whether learning rule B corrupts your memory of rule A. When retested on the original plants, do you still apply rule A correctly, or do you now mistakenly use rule B?</p>
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<p>&nbsp;</p>
<p>In most learning paradigms, these processes are confounded. Here, the structure makes different algorithmic strategies visible through their distinct signatures across transfer and interference conditions.</p>
<h2 class="header-anchor-post">Heterogeneity Hidden Beneath Averaged Performance</h2>
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<p>There are many interesting things about this paper (and some things that require taking with a grain of salt that I will point out in due course), but one big takeaway is how well individual differences in strategy choice was revealed by this work.</p>
<p>If you average across all participants (which many people routinely do), behavior would almost certainly appear to be relatively uniform. This is because people learn both tasks to high accuracy. Mean performance in Task B appears similar across individuals.</p>
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<p>But beneath this averaged regularity lies what appears to be substantial heterogeneity in <em>how</em> people solve the task. This becomes most visible when the rules are similar enough to create genuine ambiguity (the 30-degree shift condition, though patterns may exist in the far condition too). The Near condition participants seem to split into two distinct behavioral profiles:</p>
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<p>&nbsp;</p>
<p>Some participants (called “lumpers”) showed high transfer but high interference. They immediately benefited from rule A when learning rule B, suggesting they were applying the previous rule to new stimuli. But when retested on task A, they were more likely to apply rule B, apparently overwriting their memory of the original rule.</p>
<p>Other participants (“splitters”) showed the opposite pattern: low transfer but low interference. They treated task B as effectively novel, gaining little immediate advantage from prior learning, and made many more mistakes upon transitioning (higher switching cost). But they maintained rule A better at retest, showing no contamination from the recently learned rule B.</p>
<p>This dissociation extended across multiple behavioral measures beyond the primary transfer and interference metrics. Lumpers appeared to generalize the rule better to held-out stimuli they’d never received feedback on, applying the angular relationship to infer correct responses. But they performed worse on memory for unique stimulus features, specifically the summer locations that had to be memorized rather than inferred from a rule. Splitters showed what looks like the mirror pattern: better memory for unique features, poorer rule-based generalization.</p>
<p>The groups even differed on explicit temporal memory tested at the end of the experiment. Splitters were better at reporting which half of the study they’d first encountered each stimulus in, as if they maintained more distinct representations of the two task contexts.</p>
<p>These differences appear to reflect distinct computational strategies rather than differences in overall ability. Splitters actually outperformed lumpers on some measures (the unique feature memory). Both groups achieved high final accuracy. These look like fundamentally different approaches to solving the same problem, each with complementary strengths and opposite vulnerability profiles.</p>
<p>An important unresolved question is whether these represent stable individual traits or context-dependent strategies. Does someone who lumps in this task lump everywhere? Or do people flexibly shift between strategies based on task structure, recent experience, or environmental statistics? The study design can’t answer this, and it’s a genuinely significant open issue for understanding what these behavioral patterns mean.</p>
<h2 class="header-anchor-post">Formalizing the Trade-off Through Neural Networks</h2>
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<p>&nbsp;</p>
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<p>One of the interesting aspects of this work (and <a href="https://www.psy.ox.ac.uk/people/christopher-summerfield" rel="noopener" target="_blank">Summerfield’s work</a> more generally) is the use of AI architectures to gain insight into the human mind (and sometimes the brain too).</p>
<p>In this study, the authors used a surprisingly simple neural architecture, a two-layered feedforward network to make some inferences about individual differences in cognitive strategy usage. By manipulating the scale of initial weights, the authors could guide networks toward “rich” (small initial weights leading to structured, low-dimensional representations) or “lazy” (large initial weights leading to high-dimensional, task-agnostic representations) learning regimes.</p>
<p>Networks in the rich regime captured what looks like the lumper behavioral profile: high transfer to Task B, high interference when retested on Task A, strong rule generalization to held-out stimuli, poor unique feature memory, and collapsed representations of the two tasks in neural space (measured via principal angles between task subspaces).</p>
<p>Networks in the lazy regime mirrored what appears to be the splitter pattern on every measure: low transfer, low interference, poor generalization, better unique feature memory, and maintenance of orthogonal representations for the two tasks.</p>
<p>This computational modeling makes the underlying trade-off explicit. Low-dimensional structured representations might enable efficient generalization by extracting shared rules, but precisely because representations are shared across tasks, new learning could overwrite old knowledge. High-dimensional representations might maintain separability between tasks, protecting against interference, but at the potential cost of failing to extract transferable structure.</p>
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<p>The value of this modeling approach is that it formalizes what could otherwise remain a vague intuition about competing strategies. The networks make testable predictions: if someone shows high interference, they should also show better generalization within tasks. If they maintain orthogonal task representations, they should show poor transfer but preserved memory.</p>
<p>But we need to be careful about how far we extend the analogy. Having interacted with many humans, I can confirm that none are a two-layered neural networks trained via gradient descent on a single task. How people implement rapid learning likely involves multiple interacting brain systems: prefrontal cortex implementing gating policies that determine when to update versus maintain representations, hippocampus providing rapid contextualization that could separate task episodes, thalamocortical circuits routing information through different processing channels based on task demands. Our behavior, as individuals, probably emerges from how each of our systems interact and are weighted together, something like a mixture of experts where different computational solutions contribute to the final output. Also, the rich and lazy networks are unlikely to describe individual neural systems wholesale, instead, they demonstrate a computational principle that illustrate the trade-off at work (although who knows, maybe each of our brains has a mixture of these principles at work).</p>
<p>The neural network analysis also reveals something interesting about task similarity and strategy visibility. In the Same condition (where both tasks use identical rules), the distinction between lumpers and splitters may not matter much. Everyone can apply the same rule to new stimuli without cost. In the Far condition (180-degree shift), the rules are different enough that most people might naturally treat them as separate tasks, though there may still be some heterogeneity that’s less pronounced. It’s in the Near condition (30-degree shift) where the ambiguity forces different strategies into stark relief, creating the bimodal distribution.</p>
<p>This suggests that algorithmic heterogeneity in how people approach learning might be widespread but often invisible, only becoming apparent when task structure creates the right conditions to pull different strategies apart.</p>
<h2 class="header-anchor-post">Environmental Contingency and the Absence of a Single Optimal Strategy</h2>
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<p>&nbsp;</p>
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<p>The environmental context matters critically for which strategy succeeds. Lumping might dominate when the world has stable structure worth extracting. If rules genuinely repeat across contexts, you learn faster by recognizing and applying patterns. The cost of occasional interference could be outweighed by the acceleration in learning new tasks that share structure with old ones.</p>
<p>Splitting might win when rules change frequently or when maintaining distinct memories is crucial. If what you learned before is often misleading rather than helpful, protecting each memory from contamination becomes more valuable than speed of transfer. The cost of slow learning could be justified by the accuracy of what you retain.</p>
<p>There may be no single “correct” strategy. The optimal approach likely depends on the statistics of the environment you’re navigating. A world with high regularity and stable rules rewards generalization. A world with frequent rule changes or high context-specificity rewards separation and protection of distinct memories. There is also the possibility that luck plays a role; what you just encountered may predispose you to lump or split depending on how successful you just were.</p>
<p>This raises an interesting possibility about cognitive diversity. Rather than representing noise around some ideal cognitive architecture, the coexistence of lumpers and splitters might reflect adaptation to environmental variability. A population containing both strategies might outperform a homogeneous population, with pattern-extractors thriving in stable domains and detail-preservers succeeding in volatile ones. Different algorithmic profiles suited to different ecological niches.</p>
<p>This interpretation is speculative, but it suggests we might want to think about individual differences in learning strategies as positions on a trade-off curve that evolution or development has explored.</p>
<h2 class="header-anchor-post">Relevance for Understanding Disorders</h2>
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<p>This level of behavioral dissection seems important for understanding psychiatric heterogeneity. If we only measured final task accuracy, lumpers and splitters would be indistinguishable. Standard neuropsychological testing focused on whether performance is intact or impaired would miss this entirely. It’s only by examining the <em>pattern</em> of performance across multiple measures (transfer, interference, generalization, unique feature memory, temporal context) that the different strategies become visible.</p>
<p>This has potentially important implications for how we study psychiatric conditions. Consider what we know about cognitive function in schizophrenia. Working memory deficits are consistently documented. Counterfactual reasoning appears impaired. Context processing shows abnormalities. But we typically describe these as simple deficits, performance falling below some normative threshold.</p>
<p>What if some of this heterogeneity reflects people navigating trade-offs differently, perhaps forced toward one extreme by underlying capacity constraints? Someone with severe working memory limitations might be pushed toward splitting strategies, unable to maintain the flexible representations needed for successful generalization. Alternatively, they might be pushed toward excessive lumping, overgeneralizing because they can’t maintain distinct context representations. Or the computational machinery for balancing these strategies might be disrupted in ways that don’t map onto the healthy spectrum at all.</p>
<p>Without tasks that can behaviorally dissect these possibilities, separating transfer from interference, generalization from discrimination, rule application from memory for specifics, we can’t distinguish these accounts. We end up with general statements about “cognitive deficits” when we should be asking about specific algorithmic profiles and how they interact with task demands.</p>
<p>The implications here concern the level of analysis we need. Detailed statistics of behavior, comparison to normative computational models when available, careful dissection of performance patterns across conditions that pull different strategies apart.</p>
<p>And we should remember the limitations of the analogy: people aren’t one big neural network. Multiple brain systems (prefrontal gating, hippocampal context coding, thalamic routing) likely contribute to how we handle sequential learning. These computational principles might illuminate trade-offs and formalize what behavioral patterns we should look for, but the implementation almost certainly involves coordination across systems rather than a single learning mechanism. The behavioral signature we observe is likely the weighted output of this complex architecture.</p>
<h2 class="header-anchor-post">Emphasis on Methodological Insight</h2>
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<p>The methodological contribution here may be as important as the empirical findings. Averaged behavior can look normal, even optimal, while masking fundamentally different computational strategies operating beneath the surface. Standard cognitive assessments can show intact performance while missing the algorithmic heterogeneity that might matter for understanding vulnerability, predicting treatment response, or matching individuals to environments.</p>
<p>The sequential learning paradigm with separate measures of transfer and interference provides one example of how to pull these strategies apart. The neural network modeling provides a formal framework for understanding what different patterns of performance might mean. The combination suggests a path forward for computational psychiatry that goes beyond asking whether performance is impaired.</p>
<p>The next generation of this work might ask: Which specific algorithmic trade-offs are being navigated differently? What behavioral signatures reveal underlying strategy? How do these computational profiles interact with environmental demands? When does being a lumper become maladaptive, and when does being a splitter limit learning?</p>
<p>The demonstration that careful behavioral dissection can reveal hidden heterogeneity in how people learn suggests we might be missing similar structure in other domains by averaging too quickly and testing too coarsely.</p>
<h2 class="header-anchor-post">Synthesis: What We Learn About Learning</h2>
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<p>The capacity to abstract while preserving details represents a fundamental computational challenge for any learning system, biological or artificial. The tension appears intrinsic to intelligence itself.</p>
<p>What optimal looks like depends entirely on environmental structure. Stable worlds with repeating patterns reward those who extract and apply rules quickly. Volatile worlds where yesterday’s pattern misleads today reward those who maintain distinct memories and avoid overgeneralization. The problem itself changes based on context, making any single solution inadequate.</p>
<p>This has profound implications for how we think about cognitive diversity. Individual differences in learning strategies might represent different positions on a fundamental trade-off curve, possibly shaped by recent experience, current capacity constraints, or the statistics of environments people have navigated. Cognitive styles might reflect computational strategies. Apparent deficits might be extreme positions on trade-offs that have no objectively correct answer.</p>
<p>The biological implementation is almost certainly a weighted mixture of multiple brain systems coordinating to produce the behavior we observe. Disruption might affect these systems differently, push trade-offs toward extremes, or create computational patterns that don’t exist in healthy populations at all.</p>
<p>The brain’s solution to the quiet war between abstraction and detail appears to be “it depends on the brain and it depends on the world.” Understanding both dependencies seems necessary for making sense of how learning works, why it sometimes fails, and what interventions might actually help.</p>
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<p><em>Work like this only matters if it reaches people wondering about these questions. If you found value here, consider sharing it with your community. Subscribe to michaelhalassa.substack.com if you want to see where this type of thinking and analysis goes.</em></p>
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		<title>The Currency of the Mind</title>
		<link>https://michaelhalassa.net/the-currency-of-the-mind/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Mon, 16 Mar 2026 22:22:43 +0000</pubDate>
				<category><![CDATA[Cognitive flexibility]]></category>
		<category><![CDATA[Cognitive Processing]]></category>
		<category><![CDATA[Computational neuroscience]]></category>
		<category><![CDATA[Halassa Lab]]></category>
		<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neural circuits]]></category>
		<category><![CDATA[cingulate cortex]]></category>
		<category><![CDATA[cognitive flexibility]]></category>
		<category><![CDATA[cognitive processing]]></category>
		<category><![CDATA[orbitofrontal cortex]]></category>
		<category><![CDATA[value computation]]></category>
		<guid isPermaLink="false">https://michaelhalassa.net/?p=794</guid>

					<description><![CDATA[How the brain constructs the values that guide everyday decisions reveals one of neuroscience’s most fascinating puzzles. Think about it: your brain adds and subtracts quantities that share no common unit! It can add morning light through kitchen windows to forty minutes in traffic, subtract image and status from a car’s reliability and comfort. These [&#8230;]]]></description>
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<p>How the brain constructs the values that guide everyday decisions reveals one of neuroscience’s most fascinating puzzles. Think about it: your brain adds and subtracts quantities that share no common unit! It can add morning light through kitchen windows to forty minutes in traffic, subtract image and status from a car’s reliability and comfort. These things exist in completely different dimensions (light, time, dollars, social signals) yet somehow the brain is constantly adding and subtracting them when making decisions.</p>
<p>Let’s take buying a home as a concrete example. On paper it looks like a financial transaction, but in practice it’s a clash of incomparable currencies. Square footage gets weighed against school districts, the energy of a neighborhood against the stability of an investment. Walk through one house and you can already imagine your life there, until you realize it means your partner endures an extra hour of daily commute. Somewhere in this mix of clear measurements and ones that are hard to describe, the brain assembles a decision.</p>
<p>To make matters more complicated, think about how volatile our internal value estimates can be. During COVID, when daily commutes vanished, the value of space ballooned, potentially trumping distance; the same house that once felt impractical now seemed like a refuge. A new context can make ostensibly identical attributes exhibit radically different valuations.</p>
<p>The types of contextual changes that shift valuation are also themselves diverse. A genuine Monet may sell for $70 million. A forgery, indistinguishable to the eye and identical to anyone but the equipment of an art authenticator, might fetch $5,000. Same paint, same canvas, same aesthetic experience. Yet the (inferred) backstory behind it transforms its value by four orders of magnitude.</p>
<p>This is the computational puzzle at the heart of value-based decision-making: how does the brain make incomparable things comparable? What neural mechanism allows attributes measured in completely different dimensions (light, time, dollars, authenticity) to compete on the same playing field? And how does this mechanism remain stable enough to produce coherent choices yet flexible enough to radically reweight those same attributes when context shifts?</p>
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<p><em>From the Wall Street Journal. <a href="https://www.wsj.com/arts-culture/fine-art/a-claude-monet-water-lilies-scene-sold-for-65-5-million-6af15ce4" rel="noopener" target="_blank">Read the story here</a></em></p>
<h2 class="header-anchor-post"><strong>The Construction of Value</strong></h2>
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<p>Here’s the truth: we don’t really understand how this problem is solved. The question of how brains integrate diverse attributes into unified decisions remains one of the deep mysteries in neuroscience. Ultimately, the answer will likely come in cognitive and neural forms; two faces of the same computational coin. These perspectives will jointly explain the magic of turning morning light into a quantity that can be compared with commute time.</p>
<p>Despite the lack of a satisfying single narrative, we do know some fascinating pieces of the puzzle. Let’s start with what psychology has revealed about how value gets constructed.</p>
<p>One of the most striking findings is that arbitrary starting points can anchor our entire valuation system. Dan Ariely and colleagues demonstrated this by asking students to write down the last two digits of their Social Security number before bidding on items like wine, chocolate, and computer accessories. Students with Social Security numbers ending in 80-99 bid nearly three times more than those with numbers ending in 00-19. For a cordless keyboard, high-number students offered $56 while low-number students offered just $16. The same pattern held across all items. The initial number, though completely unrelated to the products’ worth, set an implicit scale that influenced all subsequent valuations. Once the brain latches onto a reference point, even a meaningless one, it builds an internally consistent preference structure around it.</p>
<p>Beyond arbitrary anchors, our sense of ownership profoundly alters how we value objects. In experiments by Kahneman, Knetsch, and Thaler, students were randomly given coffee mugs and then asked to name their selling price. These new “owners” demanded about $7 to part with their mugs, while students without mugs were only willing to pay about $3 to acquire one. The mug itself hadn’t changed. What changed was the relationship: once something becomes “mine,” its value doubles in my eyes. Norton, Mochon, and Ariely extended this finding by having people assemble IKEA furniture or fold origami cranes. Participants valued their own creations at nearly the same price as expert-made versions, even when their handiwork was visibly inferior. The act of creation adds a new attribute to the value calculation: the effort invested becomes part of what we’re evaluating, not just the object itself.</p>
<p>Even memory rewrites value. Daniel Kahneman and Donald Redelmeier studied patients undergoing colonoscopies. Some patients had longer procedures that ended less painfully, others had shorter ones that ended abruptly at peak pain. Counterintuitively, patients preferred the longer procedures. Their memories followed the “peak-end rule”: they judged the whole experience not by its average pain but by its worst moment and how it ended (Redelmeier &amp; Kahneman, 1996). How we remember an experience, not the experience itself, determines how we’ll value similar choices in the future.</p>
<p>And value is deeply social. In a massive online experiment, Matthew Salganik and colleagues (2006) created artificial “music markets.” When download counts were hidden, songs rose or fell on their own. But when popularity information was visible, some songs snowballed into “hits” while others languished, even though the songs were the same across markets. Popularity itself became an attribute folded into value, warping what people genuinely preferred.</p>
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<p><em><a href="https://en.wikipedia.org/wiki/Daniel_Kahneman" rel="noopener" target="_blank">Daniel Kahneman</a>: A most incredible thinker and contributor to the science of decision making (also a Nobel Laureate)</em></p>
<h2 class="header-anchor-post"><strong>The Computational Challenge</strong></h2>
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<p>If value is constructed rather than retrieved from a fixed look-up table, what exactly is the brain computing? Consider what components must somehow become commensurable:</p>
<p><strong>Sensory attributes</strong>: The warmth of sunlight, the bitterness of coffee, the smoothness of silk. These arrive in different neural codes from different sensory systems, yet must be integrated into unified preferences.</p>
<p><strong>Abstract properties</strong>: Distance (20 minutes), quantity (800 square feet), probability (70% chance). The brain lacks sensory receptors for these dimensions, yet they powerfully shape value.</p>
<p><strong>Social signals</strong>: Status, belonging, reputation. That Monet carries social meaning that a forgery doesn’t. A Harvard degree signals something another school may not. These intangible attributes somehow get converted into the same currency as tangible ones.</p>
<p><strong>Temporal projections</strong>: Future pleasure, anticipated regret, imagined satisfaction. The brain must evaluate things that haven’t happened yet, experiences it can only simulate.</p>
<p><strong>Effort and ownership</strong>: The IKEA table you assembled, the garden you planted, the thesis you wrote. Investment of effort literally changes the computed value, as if the brain adds your labor to the object’s attributes.</p>
<p><strong>Comparison context</strong>: The same option valued differently depending on what else is available. That $2,500 apartment seems expensive or cheap depending entirely on the alternatives, even irrelevant ones.</p>
<p>The remarkable thing is that the brain somehow integrates these components despite their fundamental incomparability. One prominent theory suggests the brain converts everything into a “common currency”; perhaps the firing rates (or patterns) of neurons in valuation regions. But how does social status get converted into the same neural code as commute time? What algorithm transforms the warmth of sunlight into the same units as financial security? Even if there is a common currency at the point of decision, the translation process remains mysterious. When you’re choosing between jobs or homes or life partners, all these incomparable attributes must somehow become comparable. One option just feels better.</p>
<p>How does the brain perform this translation and integration? That’s what the neural machinery must somehow accomplish.</p>
<h2 class="header-anchor-post"><strong>The Neural Implementation</strong></h2>
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<p>If value is constructed, where in the brain does this happen, and in what format? Researchers often divide into two camps. One view is that the brain collapses everything into a single “common currency,” a scalar signal that can be compared across apples, Monets, and commutes. The other is that value is represented in a richer, multidimensional code, more like a map of attributes than a single number, with scalar readouts emerging only when a choice is required. The best available evidence points to the orbitofrontal cortex (OFC) and the adjacent ventromedial prefrontal cortex (vmPFC) as being central to these computations.</p>
<h3 class="header-anchor-post"><strong>OFC and vmPFC: a value map with scalar readouts</strong></h3>
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<p>Work in macaques by Camillo Padoa-Schioppa and colleagues showed that OFC neurons encode subjective value in a way that is “menu invariant,” meaning the value signal for one option stays stable regardless of what it is paired against (Padoa-Schioppa &amp; Assad, 2006). This stability supports transitive choice: if juice A is valued more than B, and B more than C, then A will be valued more than C. Human neuroimaging extends this by showing vmPFC activity tracks subjective value across many domains, including money, food, and social approval (Chib et al., 2009; Bartra et al., 2013).</p>
<p>However, newer analyses suggest the OFC does not simply produce one number. Instead, its population activity preserves multiple dimensions of value, such as taste versus health or probability versus magnitude (Schuck et al., 2016; Hunt &amp; Hayden, 2017). In this view, the OFC is a “map-maker,” maintaining a structured representation of options that can be flexibly reweighted depending on context. Scalar value signals may still emerge, but only as a projection of this richer map.</p>
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<p><em>Figure 1 of <a href="https://www.cell.com/trends/neurosciences/fulltext/S0166-2236(24)00202-9#f0005" rel="noopener" target="_blank">Moneta et al., 2024 Trends in Neuroscience</a></em>.</p>
<h3 class="header-anchor-post"><strong>The role of dorsal prefrontal regions: shaping and acting on value</strong></h3>
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<p>Other prefrontal areas contribute in complementary ways. The dorsolateral prefrontal cortex (dlPFC), which is heavily involved in executive control, appears to adjust the weights assigned to different attributes. When people are instructed to prioritize health over taste, dlPFC activity reflects health information more strongly, and when told to focus on taste it reflects taste (Hare et al., 2009). Under certain conditions, the dlPFC may determine which dimensions of the map matter in a given context.</p>
<p>The dorsal anterior cingulate cortex (dACC), which monitors conflict and effort, often signals the difficulty or cost of a decision. Because it connects closely to premotor regions, it is well placed to bind abstract values to concrete actions. In challenging or effortful choices, dACC appears to integrate both the value of options and the anticipated cost of exerting control (Shenhav et al., 2013, <em>Neuron</em>).</p>
<h3 class="header-anchor-post"><strong>A distributed system</strong></h3>
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<p>The emerging picture is that value does not reside in a single place or a single format. The OFC and vmPFC maintain a flexible, map-like code of options. dlPFC helps determine which axes of that map to emphasize. dACC translates the chosen value into action, especially when the choice is close or costly. Striatal circuits and dopamine signals supply the learning machinery that updates the map when outcomes deviate from expectations.</p>
<p>Understanding this distributed system may ultimately reconcile the debate between scalar and map-like coding. The brain can preserve a rich geometry of attributes while also collapsing them into a scalar readout when a choice demands it. That dual capacity may be the key to how incomparable things become comparable.</p>
<h2 class="header-anchor-post"><strong>Broader Implications</strong></h2>
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<p>Understanding value as construction rather than retrieval has implications beyond individual choice. It might explain why the same economic conditions feel catastrophic or manageable depending on narrative framing. Why social media can shift entire populations’ valuation of political candidates through selective attribute highlighting. Why depression involves not just sadness but a fundamental inability to construct positive value from available attributes. Why cultural differences in what matters, individual achievement versus group harmony, lead to genuinely different experiences of the same situations.</p>
<p>The framework suggests that many societal conflicts aren’t really about different goals but about different attribute weightings. The same policy gets valued completely differently depending on whether you weigh “personal freedom” or “collective safety” more heavily. The same scientific finding gets valued differently depending on whether you weigh “economic growth” or “environmental protection.” Instead of viewing these as failures of rationality, an opposing view can be simply a different construction stemming from different weightings.</p>
<h2 class="header-anchor-post"><strong>The Mystery of Value</strong></h2>
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<p>We started with a puzzle: how does your brain weigh the comfort of a home’s interior against commute distance, nearby amenities against your spouse’s preferences? How do incomparable attributes become comparable values?</p>
<p>The evidence points toward construction rather than retrieval from a look-up table. Ariely’s anchoring studies show that random numbers shape our valuations. The endowment effect reveals that ownership doubles perceived worth. The peak-end rule demonstrates that certain details of memory encoding determine future value. Eye-tracking shows that attention creates preference. The Monet example shows that authentication can change value by four orders of magnitude.</p>
<p>These phenomena make sense if the brain builds value from available attributes, weights them according to context and goals, and compares through some form of competition. The neural data provides pieces: OFC/vmPFC may house a flexible map-like code that may be collapsed to a common currency scalar value for comparison. dlPFC circuitry may shape which axes of that map matter and dACC circuitry may read out the winning option in preparation for action.</p>
<p>But the core mystery remains. How does the brain actually perform the integration? What computation transforms sunshine through kitchen windows into a quantity that can be weighed against minutes of commute? How do narrative attributes like “painted by Monet himself” get converted into the same currency as visual beauty or investment potential?</p>
<p>The $70 million Monet shows how backstory can outweigh brushstrokes. What we don’t yet know is how the brain pulls off the trick of weighing sunlight against commute time, or authenticity against aesthetics. That algorithm remains one of neuroscience’s deepest mysteries.</p>
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<p><em>If you enjoyed this piece, please consider subscribing to michaelhalassa.substack.com to follow along as I write about the brain, computation, and psychiatry. Some posts dive into the neuroscience of a particular mental phenomenon (like this one), while others deal with more clinically-relevant issues.</em></p>
<p><em>You can also share this post with a friend or colleague who might be curious about how our brains turn sunlight, stories, and symbols into value.</em></p>
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<p>Bibliography:</p>
<p>Ariely, D., Loewenstein, G., &amp; Prelec, D. (2003). <em>“Coherent arbitrariness”: Stable demand curves without stable preferences.</em> Quarterly Journal of Economics, 118(1), 73–106. https://doi.org/10.1162/00335530360535153</p>
<p>Bartra, O., McGuire, J. T., &amp; Kable, J. W. (2013). <em>The valuation system: A coordinate-based meta-analysis of BOLD fMRI experiments examining neural correlates of subjective value.</em> NeuroImage, 76, 412–427. https://doi.org/10.1016/j.neuroimage.2013.02.063</p>
<p>Chib, V. S., Rangel, A., Shimojo, S., &amp; O’Doherty, J. P. (2009). <em>Evidence for a common representation of decision values for dissimilar goods in human ventromedial prefrontal cortex.</em> Journal of Neuroscience, 29(39), 12315–12320. https://doi.org/10.1523/JNEUROSCI.2575-09.2009</p>
<p>Hare, T. A., Camerer, C. F., &amp; Rangel, A. (2009). <em>Self-control in decision-making involves modulation of the vmPFC valuation system.</em> Science, 324(5927), 646–648. https://doi.org/10.1126/science.1168450</p>
<p>Hunt, L. T., &amp; Hayden, B. Y. (2017). <em>A distributed, hierarchical and recurrent framework for reward-based choice.</em> Neuron, 96(2), 355–362. https://doi.org/10.1016/j.neuron.2017.09.031</p>
<p>Kahneman, D., Knetsch, J. L., &amp; Thaler, R. H. (1991). <em>Anomalies: The endowment effect, loss aversion, and status quo bias.</em> Journal of Economic Perspectives, 5(1), 193–206. https://doi.org/10.1257/jep.5.1.193</p>
<p>Kaplan, R., &amp; Friston, K. J. (2018). <em>Planning and navigation as active inference.</em> Biological Cybernetics, 112(4), 323–343. https://doi.org/10.1007/s00422-018-0753-2</p>
<p>Norton, M. I., Mochon, D., &amp; Ariely, D. (2012). <em>The IKEA effect: When labor leads to love.</em> Journal of Consumer Psychology, 22(3), 453–460. https://doi.org/10.1016/j.jcps.2011.08.002</p>
<p>Padoa-Schioppa, C., &amp; Assad, J. A. (2006). <em>Neurons in the orbitofrontal cortex encode economic value.</em> Nature Neuroscience, 9(3), 367–373. https://doi.org/10.1038/nn1726</p>
<p>Redelmeier, D. A., &amp; Kahneman, D. (1996). <em>Patients’ memories of painful medical treatments: Real-time and retrospective evaluations of two minimally invasive procedures.</em> Pain, 66(1), 3–8. https://doi.org/10.1016/0304-3959(96)02994-6</p>
<p>Salganik, M. J., Dodds, P. S., &amp; Watts, D. J. (2006). <em>Experimental study of inequality and unpredictability in an artificial cultural market.</em> Science, 311(5762), 854–856. https://doi.org/10.1126/science.1121066</p>
<p>Schuck, N. W., Cai, M. B., Wilson, R. C., &amp; Niv, Y. (2016). <em>Human orbitofrontal cortex represents a cognitive map of state space.</em> Neuron, 91(6), 1402–1412. https://doi.org/10.1016/j.neuron.2016.08.019</p>
<p>Shenhav, A., Botvinick, M. M., &amp; Cohen, J. D. (2013). <em>The expected value of control: An integrative theory of anterior cingulate cortex function.</em> Neuron, 79(2), 217–240. https://doi.org/10.1016/j.neuron.2013.07.007</p>
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		<title>Time is Memory</title>
		<link>https://michaelhalassa.net/time-is-memory/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Mon, 01 Sep 2025 12:08:25 +0000</pubDate>
				<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neural circuits]]></category>
		<category><![CDATA[Neuroscience]]></category>
		<category><![CDATA[Science]]></category>
		<category><![CDATA[Working memory]]></category>
		<category><![CDATA[Cognitive Research]]></category>
		<category><![CDATA[Cognitive Science]]></category>
		<category><![CDATA[Computational Neuroscience]]></category>
		<category><![CDATA[Memory]]></category>
		<category><![CDATA[Temporal Memory]]></category>
		<category><![CDATA[Time]]></category>
		<guid isPermaLink="false">https://michaelhalassa.net/?p=789</guid>

					<description><![CDATA[Michael Halassa discusses how the brain may create the sense of memory and why time distortions happen in experience]]></description>
										<content:encoded><![CDATA[<p>Over the past year, I’ve found a new favorite running trail. It winds through woods, follows riverbanks, and slips through an old industrial complex. The scenery shifts constantly, broken into short, distinct segments.</p>
<p>I was surprised to discover that the run takes about an hour, almost exactly the same as my old trail from the year before. The distances are nearly identical too, which makes sense given that my pace hasn’t changed. And yet, the new trail <em>feels</em> much longer. How come?</p>
<p>The old route was simpler. It had three long, straight stretches where I could see the end from the beginning. Easy to remember, easy to chunk. The new one is nothing like that: shorter segments, sharper turns, and ever-changing backdrops. Every few minutes you’re in a completely new setting, never quite sure what’s around the bend.</p>
<p>That difference got me thinking about how we perceive time. We’ve all had those strange distortions: a memory from years ago that feels recent, or something from last week that feels impossibly distant. Time in the brain is slippery.</p>
<p>So how do we actually track it? Is there an internal clock ticking away? Probably not: decades of searching haven’t turned one up. A more likely explanation is that time is tied to how memories are organized and indexed. Let’s dig into what we actually know.</p>
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<h2 class="header-anchor-post"><strong>How Memory Creates Time</strong></h2>
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<p>The first clue comes from studying what happens when we remember. In a clever set of experiments, Olivier Jeunehomme and Arnaud D’Argembeau asked people to wear small automatic cameras while walking around a university campus. The cameras snapped photos every few seconds, creating an objective record of the experience. Later, participants were asked to verbally recall their walks while being audio-recorded.</p>
<p>The campus walks lasted around 40 minutes, but when participants replayed them aloud in memory, the descriptions only took about 5 minutes on average. That is roughly an eightfold compression of time.</p>
<p>The compression, however, was uneven. The researchers compared the recall transcripts to the time-stamped camera sequences and divided the narratives into what they called “experience units.” These were discrete remembered moments, such as buying a coffee, turning into a courtyard, or chatting with a classmate. Each unit was mapped back to the original footage so they could calculate how much real-world time it spanned.</p>
<p>The pattern was striking. Short, bounded activities with a clear goal, like making a purchase or opening a door, tended to be preserved in relatively high detail, replayed at about four to five times compression. In contrast, transitional stretches of locomotion, like walking from one building to the next, were compressed far more, sometimes by a factor of twenty or more. Long, uneventful stretches collapsed into a single unit, while activity-rich episodes retained much finer granularity.</p>
<p>These experience units appear to be the basic building blocks of episodic memory. The density of such units determines how long an episode feels in retrospect. More units per minute of clock time make for a richer memory trace and an expanded sense of duration. Fewer units create a thinner trace and a contracted sense of time.</p>
<p>Follow-up studies have highlighted the special role of event boundaries. Jeunehomme and D’Argembeau found that moments marking a change in context, such as entering a building, turning a corner, or meeting a person, were about five times more likely to be recalled than stretches in between. Boundaries act like bookmarks, segmenting the stream of experience and anchoring the flow of time in memory. These anchors not only determine what is remembered, but also shape how long the remembered experience feels.</p>
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<h2 class="header-anchor-post"><strong>The Paradox of Event Boundaries</strong></h2>
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<p>Experience units and event boundaries create a fundamental paradox in how we perceive time. Bangert and colleagues (2019, 2020) ran a series of experiments in which participants watched short films of everyday activities while making timing judgments. The films were paused at different points, and participants were asked to estimate whether a brief interval, usually around five seconds, had just passed. The twist was that sometimes the interval contained an event boundary, such as finishing washing dishes and beginning to dry them, and sometimes it did not. Intervals that contained a boundary were consistently judged as shorter than otherwise identical spans without one.</p>
<p>The mechanism behind this compression may become clearer when considering what&#8217;s happening in working memory. Swallow and colleagues (2009) tracked this directly by having participants watch movie clips while objects appeared on screen, a knife during sandwich-making, a towel during dishwashing. Five seconds later, the movie would pause for a recognition test. Objects present at event boundaries were recognized significantly better than those at non-boundaries. But this enhancement came with a cost: memory for objects from just before a boundary dropped dramatically. The boundary created a barrier, making it harder to retrieve information from the previous event even though it had occurred mere seconds earlier.</p>
<p>Event Segmentation Theory, developed by Jeffrey Zacks and colleagues in 2007, provides the framework. According to their theory, event boundaries are when the brain discards its current &#8220;event model&#8221; from working memory and uploads a new one. This updating process requires attention, which leaves fewer resources available for keeping track of time. As Bangert and colleagues (2020) demonstrated using dual-task paradigms, devoting attention to updating perceptual and conceptual features of the activity left fewer attentional resources for accumulating temporal information. It&#8217;s like trying to count seconds while also solving a puzzle &#8211; each boundary forces you to solve a new puzzle, and your counting falters.</p>
<p>The paradox is that the very same boundaries that compress time during experience expand it in memory. They serve as landmarks that structure recall, making events feel more spacious in retrospect. This dual effect helps explain a familiar puzzle: why the drive home from a new place usually feels longer than the drive there. On the outbound trip, the brain is constantly updating its models: pass the gas station (boundary), turn at the intersection (boundary), merge onto the highway (boundary). Each update reduces attention for tracking duration, so the drive feels shorter while you are in it. Yet those boundaries also create anchors that expand the memory of the trip. On the return drive the route is familiar, there are fewer surprises, and the brain needs fewer updates. With less attention diverted, duration is tracked more faithfully, so the drive feels longer in the moment but compresses more in memory.</p>
<p>Bangert and colleagues (2019) also tested temporal proximity, asking participants to judge how far apart two moments in the film felt. Boundaries made items seem further apart in time, even when the objective duration was identical. In this sense, boundaries insert psychological distance between moments. They stretch the remembered timeline even while compressing the lived experience of duration.</p>
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<h2 class="header-anchor-post"><strong>The Implications</strong></h2>
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<p>This framework explains a wide range of everyday paradoxes. Vacations, filled with novelty, fly by while they happen but expand richly in memory. Daily routines, stripped of boundaries, drag while we live them but collapse into nothing when recalled. Clewett and Davachi (2017) argued that the ebb and flow of experience itself determines the temporal structure of memory. Lositsky and colleagues (2016) showed that the greater the number and diversity of boundaries, the more time expands in recall.</p>
<p>It explains my running puzzle. My old trail was made up of long, predictable stretches, so it generated relatively few event boundaries. My new trail, by contrast, forced segmentation at every turn: woods to riverbank, riverbank to industrial ruins, sharp corner, sudden hill, unexpected vista. Each transition became a boundary, a new chunk in memory. The clock says both trails take about an hour, but memory disagrees. The old one collapses into a few coarse segments, while the new one expands into a much longer-feeling journey.</p>
<p>The principle is simple: if you want something to feel substantial in memory, add boundaries. Change contexts, vary activities, create moments that require updates. If you want time to flow by quickly, keep it continuous and predictable.</p>
<p>But the implications go deeper than personal experience design. This mechanism may explain why time seems to accelerate as we age. Childhood is packed with firsts, each creating boundaries: first day of school, first sleepover, first kiss. Adult life, especially in stable careers and relationships, can become a series of similar days bleeding into each other. The years feel shorter not because our metabolism changes or because of some cosmic injustice, but because we&#8217;re creating fewer distinct memory segments.</p>
<p>The brain doesn&#8217;t keep time like a clock. It builds time from its internal dynamics. The elasticity of time isn&#8217;t an illusion; it&#8217;s how the mind constructs a temporal dimension from the boundaries of experience.</p>
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<p><em>If you enjoyed this piece, let me know. I’d love to hear how you’ve experienced time stretching or compressing in your own life. I’ll also be following up with another post that digs into the neural substrates of time perception, exploring how brain circuits generate these distortions.</em></p>
<p><em>If you’d like to read that when it comes out, consider subscribing or sharing this piece with someone who might find it interesting.</em></p>
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<h2 class="header-anchor-post"><strong>Bibliography</strong></h2>
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<p>Bangert, A. S., Kurby, C. A., Hughes, A. S., &amp; Carrasco, O. (2019). Crossing event boundaries changes prospective perceptions of temporal length and proximity. <em>Attention, Perception, &amp; Psychophysics</em>, 81(8), 2459-2472.</p>
<p>Block, R. A., &amp; Zakay, D. (1997). Prospective and retrospective duration judgments: A meta-analytic review. <em>Psychonomic Bulletin &amp; Review</em>, 4(2), 184-197.</p>
<p>Clewett, D., &amp; Davachi, L. (2017). The ebb and flow of experience determines the temporal structure of memory. <em>Current Opinion in Behavioral Sciences</em>, 17, 186-193.</p>
<p>Jeunehomme, O., &amp; D&#8217;Argembeau, A. (2020). Event segmentation and the temporal compression of experience in episodic memory. <em>Psychological Research</em>, 84(2), 481-490.</p>
<p>Lositsky, O., Chen, J., Toker, D., Honey, C. J., Shvartsman, M., Poppenk, J. L., &#8230; &amp; Norman, K. A. (2016). Neural pattern change during encoding of a narrative predicts retrospective duration estimates. <em>eLife</em>, 5, e16070.</p>
<p>Swallow, K. M., Zacks, J. M., &amp; Abrams, R. A. (2009). Event boundaries in perception affect memory encoding and updating. <em>Journal of Experimental Psychology: General</em>, 138(2), 236-257.</p>
<p>Zacks, J. M., Speer, N. K., Swallow, K. M., Braver, T. S., &amp; Reynolds, J. R. (2007). Event perception: A mind-brain perspective. <em>Psychological Bulletin</em>, 133(2), 273-293.</p>
<p>Zacks, J. M., Kurby, C. A., Eisenberg, M. L., &amp; Haroutunian, N. (2011). Prediction error associated with the perceptual segmentation of naturalistic events. <em>Journal of Cognitive Neuroscience</em>, 23(12), 4057-4066.</p>
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		<title>The Brain&#8217;s &#8220;What If&#8221; Engine: Why Counterfactuals Are Key to Human Intelligence</title>
		<link>https://michaelhalassa.net/counterfactuals-human-intelligence/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Sun, 03 Aug 2025 23:04:06 +0000</pubDate>
				<category><![CDATA[Cognitive flexibility]]></category>
		<category><![CDATA[Computational neuroscience]]></category>
		<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neural circuits]]></category>
		<category><![CDATA[NeuroAI]]></category>
		<category><![CDATA[Neuroscience]]></category>
		<category><![CDATA[Prefrontal cortex]]></category>
		<category><![CDATA[Working memory]]></category>
		<category><![CDATA[Computational Neuroscience]]></category>
		<category><![CDATA[neuroscience]]></category>
		<category><![CDATA[Recurrent Neural Networks]]></category>
		<category><![CDATA[research paper]]></category>
		<category><![CDATA[Science]]></category>
		<guid isPermaLink="false">https://michaelhalassa.net/?p=785</guid>

					<description><![CDATA[Michael Halassa discusses recent work on counterfactual reasoning and its contribution to human cognition]]></description>
										<content:encoded><![CDATA[<p>I&#8217;ve always been fascinated by the kinds of thoughts we <em>don&#8217;t</em> act on. In psychiatry, they shape regret, resilience, and rumination. In neuroscience, they reveal a deep truth about how the brain handles uncertainty. Every morning when I&#8217;m running late, I catch myself thinking: &#8220;If only I&#8217;d left five minutes earlier.&#8221; It&#8217;s a fleeting thought, but it represents one of the most computationally sophisticated processes our brains perform: imagining alternative realities that never happened.</p>
<p>Every day, your brain performs millions of &#8220;what if&#8221; calculations without you even noticing. What if I had taken the other route to work? What if I hadn&#8217;t said that in the meeting? What if the ball bounces differently than expected? This capacity for <strong>counterfactual reasoning</strong>, imagining alternative realities that never actually occurred, represents one of the most sophisticated computational achievements of biological intelligence.</p>
<p>A groundbreaking new study published in <em>Nature Human Behaviour</em> by Ramadan, Tang, Watters, and Jazayeri has shed new light on why humans rely on these mentally expensive &#8220;what if&#8221; simulations, revealing computational constraints that force our brains into remarkably clever problem-solving strategies. Their findings illuminate human cognition and change how we understand intelligence itself.</p>
<h2>The Computational Mystery: Why Do We Think in &#8220;What Ifs&#8221;?</h2>
<p>From a purely computational standpoint, counterfactual reasoning seems inefficient. When facing complex decisions, optimal algorithms should simply compute the joint probability of all possible outcomes and pick the best option. So why do humans constantly engage in the seemingly wasteful exercise of imagining alternatives?</p>
<p>The answer, as Ramadan and colleagues discovered, lies in the fundamental constraints that shape how our brains process information. Using an ingenious H-maze task where participants had to track an invisible ball through branching pathways, they uncovered three critical computational bottlenecks that force human cognition into hierarchical and counterfactual strategies:</p>
<p><strong>1. Parallel Processing Bottleneck</strong>: Our brains cannot track all possible trajectories simultaneously. We must break complex problems into sequential, hierarchical steps.</p>
<p><strong>2. Counterfactual Processing Noise</strong>: When we engage in &#8220;what if&#8221; thinking, our working memory introduces noise that degrades the fidelity of these mental simulations.</p>
<p><strong>3. Rational Resource Allocation</strong>: Humans adaptively adjust their reliance on counterfactuals based on how much these mental simulations cost them.</p>
<h2>Very Clever Use of Recurrent Neural Networks in Modeling Features of the Human Mind</h2>
<p>The research reveals profound insights about intelligence itself. When Ramadan et al. created artificial neural networks and subjected them to the same computational constraints humans face, something remarkable happened: only the networks constrained by all three bottlenecks reproduced human-like behavior.</p>
<p>This finding demonstrates the power of using recurrent neural networks to model human cognition. By constraining artificial networks with the same limitations that shape human thinking, Ramadan et al. created systems that behave remarkably like people. The key insight is that RNNs can capture mental processes like hierarchical and counterfactual reasoning when they face the same computational bottlenecks humans do.</p>
<h3>Neural Architecture of Counterfactual Reasoning</h3>
<p>The neural implementation of counterfactual reasoning tells a more complex story beyond frontal control. Van Hoeck and colleagues&#8217; landmark fMRI study revealed that counterfactual thinking engages a distributed network that hijacks the brain&#8217;s episodic memory system.</p>
<p>When participants imagined &#8220;upward counterfactuals&#8221; (better outcomes for negative past events), their brains activated the same core memory network used for remembering the past and imagining the future: hippocampus, posterior cingulate, inferior parietal lobule, lateral temporal cortices, and medial prefrontal cortex.</p>
<p>What makes counterfactual reasoning computationally expensive becomes clear in this neural architecture. Counterfactual thinking recruited these memory regions more extensively than episodic past or future thinking, and additionally engaged bilateral inferior parietal lobe and posterior medial frontal cortex.</p>
<p>The extra brain activity reflects just how demanding this kind of mental juggling really is: counterfactual reasoning requires simultaneously maintaining factual and contrafactual representations while actively inhibiting the dominant factual reality.</p>
<p>The brain has evolved specialized circuitry for tracking &#8220;what might have been.&#8221; Boorman and colleagues discovered that lateral frontopolar cortex, dorsomedial frontal cortex, and posteromedial cortex form a dedicated network for encoding counterfactual choice values: tracking not just what happened, but whether alternative options might be worth choosing in the future.</p>
<p>This network operates in parallel to the ventromedial prefrontal system that tracks the value of chosen options, suggesting that the brain maintains separate computational channels for factual and counterfactual value processing.</p>
<p>Perhaps most remarkably, recent work has shown that counterfactual information fundamentally transforms how the brain codes value itself. When counterfactual outcomes are available, medial prefrontal and cingulate cortex shift from absolute to relative value coding.</p>
<p>Think of it this way: losing $10 feels terrible if you could have won $50, but feels great if you could have lost $100. The same neural outcome is processed as positive in a loss context (absence of punishment) but negative in a gain context (absence of reward).</p>
<p>This neural flexibility mirrors the adaptive computational strategies revealed in behavioral studies: the brain dynamically reconfigures its representational schemes based on available information and processing constraints.</p>
<p>These findings illuminate why counterfactual reasoning is both computationally expensive and evolutionarily preserved. The enhanced neural demands reflect genuine computational costs: maintaining multiple alternative representations, binding novel scenario elements, and managing conflict between factual and counterfactual worlds. Yet this system enables the kind of flexible, context-sensitive reasoning that allows humans to learn from paths not taken and adapt behavior based on imagined alternatives.</p>
<h2>The Bounded Rationality Renaissance</h2>
<p>These discoveries are part of a broader renaissance in understanding <strong>bounded rationality</strong>, the idea that intelligent behavior emerges not from perfect optimization, but from smart adaptations to computational limitations.</p>
<p>Herbert Simon&#8217;s revolutionary concept of bounded rationality challenged the assumptions of perfect rationality in classical economic theory, proposing instead that individuals &#8220;satisfice&#8221; (seeking good enough solutions rather than optimal ones) due to limitations in computation, time, information, and cognitive resources.</p>
<p>Simon&#8217;s work recognized that &#8220;perfectly rational decisions are often not feasible in practice because of the intractability of natural decision problems and the finite computational resources available for making them.&#8221; This insight has profound implications for both understanding human cognition and designing artificial intelligence systems.</p>
<h3>The Bigger Picture</h3>
<p>The Ramadan study reveals something profound: the cognitive strategies we think of as distinct (hierarchical reasoning, counterfactual thinking, simple optimization) actually lie along a continuum. Human intelligence dynamically shifts between these approaches based on available mental resources and task demands.</p>
<p>This has implications beyond neuroscience. If counterfactual reasoning emerges from computational constraints rather than being hardwired, it suggests these &#8220;what if&#8221; processes might be fundamental to any sufficiently complex intelligence, biological or artificial.</p>
<h2>Clinical Frontiers: When Counterfactuals Break Down</h2>
<p>From a clinical perspective, this research offers new windows into psychiatric and neurological conditions. Counterfactual reasoning depends on integrative networks for affective processing, mental simulation, and cognitive control. These are systems that are systematically altered in psychiatric illness and neurological disease.</p>
<p>Consider a patient with OCD who gets trapped in endless loops of &#8220;what if I didn&#8217;t check the door?&#8221; or someone with depression whose counterfactual thinking spirals into &#8220;if only I were different, everything would be better.&#8221; Understanding the computational basis of these patterns could lead to more targeted therapeutic approaches.</p>
<p>Patients with schizophrenia show specific deficits in counterfactual reasoning when complex non-factual elements are needed to understand social environments. By mapping how these computational processes break down, we&#8217;re gaining new tools for both diagnosis and treatment.</p>
<h2>The Bottom Line: Constraints as Features</h2>
<p>The story of counterfactual reasoning is a story about the power of constraints. What initially appears to be a computational limitation (our inability to process all information in parallel) turns out to be the very foundation of human cognitive flexibility.</p>
<p>The human brain&#8217;s &#8220;what if&#8221; engine represents an elegant solution that emerges from the interplay between computational constraints and adaptive intelligence. As we stand on the brink of artificial general intelligence, perhaps the secret lies not in building systems that can process everything at once, but systems that can gracefully adapt to the fundamental constraints that shape all intelligence.</p>
<p>The future of AI may not lie in eliminating human limitations, but in understanding why those limitations exist and what remarkable capabilities they make possible.</p>
<hr />
<p><em>This convergence of neuroscience, cognitive science, and AI represents a fundamental shift in how we understand intelligence. Rather than seeing computational constraints as problems to solve, we&#8217;re beginning to recognize them as the very features that make flexible, adaptive intelligence possible. The brain&#8217;s &#8220;what if&#8221; engine may be a blueprint for the next generation of truly intelligent machines.</em></p>
<p>The next time you wonder what might have been, remember: that question may be the very core of what makes you human.</p>
<hr />
<h2>Bibliography</h2>
<p>Boorman, E. D., Behrens, T. E., &amp; Rushworth, M. F. (2011). Counterfactual choice and learning in a neural network centered on human lateral frontopolar cortex. <em>PLoS Biology</em>, 9(6), e1001093.</p>
<p>Pischedda, D., Palminteri, S., &amp; Coricelli, G. (2020). The effect of counterfactual information on outcome value coding in medial prefrontal and cingulate cortex: From an absolute to a relative neural code. <em>Journal of Neuroscience</em>, 40(16), 3268-3277.</p>
<p>Ramadan, M., Tang, C., Watters, N., &amp; Jazayeri, M. (2025). Computational basis of hierarchical and counterfactual information processing. <em>Nature Human Behaviour</em>. doi:10.1038/s41562-025-02232-3.</p>
<p>Simon, H. A. (1955). A behavioral model of rational choice. <em>Quarterly Journal of Economics</em>, 69(1), 99-118.</p>
<p>Van Hoeck, N., Ma, N., Ampe, L., Baetens, K., Vandekerckhove, M., &amp; Van Overwalle, F. (2013). Counterfactual thinking: An fMRI study on changing the past for a better future. <em>Social Cognitive and Affective Neuroscience</em>, 8(5), 556-564.</p>
<p>Van Hoeck, N., Watson, P. D., &amp; Barbey, A. K. (2015). Cognitive neuroscience of human counterfactual reasoning. <em>Frontiers in Human Neuroscience</em>, 9, 420.</p>
<p>Zador, A., Escola, S., Richards, B., et al. (2023). Catalyzing next-generation Artificial Intelligence through NeuroAI. <em>Nature Communications</em>, 14, 1597.</p>
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		<title>The Next Chapter of AI: Leveraging the Evolutionary Principles Powering Human Intelligence</title>
		<link>https://michaelhalassa.net/neuroai2025/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Thu, 17 Jul 2025 09:04:13 +0000</pubDate>
				<category><![CDATA[Cognitive flexibility]]></category>
		<category><![CDATA[Cognitive Processing]]></category>
		<category><![CDATA[Computational neuroscience]]></category>
		<category><![CDATA[Halassa Lab]]></category>
		<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neural circuits]]></category>
		<category><![CDATA[NeuroAI]]></category>
		<category><![CDATA[Neuroscience]]></category>
		<category><![CDATA[Science]]></category>
		<category><![CDATA[Brain scientist]]></category>
		<category><![CDATA[Computational Neuroscience]]></category>
		<category><![CDATA[Recurrent Neural Networks]]></category>
		<guid isPermaLink="false">https://michaelhalassa.net/?p=772</guid>

					<description><![CDATA[Michael Halassa explores the intersection between Neuroscience and AI (NeuroAI) highlighting research on flexible cognition.]]></description>
										<content:encoded><![CDATA[<p>A mouse can explore a new environment, find food and adapt when the rules change, all using less energy than a lightbulb. Meanwhile, our most powerful computers can solve chess and master protein folding, but still can’t walk across a messy room without crashing into a chair.</p>
<p>This contrast reveals something profound about intelligence itself and where we need to go next. As we celebrate Geoffrey Hinton and John Hopfield&#8217;s recent Nobel Prize in Physics for their foundational work on neural networks, it&#8217;s the perfect time to ask: what&#8217;s the next chapter in understanding intelligence?</p>
<p><strong>The Great Intelligence Paradox</strong></p>
<p>We&#8217;re living through what some call the &#8220;Great Intelligence Paradox.&#8221; Our most advanced computational systems can master protein folding and beat world champions at Go, tasks that require incredible sophistication. But they&#8217;re surprisingly brittle when faced with the kind of flexible, real-world intelligence that any animal takes for granted.</p>
<p>Consider this: no machine can build a nest, forage for berries, or care for young. Today&#8217;s computational systems cannot compete with the sensorimotor capabilities of a four-year old child or even simple animals. The reason isn&#8217;t that we lack computational power. It&#8217;s that we&#8217;ve been approaching intelligence from a different angle.</p>
<p>As researcher Hans Moravec put it, abstract thought &#8220;is a new trick, perhaps less than 100 thousand years old, effective only because it is supported by this much older and much more powerful, though usually unconscious, sensorimotor knowledge.&#8221; In other words, when trying to capture natural intelligence, we&#8217;ve been focusing on the penthouse without first understanding the foundation.</p>
<p><strong>The Deep History of NeuroAI: A 70-Year Symbiosis</strong></p>
<p>This realization has sparked the emergence of NeuroAI, a field that recognizes something remarkable: evolution has already solved many of the problems we&#8217;re struggling with in artificial intelligence. But the connection between neuroscience and computing isn&#8217;t new. It can be traced to the very foundations of modern computer science itself.</p>
<p>John von Neumann&#8217;s seminal 1945 report outlining the first computer architecture (EDVAC) dedicated an entire chapter to discussing whether the proposed system was sufficiently brain-like. Remarkably, the only citation in this foundational document was to Warren McCulloch and Walter Pitts&#8217; 1943 paper, widely considered the first work on neural networks. This early cross-pollination between neuroscience and computer science set the stage for decades of mutual inspiration.</p>
<p>The relationship deepened with Frank Rosenblatt&#8217;s introduction of the perceptron in 1958. The revolutionary idea here wasn&#8217;t just that machines could learn, but that they should learn from data rather than being explicitly programmed. Rosenblatt established synaptic connections as the primary locus of learning in artificial neural networks, a concept heavily influenced by Donald Hebb&#8217;s 1949 work highlighting the importance of the synapse as the physical basis of learning and memory.</p>
<p>This neuroscience-inspired principle that synapses are the plastic elements of neural networks has remained absolutely central to modern computation. Even when Marvin Minsky and Seymour Papert&#8217;s 1969 critique of perceptrons triggered the first &#8220;neural network winter,&#8221; the core insight persisted.</p>
<p>The symbiosis between artificial and biological neural network research has produced numerous breakthrough success stories. Perhaps the most celebrated is the convolutional neural network (CNN), which powers many of today&#8217;s most successful artificial vision systems. CNNs were directly inspired by David Hubel and Torsten Wiesel&#8217;s model of the visual cortex, work that earned them a Nobel Prize more than four decades ago.</p>
<p>Another home run is reinforcement learning, which has driven groundbreaking achievements including Google DeepMind&#8217;s AlphaZero and AlphaGo. The computational principles underlying these systems mirror the dopamine-mediated learning circuits in biological brains. When a monkey reaches for a reward and receives more than expected, dopamine neurons fire in patterns that precisely match the temporal difference learning algorithms used in these game-playing systems.</p>
<p>More recently, the concept of &#8220;dropout&#8221; has gained prominence in artificial neural networks. This technique, in which individual neurons are randomly deactivated during training to prevent overfitting, draws inspiration from the brain&#8217;s use of stochastic processes. By mimicking the occasional misfiring of neurons, dropout encourages networks to develop more robust and resilient representations.</p>
<p>Critically, this relationship is truly mutualistic, not parasitic. Computational advances have revolutionized neuroscience as much as neuroscience has inspired computation. Artificial neural networks now form the backbone of state-of-the-art models of the visual cortex. The success of these models in solving complex perceptual tasks has generated new hypotheses about how biological brains might perform similar computations.</p>
<p><strong>Why Animals Are the Ultimate Intelligence Teachers</strong></p>
<p>Instead of trying to replicate what makes humans special, we should look at what makes all animals successful. These are the capabilities that have been tested and refined over 500 million years of evolution.</p>
<p>This is where Tony Zador and his colleagues propose the &#8220;embodied Turing test.&#8221; The idea is straightforward but profound: instead of asking whether computation can fool us in conversation, we should ask whether an artificial beaver can build a dam as skillfully as a real one, or whether an artificial squirrel can navigate through trees with the same agility.</p>
<p>This shift in perspective reveals three crucial capabilities that current computational systems lack:</p>
<p><strong>They Engage Their Environment</strong></p>
<p>The defining feature of animals is their ability to move around and interact with their environment in purposeful ways. It&#8217;s about understanding how actions affect the world and using that understanding to achieve goals.</p>
<p>Consider the computational challenge this represents. When you watch a cat stalking prey, you&#8217;re witnessing real-time integration of visual tracking, motor prediction, uncertainty estimation, and action selection. The cat must predict the prey&#8217;s trajectory, estimate the optimal interception point, account for its own motor delays, and continuously update its strategy as the situation evolves. This requires what computational scientists call forward models, inverse models, and optimal control, all running simultaneously in a brain that weighs 30 grams.</p>
<p>Or take nest building in birds. A Baltimore oriole weaves together hundreds of individual grass fibers, each requiring precise motor control and spatial reasoning. The bird must estimate structural integrity in real-time, adapt to varying material properties, and maintain a global architectural plan while executing thousands of local actions. No current robotic system can approach this level of sensorimotor sophistication.</p>
<p><strong>They Behave Flexibly</strong></p>
<p>Animals are born with most of the skills needed to thrive or can rapidly acquire them from limited experience, thanks to their strong foundation in real-world interaction, courtesy of evolution and development. Unlike computational systems that catastrophically fail when encountering scenarios outside their training data, animals excel at handling novel situations by drawing on their general understanding of how the world works.</p>
<p>This flexibility emerges from what neuroscientists call compositional representation. Rather than memorizing specific stimulus-response patterns, animals build internal models of causal structure that can be recombined in novel ways. A squirrel encountering an unfamiliar tree can still navigate it by applying general principles of branch mechanics, gravity, and momentum.</p>
<p>Recent work by Rajalingham and colleagues has provided a striking demonstration of this principle. They trained monkeys to play &#8220;mental Pong,&#8221; where a ball disappeared behind a barrier and the animal had to predict where it would emerge. Neural recordings from the monkeys&#8217; frontal cortex revealed that the brain was running a mental physics engine, maintaining an internal trajectory that matched physical reality even when the ball was invisible.</p>
<p>Even more remarkably, when computational systems were trained on the same task but required to infer the ball&#8217;s hidden path, they produced patterns of activity that mirrored the monkey frontal cortex. This suggests that both biological and artificial systems converge on similar computational solutions when solving similar problems, but biological systems achieve this with far greater efficiency and flexibility.</p>
<p><strong>They Compute Efficiently</strong></p>
<p>Here&#8217;s a staggering comparison that reveals the depth of the efficiency gap: training a large language model such as GPT-3 requires over 1000 megawatt-hours, enough electricity to power a small town for a day. The human brain uses about 20 watts, roughly the same as a bright light bulb.</p>
<p>This efficiency gap points to fundamentally different computational principles. Biological circuits operate in a regime where spikes are sparse and energy-efficient, using asynchronous communication protocols that bear little resemblance to the synchronous, dense matrix operations that characterize current computational systems.</p>
<p>The brain achieves this efficiency through several key innovations. First, it uses event-driven computation, where neurons only consume energy when they have something important to communicate. Second, it employs local learning rules that don&#8217;t require global coordination or backpropagation of error signals. Third, it multiplexes different types of information in the same circuits, allowing the same neural hardware to support multiple functions depending on context.</p>
<p>Recent advances in neuromorphic engineering are beginning to capture some of these principles. Intel&#8217;s Loihi chip and IBM&#8217;s TrueNorth processor implement spiking neural networks that dramatically reduce power consumption for certain tasks. But we&#8217;re still far from achieving the full computational elegance of biological systems.</p>
<p><strong>Our Research: Natural Architectures for Cognitive Flexibility</strong></p>
<p>This broader NeuroAI vision connects directly to collaborative research efforts my colleagues and I have been pursuing through the Thalamus Conte Center at Princeton. Working alongside talented investigators, we&#8217;ve been studying how thalamic circuits, particularly the mediodorsal thalamus, regulate uncertainty and cognitive flexibility.</p>
<p>The thalamus has long been thought of as a simple relay station, passively transferring information between brain regions. Our work reveals a far more sophisticated picture: the thalamus acts as a regulator of cortical representations, actively regulating the flow of information based on context, confidence, and computational demands.</p>
<p>Recent findings show that the mediodorsal thalamus exhibits distinct coding properties from prefrontal cortex. While prefrontal areas represent information in high-dimensional, mixed formats that can support many different behaviors, the thalamus compresses this information into lower-dimensional representations focused on key contextual variables like task rules and uncertainty estimates.</p>
<p>This architectural arrangement resembles what computational scientists call &#8220;regularization,&#8221; where a system constrains its processing to focus on the most relevant dimensions of a problem. The thalamus appears to provide this kind of regularization to prefrontal networks, helping them avoid getting lost in irrelevant details while maintaining the flexibility to handle novel situations.</p>
<p>This has direct implications for understanding psychiatric disorders. Schizophrenia, for instance, involves difficulties with cognitive flexibility and context processing. Our work suggests that these may reflect specific disruptions in thalamic computation rather than global deficits in learning or reasoning.</p>
<p>Understanding how evolution solved the uncertainty problem in biological brains could be the key to creating computational systems that are truly adaptive and robust in the face of novel situations. Current systems struggle precisely because they lack principled ways to handle uncertainty and adjust their confidence based on context.</p>
<p><strong>The Road Ahead: From Lab to Life</strong></p>
<p>The implications of this NeuroAI approach extend far beyond academic laboratories. The convergence of insights from biological intelligence and computational innovation points toward systems that could:</p>
<p><strong>Adapt like animals</strong>: Robots that learn to navigate new environments with the flexibility of a mouse exploring new territory. Imagine search and rescue robots that can adapt to novel disaster scenarios, or autonomous vehicles that can handle completely unprecedented road conditions by drawing on fundamental principles of navigation and obstacle avoidance rather than memorized patterns.</p>
<p><strong>Learn efficiently</strong>: Systems that acquire new skills from limited examples, like how animals quickly adapt to new food sources or threats. A key insight from biological learning is the importance of strong inductive biases, the built-in assumptions that help guide learning in the right direction. Animals don&#8217;t start from scratch; they leverage millions of years of evolutionary optimization.</p>
<p><strong>Handle uncertainty gracefully</strong>: Systems that know when they don&#8217;t know, actively seeking information to improve their decisions rather than confidently making wrong choices. This requires implementing something like the thalamic uncertainty computation we&#8217;ve been studying, a principled way to calibrate confidence and adjust exploration strategies based on current knowledge state.</p>
<p><strong>Integrate seamlessly</strong>: Computation that works alongside humans as naturally as animals coordinate in flocks or herds. This requires understanding not just individual intelligence but collective intelligence, how multiple agents can share information and coordinate actions without centralized control.</p>
<p>Recent experimental work provides concrete examples of how these principles might be implemented. Researchers at DeepMind have developed systems that can learn to play multiple Atari games using the same general-purpose algorithm, rather than requiring game-specific training. Their success comes from incorporating biological principles like replay (reactivating and reorganizing memories during rest) and curiosity-driven exploration.</p>
<p>Similarly, researchers at OpenAI have shown that large language models can exhibit emergent reasoning capabilities when scaled up, suggesting that some aspects of flexible intelligence might emerge from sufficient computational scale combined with appropriate architectural principles.</p>
<p>But perhaps the most promising developments come from robotics, where researchers are beginning to implement embodied learning principles. Boston Dynamics&#8217; robots can navigate complex terrain and recover from perturbations in ways that would have been impossible just a few years ago. Their success comes from combining traditional control theory with machine learning approaches that can adapt to novel situations.</p>
<p><strong>A New Kind of Intelligence</strong></p>
<p>Building models that can pass the embodied Turing test requires more than tweaking existing algorithms. As Zador and colleagues argue, we need a &#8220;large-scale effort to identify and understand the principles of biological intelligence and abstract those for application in computer and robotic systems.&#8221;</p>
<p>Two key insights emerge from this challenge. First, intelligence isn&#8217;t about building internal representations—it&#8217;s about discovering affordances, the opportunities for action that emerge from the interaction between an agent and its environment. Second, animals don&#8217;t just learn; they develop, with their learning capabilities changing over time. Understanding how biological systems bootstrap from simple reflexes to sophisticated reasoning could transform how we build adaptive computational systems.</p>
<p>The convergence of neuroscience and computation offers concrete opportunities for progress. Animals solve computational problems that current systems struggle with, using principles refined over hundreds of millions of years of evolution. The mouse exploring a maze demonstrates flexible navigation, efficient learning from limited experience, and robust generalization. These capabilities emerge from biological circuits that balance exploration with exploitation, build and update internal maps, and adapt to novel situations.</p>
<p>Progress will require sustained collaboration between neuroscientists, computer scientists, and engineers. The questions are concrete: How do biological systems achieve such efficiency? What computational principles underlie adaptive behavior? How can we implement these in artificial systems?</p>
<p>Want to dive deeper into these ideas? Join us at CNS2025 in Florence, Italy (July 5-9, 2025) for our NeuroAI workshop, where we&#8217;ll explore how the convergence of neuroscience and computation is shaping the future of both fields. More details at cnsorg.org/cns-2025.</p>
<p><strong>References</strong></p>
<p>Zador, A., Escola, S., Richards, B., Ölveczky, B., Bengio, Y., Boahen, K., Botvinick, M., Chklovskii, D., Collins, A., Doya, K., Hassabis, D., Kording, K., Konidaris, G., Marblestone, A., Olshausen, B., Pouget, A., Sejnowski, T., Simoncelli, E., Solla, S., Sussillo, D., Tsao, D., &amp; Tsodyks, M. (2023). Catalyzing next-generation Artificial Intelligence through NeuroAI. <em>Nature Communications</em>, 14, 1597. https://doi.org/10.1038/s41467-023-37180-x</p>
<p>Zador, A. (2024). NeuroAI: A field born from the symbiosis between neuroscience and computation. <em>The Transmitter</em>. https://www.thetransmitter.org/neuroai/neuroai-a-field-born-from-the-symbiosis-between-neuroscience-ai/</p>
<p>Rajalingham, R., Sohn, H. &amp; Jazayeri, M. (2025). Dynamic tracking of objects in the macaque dorsomedial frontal cortex. <em>Nature Communications</em>, 16, 346. https://doi.org/10.1038/s41467-024-54688-y</p>
<p>Thalamus Conte Center. (2024). Princeton University. https://conte.thalamus.princeton.edu/</p>
<p>Hubel, D. H., &amp; Wiesel, T. N. (1962). Receptive fields, binocular interaction and functional architecture in the cat&#8217;s visual cortex. <em>The Journal of Physiology</em>, 160(1), 106-154.</p>
<p>von Neumann, J. (1945). First Draft of a Report on the EDVAC. University of Pennsylvania.</p>
<p>McCulloch, W. S., &amp; Pitts, W. (1943). A logical calculus of the ideas immanent in nervous activity. <em>Bulletin of Mathematical Biophysics</em>, 5(4), 115-133.</p>
<p>Rosenblatt, F. (1958). The perceptron: A probabilistic model for information storage and organization in the brain. <em>Psychological Review</em>, 65(6), 386-408.</p>
<p>Hebb, D. O. (1949). <em>The Organization of Behavior: A Neuropsychological Theory</em>. Wiley.</p>
<p>Minsky, M., &amp; Papert, S. (1969). <em>Perceptrons: An Introduction to Computational Geometry</em>. MIT Press.</p>
<p>Moravec, H. (1988). <em>Mind Children: The Future of Robot and Human Intelligence</em>. Harvard University Press.</p>
<p>&nbsp;</p>
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		<title>When the Brain&#8217;s Uncertainty computer goes offline: New Human Evidence for Thalamic Regulation of Decision-Making</title>
		<link>https://michaelhalassa.net/thalamus-uncertainty-decisions/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Fri, 11 Jul 2025 04:32:22 +0000</pubDate>
				<category><![CDATA[Cognitive flexibility]]></category>
		<category><![CDATA[Cognitive Processing]]></category>
		<category><![CDATA[Computational neuroscience]]></category>
		<category><![CDATA[Halassa Lab]]></category>
		<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neural circuits]]></category>
		<category><![CDATA[Prefrontal cortex]]></category>
		<category><![CDATA[Thalamocortical circuits]]></category>
		<category><![CDATA[Halassa]]></category>
		<category><![CDATA[MD thalamus]]></category>
		<category><![CDATA[Thalamocortical]]></category>
		<guid isPermaLink="false">https://michaelhalassa.net/?p=754</guid>

					<description><![CDATA[New research reveals how the mediodorsal thalamus regulates decision-making confidence and exploration behavior in humans. Michael Halassa discusses breakthrough findings from focused ultrasound studies showing thalamic control of belief updating and cognitive flexibility.]]></description>
										<content:encoded><![CDATA[<h2>An Elegant Natural Experiment</h2>
<p>The study by Mackenzie et al. (2025, bioRxiv) represents a particularly clever approach to understanding human thalamic function. Rather than relying on correlational neuroimaging, the researchers capitalized on an unintended consequence of focused ultrasound thalamotomy for essential tremor. When post-surgical vasogenic edema extended beyond the intended motor target into cognitive thalamic regions, it created a rare opportunity to assess the causal contribution of different thalamic nuclei to decision-making behavior.</p>
<p>What makes this approach so powerful is the precision it affords. Patients served as their own controls, tested before and after surgery on a sophisticated decision-making paradigm that probes the exploration-exploitation trade-off under uncertainty.</p>
<h2>Computational Dissection of Behavioral Changes</h2>
<p>Using the restless four-armed bandit task, which requires continuous adaptation to changing reward contingencies, the researchers could probe multiple aspects of decision-making simultaneously. The task&#8217;s Gaussian random walk structure creates ongoing uncertainty, forcing participants to balance between exploiting currently favored options and exploring alternatives that might yield better outcomes.</p>
<p>The key innovation came from their computational modeling approach. Rather than simply observing that patients made more &#8220;stay&#8221; choices post-surgery, the authors fitted multiple reinforcement learning model variants to decompose the underlying decision processes. This revealed that the behavioral shift was best captured by a Bayesian learning model with increased reward sensitivity (β) but eliminated exploration bonus—suggesting that patients weren&#8217;t simply perseverating, but had fundamentally altered confidence in their value estimates.</p>
<p>Most strikingly, when using their winning model to classify choice types, the researchers found a dramatic reduction in <strong>directed exploration</strong>—the strategic sampling of uncertain options to gain information (Mackenzie et al., 2025). This wasn&#8217;t random exploration or simple indecision, but the specific loss of information-seeking behavior that would normally help resolve uncertainty about option values.</p>
<h2>Anatomical Precision and Circuit Specificity</h2>
<p>The neuroimaging analysis provided crucial anatomical specificity. The degree of behavioral change correlated specifically with edema extension into the <strong>lateral (parvocellular) mediodorsal nucleus</strong>—not other thalamic regions including the intended surgical target (VIM). This specificity is important given the known functional subdivisions within MD:</p>
<ul>
<li><strong>Lateral MD → DLPFC/Frontal Pole</strong>: Dense reciprocal connectivity with areas involved in cognitive control and abstract rule formation</li>
<li><strong>Medial MD → OFC/vmPFC</strong>: Connections with valuation and reward-processing regions</li>
</ul>
<p>The functional connectivity analysis further supported this anatomical specificity. Using normative connectome data, individual patients&#8217; behavioral changes could be predicted from the connectivity profile between their lesioned MD voxels and prefrontal cortex—but only for MD, not other thalamic nuclei.</p>
<p>&nbsp;</p>
<h2>Mechanistic Insights: From Confidence Calibration to Circuit Function</h2>
<p>The computational framework reveals something more nuanced than simple &#8220;inflexibility.&#8221; The post-lesion behavioral profile suggests a specific breakdown in <strong>uncertainty representation</strong>—what we might call miscalibrated confidence. When MD-PFC communication is compromised, the system appears to default to high confidence in existing value representations, reducing sensitivity to contradictory information.</p>
<p>This aligns with emerging theoretical frameworks positioning the MD thalamus as a critical node in hierarchical inference, helping to coordinate distributed computations for flexible and efficient learning (Scott et al., 2024). The loss of directed exploration particularly supports this view, as this behavior specifically emerges when agents are uncertain about their value estimates and seek information to reduce that uncertainty.</p>
<h2>Bridging Animal Models and Human Neuroscience</h2>
<p>The convergence with our rodent findings is striking:</p>
<p><strong>Animal Studies (MD inactivation/optogenetics)</strong>:</p>
<ul>
<li>Reduced flexibility in volatile environments</li>
<li>Animals fail to revise beliefs when contingencies change</li>
<li>Inflated certainty in action values</li>
<li>Deficit specific to belief updating, not initial learning</li>
</ul>
<p><strong>Human Study (accidental MD lesions)</strong>:</p>
<ul>
<li>Reduced switching in uncertain environments</li>
<li>Patients fail to explore when exploration would be beneficial</li>
<li>Increased confidence in value estimates</li>
<li>Preserved basic learning ability</li>
</ul>
<p>This cross-species convergence suggests we&#8217;ve identified a fundamental computational principle rather than a species-specific curiosity.</p>
<h2>Therapeutic Implications: Beyond Motor Applications</h2>
<p>The findings suggest intriguing therapeutic possibilities, particularly for disorders characterized by altered belief updating and confidence calibration. The demonstration that MD disruption leads to overconfident exploitation with reduced information-seeking offers a compelling framework for understanding psychiatric conditions where belief revision goes awry.</p>
<p>Consider schizophrenia, where patients often exhibit <strong>pathological certainty</strong> in delusional beliefs despite contradictory evidence. The current findings suggest a potential mechanism: if MD-PFC circuits that normally regulate confidence in beliefs become dysregulated, patients might lose the capacity for adaptive doubt that would otherwise prompt belief revision. The specific loss of directed exploration observed here—the strategic sampling of information to resolve uncertainty—parallels the clinical observation that individuals with psychosis often fail to seek disconfirming evidence for their beliefs.</p>
<p>This connects to broader hypotheses about uncertainty processing in cognitive control. Rather than viewing delusions simply as &#8220;false beliefs,&#8221; they might reflect a fundamental breakdown in the brain&#8217;s ability to appropriately weight confidence in its own predictions. When the system becomes overconfident in existing representations (as seen post-thalamotomy), it loses the motivation to gather information that might challenge those representations—a hallmark of delusional thinking.</p>
<h2>Broader Significance: Rethinking Thalamic Function</h2>
<p>This work contributes to a fundamental reconceptualization of thalamic function &#8211; from simple relay station to active computational processor. The thalamus isn&#8217;t just routing information; it&#8217;s dynamically modulating the confidence and precision of cortical computations based on behavioral context.</p>
<h2>Personal Reflection: When Theory Meets Unexpected Validation</h2>
<p>For our lab, this study is quite awesome &#8211; when years of circuit-level investigation receive independent validation from an entirely different methodology. The fact that this confirmation came through a clinical study aimed at treating human suffering makes it even more meaningful.</p>
<p>It&#8217;s the rare convergence where theory and evidence transform each other: the theory gains human causal validation, while the evidence gains mechanistic understanding. Together, they point toward a future where we might not just understand the circuits of adaptive decision-making, but actively repair them when they break.</p>
<p>The patients in this study, seeking relief from debilitating tremor, graciously contributed to our understanding of one of the brain&#8217;s most fundamental computations: how to balance confidence with curiosity. Their experience shows us what happens when certainty becomes a cage &#8211; when we lose the capacity to doubt ourselves when doubt would serve us best.</p>
<p><em>The work discussed builds on extensive research into thalamocortical circuits and decision-making, offering new insights into the neural mechanisms underlying adaptive behavior and potential therapeutic applications for disorders of motivation and cognitive flexibility.</em></p>
<p><strong>References:</strong></p>
<ul>
<li>Mackenzie, G., et al. (2025). Focused ultrasound neuromodulation of mediodorsal thalamus disrupts decision flexibility during reward learning. bioRxiv.</li>
<li>Scott, D.N., Mukherjee, A., Nassar, M.R., &amp; Halassa, M.M. (2024). Thalamocortical architectures for flexible cognition and efficient learning. Trends in Cognitive Sciences, 28(7), 639-652.</li>
</ul>
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		<title>The Two Prefrontal Streams Evidence for Homology Across Species</title>
		<link>https://michaelhalassa.net/the-two-prefrontal-streams-evidence-for-homology-across-species/</link>
		
		<dc:creator><![CDATA[michaelhalassa]]></dc:creator>
		<pubDate>Thu, 12 Sep 2024 03:56:16 +0000</pubDate>
				<category><![CDATA[Michael Halassa]]></category>
		<category><![CDATA[Neuroscience]]></category>
		<category><![CDATA[Science]]></category>
		<category><![CDATA[Brain scientist]]></category>
		<category><![CDATA[neuroscience]]></category>
		<category><![CDATA[research paper]]></category>
		<category><![CDATA[studies]]></category>
		<guid isPermaLink="false">https://michaelhalassa.net/?p=722</guid>

					<description><![CDATA[I have recently had the privilege of writing a book chapter with Bob Vertes and Nicola Palemero-Ghallager on the evolution of the frontal cortex. It was an amazing intellectual journey, where I learned a lot from our interactions. The product was a new hypothesis for how this amazing part of our brain has evolved based [&#8230;]]]></description>
										<content:encoded><![CDATA[<p>I have recently had the privilege of writing a book chapter with Bob Vertes and Nicola Palemero-Ghallager on the evolution of the frontal cortex. It was an amazing intellectual journey, where I learned a lot from our interactions. The product was a new hypothesis for how this amazing part of our brain has evolved based on comparative anatomy and function. Please enjoy and reach out with any questions. Any and all input is welcome.</p>
<p><strong>Abstract</strong></p>
<p>The prefrontal cortex (PFC) plays a critical role in human cognition, but the precise mechanisms by which its circuitry accomplishes its proposed functions are unclear. Nonhuman animals are indispensable in revealing such mechanisms, as the ability to monitor and manipulate their circuitry provides necessary insights. A major impediment to linking the growing progress in animal research to insights for human cognition and applications to human health is the lack of consensus on how the PFC is homologous across species. In this perspective, we follow the classifi cation of human PFC into medial and lateral streams, with the medial being primarily evaluative and the lateral being executive. Based on anatomy, physiology and function, we advance the proposal that the rodent medial prefrontal cortex contains elements of both streams, with functional parallels between primate ventromedial and dorsolateral PFC with rodent infralimbic and prelimbic areas, respectively. To support this argument, we highlight the granular nature of the prelimbic cortex in Tupaia belangeri, a basal primate whose PFC macrostructure is rodent-like. Our perspective may help provide additional input to the debate on PFC homology and lead to new testable hypotheses.</p>
<p><strong>Introduction </strong></p>
<p>The prefrontal cortex (PFC) is a complex and highly interconnected region that engages in a wide variety of cognitive functions, including attention, working memory, decision making, and social behavior (Miller and Cohen 2001; Soltani and Koechlin 2022). In the human brain, the PFC has shown great expansion compared to even the closest primate relatives (Preuss and Wise 2022), a process thought to be key to the unparalleled cognitive expansion seen in our species. However, both the principles by which PFC circuits contribute to cognition as well as their origin/emergence are poorly understood.</p>
<p>Nonhuman animal research is poised to help fi ll this knowledge gap because, in addition to its basic scientifi c value, it off ers important insights into human health given the involvement of PFC dysfunction in several neurological and psychiatric illnesses (Liston et al. 2011; Smucny et al. 2022). Given the mechanistic accessibility aff orded by newer monitoring (Tian and Looger 2008; Wu et al. 2022; Xu et al. 2017) and causal tools (Kim et al. 2017; Rabut et al. 2020; Roth 2016), there has been an explosion in PFC animal research over the last decade focused on rodent PFC. Yet despite this progress, it is considerably challenging to relate these advances into insights applicable to understanding the human (and nonhuman primate) PFC given the considerable diff erences in macro- and microarchitecture. Specifi cally, while the human PFC has a large number of well-diff erentiated areas (Haber and Robbins 2022)—von Economo and Koskinas (1925), for example, identifi ed 39 cytoarchitectonically distinct areas on the cortex covering the lateral, medial, and orbital portions of the frontal lobe—the rodent PFC is far less diff erentiated, thus making homology assignments very challenging.</p>
<p>Here, we follow the general two-stream human PFC classifi cation (Domenech and Koechlin 2015) as a starting point. Specifi cally, this functional classifi cation suggests that the lateral stream, which is largely composed of the lateral PFC (lPFC) is involved in executive control and rule-based behavior (Friedman and Robbins 2022). In contrast, the medial stream, which is composed of the ventromedial PFC (vmPFC) and dorsomedial PFC (dmPFC), is involved in adjusting behavioral strategies based on reinforcement and selfmonitoring (Domenech and Koechlin 2015). According to the defi nition of Domenech and Koechlin (2015), the lPFC encompasses Brodmann’s (1909) areas 44 and 45, as well as the lateral portion of areas 8, 9, and 10 (although those authors do not mention areas 46 or 47 which are commonly included in the lateral stream). Their vmPFC covers Brodmann’s areas 11, 12, 14, 25, the medial part of 10, rostral part of 24, and ventral portion of 32, whereas the dmPFC encompasses the caudal and dorsal parts of 24 and 32, respectively, as well as the medial portion of areas 6, 8, and 9.</p>
<p>We present evidence that the rodent medial prefrontal cortex (mPFC) exhibits homology to both streams. Specifi cally, our thesis indicates that the rodent infralimbic cortex (i.e., area IL) is most closely related to the primate vmPFC based on both connectivity and function. On the other hand, the rodent prelimbic cortex (i.e., area PL) exhibits gradients of connectivity that makes it a likely precursor of several regions found in the primate PFC. Specifi cally, the evidence reviewed here supports that PL is a precursor of areas belonging to the primate medial and lateral stream regions such as dmPFC area 32, and dorsolateral PFC ( dlPFC) areas 10, 9, and 8. The notion of a single rodent-like precursor of several primate cortical areas is not new and has been utilized to explain evolutionary expansion and diff erentiation in the sensorimotor system (Kaas 2004). Here, we extend the notion of an evolutionary precursor to prefrontal circuitry, providing a clearer context for relating rodent functional data to primate cognition. Consistent with our proposal, we point to T. belangeri, an evolutionary intermediate whose prelimbic cortex contains an area that is granular, a microcircuit feature that establishes its correspondence to primate dlPFC.</p>
<p><strong>The Prelimbic Cortex As a Precursor of Dorsomedial and Dorsolateral Prefrontal Cortex</strong></p>
<p>The cerebral cortex has undergone signifi cant changes and diff erentiations throughout evolution, providing space for the development of distinct cortical areas with specialized functions. The evolution of somatomotor control, for example, from simple refl exive movements to highly coordinated and precise voluntary actions, is associated with a signifi cant cortical expansion and segregation as well as neuronal specialization. Indeed, the Bauplan of the brain of opossums resembles that of small-brained placental mammals in all but one aspect: it contains a “somatosensory-motor amalgam,” with a complete overlap of somatosensory representation and motor control maps (Dooley et al. 2014; Karlen and Krubitzer 2007; Wong and Kaas 2009a). Since marsupials diverged from placental mammals around 130 million years ago, Kaas (2004) proposed that this somatosensory-motor amalgam could be considered a “precursor area” of the architectonically distinct sensory and motor areas found in the brains of the latter infraclass. Small placental mammals, including tenrecs (Krubitzer et al. 1997), hedgehogs (Catania et al. 2000), or rats (Haghir et al. 2023), present a distinct primary motor cortex (M1), and in most cases their somatosensory region encompasses four areas: a primary (S1) and a secondary (S2) somatosensory area as well as rostral and caudal somatosensory belt areas. A secondary motor area has also been described in the rat brain, and some of these species present a further somatosensory area located ventrocaudally to S2 (for a comprehensive review, see Kaas 2004). In addition to these two motor and fi ve somatosensory areas, the brain of tree shrews (the closest relatives of primates) presents a rudimentary somatosensory posterior parietal area (Wong and Kaas 2009a). A further diff erentiation occurs in the brains of small primates such as galagos (Wu and Kaas 2003) and slow lorises (Carlson and Fitzpatrick 1982), which display additional somatosensory areas located in the lateral fi ssure. In macaque monkeys, but not in marmosets, the caudal somatosensory belt area developed further into areas 1 and 2 (Kaas 2004), and three subfi elds can be identifi ed within M1 (Rapan et al. 2023). This cortical segregation reaches its apex in humans, where both the motor and somatosensory cortex have expanded signifi cantly in terms of size and complexity to enable fi ner control of movements, including intricate fi nger and hand movements, as well as the production of speech, and enhance the individual’s capacity for motor planning and decision making. The gradual changes in cytoarchitecture associated with the phylogenetically related emergence of multiple areas from the marsupial somatosensory-motor amalgam are in line with the “gradation theory” postulated by Sanides (1962) to explain cortical diff erentiation in the human PFC. Specifi cally, his systematic analysis revealed that segregation in the human PFC is associated with discontinuous step-wise changes of cytoarchitectonic features which not only follow phylogenetically related cortical expansion (i.e., when moving medio-laterally from allocortical through mesocortical to neocortical areas), but also when moving in a poleward direction throughout the prefrontal neocortex (Sanides 1962). Below, we present both structural and functional evidence in support of the framework that rodent area PL could be considered a precursor of primate dmPFC area 32 and of areas belonging to the primate dlPFC.</p>
<p><strong>Structural Studies</strong></p>
<p>The prelimbic cortex occupies a very large area of the prefrontal cortex in rodents. In rats, the PL extends rostro-caudally for about 3 mm, from the anterior pole of the PFC, sitting above the medial orbital cortex, to caudally situated dorsal to IL (Swanson 2004). While PL has generally been regarded as a single entity, recent evidence leads us to propose that PL may anatomically and functionally consist of two major divisions: rostrodorsal and caudoventral divisions. Specifi cally, there are notable anatomical diff erences between these two parts of PL with respect to both their inputs and outputs. For instance, in an early examination of PFC projections to the striatum, Berendse et al. (1992) reported that the dorsal part of PL projected to mid-regions of the dorsal striatum, whereas ventrally PL selectively distributed to the nucleus accumbens (ACB), and we could confirm this distinction.</p>
<p>As is well established, the mediodorsal nucleus (MD) of the thalamus is strongly connected reciprocally with the mPFC. However, the caudoventral PL distributes specifi cally to the medial segment of MD, whereas the rostrodorsal PL projects selectively to the lateral segment of MD (Groenewegen 1988; Vertes 2004). Taken together, this pattern indicates that the rostrodorsal PL communicates primarily with action/premotor-associated structures and may therefore serve a role in executive control, similar to areas of the primate dlPFC. On the other hand, caudoventral PL is strongly interconnected with limbic structures and may accordingly be involved primarily in aff ective behaviors, comparable to those of area 32 of primates. With respect to limbic connections, the caudoventral PL receives pronounced projections from the hippocampus, mainly originating from CA1 and the subiculum of the ventral hippocampus. Thalamic aff erents to this division of PL arise predominantly from medial/central regions of the thalamus including MD (as mentioned above), rostral intralaminar nuclei, and the midline nuclei: the paraventricular, paratenial, rhomboid, and reuniens (RE) nuclei (Hoover and Vertes 2007; Vertes 2004, 2006). Finally, the caudoventral PL receives signifi cant projections from the basal nuclei of the amygdala as well as from monoaminergic nuclei (e.g., dopaminergic, noradrenergic and serotonergic) of the brainstem. It is well recognized that the monoaminergic nuclei exert pronounced modulatory eff ects on PL in aff ective and cognitive functions (Friedman and Robbins 2022).</p>
<p>With some exceptions, the output of caudoventral PL parallels its input (Hoover and Vertes 2007; Vertes 2004). Cortically, this caudoventral PL strongly targets other prefrontal cortical regions, including the medial orbital cortex, the dorsal and ventral agranular insular cortex, the anterior piriform cortex, and the entorhinal cortex. Subcortically, caudoventral PL distributes heavily to (a) the ACB, olfactory tubercle, and claustrum of the basal forebrain; (b) the central and basal nuclei of the amygdala; (c) the MD, intermediodorsal, paraventricular, paratenial, reuniens, and centromedial thalamic nuclei; and (d) the substantia nigra, pars compacta, ventral tegmental area, and dorsal and median raphe nuclei of the midbrain. In summary, the inputs and outputs of the caudoventral PL largely mirror those of area 32 of primates.</p>
<p><strong>Functional Studies</strong></p>
<p>While the debate on the rodent homologue of the dlPFC of primates may never be resolved to everyone’s satisfaction, primates (especially humans) possess abilities that undeniably exceed those of rodents, and this undoubtedly is tied to cortical evolution including that of the dlPFC. Still, it must be acknowledged that rodents exhibit executive functions that are classically attributed to primate dlPFC. In addition to the anatomical evidence discussed above, behavioral evidence suggests that rostrodorsal PL is a “ functional homologue” of primate dlPFC.</p>
<p>Granon and Poucet (2000) were among the fi rst to make this proposal. Specifi cally, they reviewed evidence showing that alterations of PL in rodents (but not other mPFC regions) produced severe impairments on various spatial and nonspatial delay tasks. This indicated a profound working memory defi cit—a hallmark of damage to the dlPFC. The working memory defi cits were part of a constellation of cognitive impairments produced by alterations of PL that included attentional defi cits. In addition, Granon and Poucet pointed out that rostrodorsal PL is reciprocally connected to the lateral subdivision of the MD, paralleling primate dlPFC projections to the lateral MD (Granon and Poucet 2000). Several other studies described similar reciprocal connections between PL and lateral MD in rodents (Bolkan et al. 2017; Mukherjee et al. 2020; Schmitt et al. 2017; Wolff et al. 2008). Granon and Poucet (2000:235) concluded that “in both species [rodents and primates], the prefrontal cortex, seems to share some common function in those aspects of cognitive processing that, in humans, are usually referred to as executive functions. Within the rat prefrontal cortex, the prelimbic area appears to play a central role in such processes.”</p>
<p>Several subsequent reports have confi rmed the role of PL of rodents in working memory and in several additional cognitive functions including attentional processes, set shifting behavior, reversal learning, and decision making (for reviews, see Chudasama 2011; Friedman and Robbins 2022). Specifically, these are all functions that in primates are associated with activation of the dlPFC.</p>
<p>Physiological evidence also supports the idea that the rostrodorsal PL and dlPFC are homologous. Classical work by Fuster, Goldman-Rakic, and others (Funahashi et al. 1993b; Fuster and Alexander 1971) have shown that neurons in the dlPFC exhibit persistent increase in spike rates in the context of working memory, which has been considered to be a cellular correlate for this cognitive process (Fuster and Alexander 1971). Newer studies have corroborated these observations, albeit they emphasize a persistent network activity pattern (rather than individual neurons) and perhaps temporally sparser patterns of working memory correlates at the level of single neurons (Lundqvist et al. 2016). Consistent with these latter observations, and with the PL homology, multiple studies have found evidence for persistent network activity patterns in the context of working memory tasks. For example, Bolkan et al. (2017) found evidence for a sequential PL activity pattern in the context of a spatial working memory task. Interestingly, this activity pattern was not spatially specifi c, potentially refl ective of the PL’s function in the generation of abstract rules, which are a known attribute of dlPFC. This was corroborated by data from Schmitt et al. (2017), who trained mice on a cross-modal attentional control task where mice selected between visual and auditory target stimuli based on a cue that varied on a trial-by-trial basis. Out of several cortical areas inactivated in the PFC, including orbitofrontal cortex, anterior cingulate cortex, and premotor cortex, only the PL showed a delay period specifi c eff ect (Wimmer et al. 2015). Recordings from the PL showed a persistent network activity pattern over the delay, where single neurons exhibited a temporally precise increase in fi ring rate tiling the delay period (sequential activity pattern). These network patterns where “rule specifi c” (Rikhye et al. 2018; Schmitt et al. 2017), consistent with the fi nding from primate dlPFC which showed the highest proportion of neurons encoding abstract rules in working memory tasks (Wallis et al. 2001). Perhaps the most compelling link to the specifi city of these observations to the rostrodorsal PL is the work by Nakajima et al. (2019), which showed that neurons in this particular region project to the dorsal striatum (Figure 3.2a) and exhibit activity patterns consistent with attentional modulation (Figure 3.2b, c).</p>
<p>Lastly, in studying the architectonic subdivisions of the neocortex of the tree shrew, T. belangeri, a close relative of primates, Wong and Kaas (2009a) found that the PL of that species (and which they designated as area MF) contained a well-developed layer 4, which was densely populated with granule cells. This suggests that area PL of rodents, which occupies the same relative position as area MF of tree shrews, dorsally on the medial wall of the PFC, could be the antecedent of the granule cell layer of primates. Consistent with this notion, we show comparative sections of this region across rats, Tupaia, and macaques.</p>
<p><strong>Homology between Infralimbic Cortex and vmPFC</strong></p>
<p>Whereas the rodent homologue to the dlPFC of primates remains controversial, there appears to be a general consensus that ventral parts of the mPFC of rodents are anatomically and functionally equivalent to the agranular ventral medial PFC ( vmPFC) of primates. More specifi cally, area IL of rodents appears anatomically homologous to area 25 (A25) of primates.</p>
<p>For instance, the IL of rodents and A25 of primates serve well-recognized roles in autonomic, visceral, and aff ective functions. IL has been described as a visceromotor cortex. The projections of IL refl ect its involvement in visceral/ aff ective functions. Specifi cally, Vertes (2004) examined IL projections in rats and showed that IL distributes to several sites of the forebrain and brainstemlinked to autonomic and aff ective behavior. These included orbitofrontal cortices, shell of nucleus accumbens (sACB), lateral septum, bed nucleus of stria terminalis (BST), medial and lateral preoptic nuclei, central nucleus of the amygdala, lateral and posterior nuclei of the hypothalamus, and the periaqueductal gray, parabrachial nucleus and solitary nucleus of the brainstem. Each of the structures has been shown to modulate autonomic/visceral activity, and thus emotional behavior, and importantly as a group, these nuclei receive input almost exclusively from IL and little from PL.</p>
<p>Although fewer reports have examined vmPFC (or A25) projections in primates, A25 projections in the monkey appear to directly parallel those of IL in rodents. Specifi cally, an early report by Chiba et al. (2001) compared the eff erent projections of A25 (IL) and A32 (PL) in the Japanese monkey and showed that the output of A25, like that of IL in rodents, strongly targeted sites involved in autonomic/visceral control, primarily including the sACB, the preoptic area, BST, central nucleus of the amygdala (CeM) and the periaqueductal gray and parabrachial nucleus of the brainstem. They thus concluded that their fi ndings “support the hypothesis that IL is a major cortical autonomic motor area.” Several subsequent examinations of A25 projections in monkeys and have similarly demonstrated that A25 prominently distributes to several “visceral-related” subcortical structures of the basal forebrain, amygdala, hypothalamus and brainstem (Barbas et al. 2003; Ghashghaei et al. 2007; Heilbronner et al. 2016; Joyce and Barbas 2018; Rios-Florez et al. 2021; Roberts et al. 2007). Major targets included the ACB, BST, central nucleus of the amygdala, posterior and lateral nuclei of the hypothalamus, periaqueductal gray and parabrachial nucleus.</p>
<p>Barbas et al. (2003) described projections from mPFC in primates, including A25, to discrete nuclei of the amygdala and hypothalamus that directly distribute to (autonomic) brainstem and spinal cord nuclei which innervate peripheral autonomic sites. This system of connections linked mPFC/A25 with autonomic eff ector sites in the modulation of visceral functions and emotional behavior. However, in subsequent studies Barbas and colleagues have suggested that the connections of posterior OFC with the intercalated cell masses of the amygdala more resemble rodent IL, than primate A25 (Zikopoulos et al. 2017).</p>
<p>In contrast, Heilbronner et al. (2016) compared the projections to the striatum from A25 in macaques and IL in rats. Specifi cally, they fi rst identifi ed a region of the sACB (termed the “striatal emotion processing network” or EPN) and conserved across these species. The EPN is a convergence zone of projections from the amygdala and hippocampus to the sACB. Importantly, they showed that both IL and A25 distributed heavily to the striatal EPN, whereas other prefrontal cortical areas (of both species) projected at best weakly to EPN. They concluded that “consistent with prior literature, the infralimbic cortex and area 25 are likely homologous” (Heilbronner et al. 2016:509). Future studies should perform whole brain connectivity fi ngerprints across species for a more comprehensive comparison. However, it should be noted that even if rodent IL and primate A25 show overall similar connectivity patterns, the evolutionary expansion of the PFC may endow primate A25 with unique interregional connectivity patterns and divergent functions.</p>
<p>Recently, Roberts and colleagues (Alexander et al. 2023) comprehensively reviewed the structural and functional properties of the vmPFC across species (rat, monkey, human) and cited evidence showing that (a) the IL of rats and A25 of primates show some functional homology/analogy in the regulation of behavior in the reward domain but not in the punishment domain. Specifi cally, they showed that A25 overactivation in marmosets blunted Pavlovian approach and motivated responding, comparable to that reported following similar manipulations in rodents. In marked contrast, the same manipulation heightened behavioral and cardiovascular responsivity to both proximal and distal threat, opposite to that reported in rodent IL. This suggests that IL and A25 may act similarly within reward networks but their roles may have diverged within threat networks illustrating the complexity of cross-species functional comparisons. Roberts and colleagues also showed (b) that IL/A25 and PL/ A32 predominantly serve distinct and separable functions, with A25 mainly involved in cardiovascular and aff ective functions and A32 in cognitive functions. A cytoarchitectonically informed meta-analysis of functional imaging studies in humans provides further evidence for this functional segregation of A25 and A32 (Palomero-Gallagher et al. 2015). For instance, with respect to diff erences between A25 and A32, Wallis et al. (2017) demonstrated that inactivation of A25 produced pronounced cardiovascular changes, whereas inactivation of A32 had no cardiovascular eff ects, and further that A25 and A32 mediated opposite eff ects on a Pavlovian fear conditioning and extinction paradigm: A25 inactivation decreased fear-elicited behavior responses promoting extinction, whereas A32 inactivation enhanced these responses thereby suppressing extinction.</p>
<p>Lastly, Diehl and Redish (2023) have performed comprehensive recordings across the rat mPFC in the context of a foraging task termed “restaurant row.” This task combines multiple cognitive elements including associative learning, working memory, switching, and value-based judgments. Although they found that all prefrontal areas encode the various relevant task variables, there was clear specialization, with the IL clearly encoding more value-related cognitive variables than executive or sensorimotor ones. This is consistent with an earlier report, in which Hardung et al. (2017) examined the neural substrates for response inhibition across areas of the rodent frontal cortex using both optogenetic inactivation and electrophysiological recordings. Strikingly, inactivation of the PL and IL had opposite eff ects on the behavior, where PL inactivation increased and IL inactivation decreased premature responses. Electrophysiological recordings were also consistent with opposing roles for these two subregions, again, consistent with the idea that PL shares functional homology with the primate lateral stream whereas the IL is medial (and evaluative).</p>
<p><strong>Conclusions</strong></p>
<p>Building on the two-stream notion of human (or generally primate) PFC, the collective evidence reviewed in this chapter argues for homology with the two major divisions of rodent PFC: the PL and IL. The argument implicitly makes a prediction about how the rostrodorsal PL may have disconnected from the IL throughout evolution, and subsequently pushed laterally to form what is currently recognized as dlPFC of primates. The fact that T. belangeri MF is granular is consistent with this idea. Overall, we hope this synthesis will stimulate further discussion and motivate the design of new experiments to test this hypothesis directly.</p>
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