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Evolution of Integrated Causal Structures in Animats Exposed to Environments of Increasing Complexity
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Natural selection favors the evolution of brains that can capture fitness-relevant features of the environment's causal structure. We investigated the evolution of small, adaptive logic-gate networks ("animats") in task environments where falling blocks of different sizes have to be caught or avoided in a 'Tetris-like' game. Solving these tasks requires the integration of sensor inputs and memory. Evolved networks were evaluated using measures of information integration, including the number of evolved concepts and the total amount of integrated conceptual information. The results show that, over the course of the animats' adaptation, i) the number of concepts grows; ii) integrated conceptual information increases; iii) this increase depends on the complexity of the environment, especially on the requirement for sequential memory. These results suggest that the need to capture the causal structure of a rich environment, given limited sensors and internal mechanisms, is an important driving force for organisms to develop highly integrated networks ("brains") with many concepts, leading to an increase in their internal complexity.
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![Figure 2 Assessing the causal structure of an animat in a state. (A) A hypothetical animat brain comprised of a logic-gate network with 2 sensors (S1S2), 4 hidden elements (ABCD), and 2 motors (M1M2) is analyzed for illustration in state 00-1010-10. (B) First, the power set of all candidate concepts in the entire animat brain is evaluated. Note that the sensors and motors cannot give rise to concepts or be part of higher order concepts since – by design - they either lack causes or effects (i.e., inputs or outputs) within the system. Each animat brain can thus maximally have 24−1 = 15 concepts (the power-set of the 4 hidden elements, excluding the empty set). “Small phi” φ measures how irreducible a mechanism's cause-effect repertoire is over a particular set of inputs and outputs. φMax is the integrated information of the most irreducible cause-effect repertoire of the mechanism. The number of concepts and ΣφMax are measures of all the brain's causal relations and their strength, both modular or feed-forward and integrated. Here, 6 concepts exist, 4 elementary concepts ([A], [B], [C], [D]) and 2 higher order concepts ([AB], [AC]). All other higher order mechanisms are reducible (φMax = 0). (C) Second, Φ (“big phi”) is evaluated for all subsets of the system (candidate complexes). Φ measures how integrated a set of elements is. It quantifies how much the concepts of the set of elements change under a unidirectional partition between elements (for example, “noising” the connections from A to the rest of the system, leaving the connections to A from the system intact, see Methods). During the analysis, elements outside of the candidate complex are taken as fixed background conditions and remain unperturbed. Note that all subsets that contain either a sensor or a motor have Φ = 0, because elements that are connected to the rest of the system in a feed-forward manner cannot be part of an integrated system (see Methods). An animat's main complex can thus contain at most the 4 hidden elements. (D) Of all subsets of elements, in this particular system state, ABC is maximally integrated (ΦMax = 0.92) and thus forms the main complex (MC). Gray arrows denote fixed background conditions, blue arrows denote functional connections within the MC. (E) Out of the power-set of ABC (maximally 23−1 = 7 possible concepts), the MC specifies 4 irreducible concepts. The number of elements of the main complex, the number of MC concepts, and ΦMax measure different aspects of how integrated the animat's brain is. For each animat at a particular generation, the analysis is performed for every state of the animat's brain, while the animat is performing its particular task. The state-dependent values are then averaged, weighted by the probability of occurrence of each state over 128 trials of different blocks falling.](/paper-figures/2958b2e2c8e04832a6387928bf55d743/evolution-of-integrated-causal-structures-in-animats-exposed-to-environments-of-figure-2.jpg)






![Figure 9 The information, integration, and exclusion postulate applied at the level of mechanisms (A–C) and systems of mechanisms (D–F). (A–F) Each node is a binary logic-gate mechanism that can be in either state ‘0’ (white) or ‘1’ (yellow). The logic-gates and their connections are represented as neural circuits rather than electronic circuits: directed connections between the nodes indicate the inputs and outputs of the logic gates. The mechanisms labeled A, B, and C correspond to system ABC = 101 shown in (D). (A) Information: Mechanism C in its current state ‘1’ generates information as it constrains its causes (the past states of its inputs AB) and effects (the future states of its outputs AB) compared to their unconstrained distributions (gray distribution). Past and future nodes whose state is unspecified are shown in gray. (B) Integration: The elements X and Y do not form an integrated higher order mechanism, since XY is reducible to its component mechanisms X and Y (φ = 0). However, the elements AB in state ‘10’ do form a higher order mechanism, since AB specifies both, irreducible causes and irreducible effects (the minimum information partition (MIP) on both, the cause and effect side leads to a loss of information). Integrated information φ of AB = 10 is evaluated as the minimum of the cause and effect integrated information: φ = min(φCause, φEffect), here φ = φEffect = 0.25, taking all inputs and outputs of AB into account. The overall MIP of AB over all its inputs and outputs is thus MIPEffect, labeled in red. (C) Exclusion: Of all input-output combinations of mechanism AB, the “concept” of AB = 10 is its maximally irreducible cause repertoire, here over all input elements ABC (φCause = 0.33, same as in (B)), together with its maximally irreducible effect repertoire, here over output element C only (φEffect = 0.5). This means that AB has its maximally irreducible effect repertoire specified on C, not on ABC or any other output combination. The concept's integrated information is φMax = min(φCause, φEffect) = φCause = 0.33, its overall MIP is MIPCause, labeled in red. (D) System information: The system ABC = 101 gives rise to a conceptual structure with 4 concepts. (E) System integration: The system WXYZ is reducible into the subsets WX and YZ. WXYZ cannot exist as a system from the intrinsic perspective. By contrast, system ABC is irreducible. Its minimum information partition (MIP) leaves the concepts of A and B intact, but destroys concepts C and AB. Integrated conceptual information Φ(ABC) is evaluated as the difference between the whole conceptual structure C and the partitioned conceptual structure CMIP (see Text S2 in [15]). (F) System exclusion: Of all sets of elements in this larger system, the set ABC has ΦMax and thus forms the main “complex”. ABCD, for example, also specifies integrated conceptual information Φ, but cannot form another complex since it overlaps with ABC and Φ(ABC)> Φ(ABCD) (see Fig. 2).](/paper-figures/5c919159aac24fc487418529d4b66063/evolution-of-integrated-causal-structures-in-animats-exposed-to-environments-of-figure-9.jpg)
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“Albantakis et al. (2014) analyzed networks with 3-6 nodes.”
From Can "consciousness" be observed from large language model (LLM) internal states? Dissecting LLM representations obtained from Theory of Mind test with Integrated Information Theory and Span Representation analysis (2025) · cites this paper as Albantakis et al., 2014
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Citations
Cites
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Integrated information theory: from consciousness to its physical substrate
Tononi, G., Boly, M., Massimini, M., & Koch, C. (2016) · Nature Reviews Neuroscience · doi:10.1038/nrn.2016.44 · Cited by 1,727
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Consciousness: here, there and everywhere?
Tononi, G., & Koch, C. (2015) · Philosophical Transactions of the Royal Society B: Biological Sciences · doi:10.1098/rstb.2014.0167 · Cited by 811
Albantakis, L., Barbosa, L., Findlay, G., Grasso, M., Haun, A. M., Marshall, W., Mayner, W. G. P., Zaeemzadeh, A., Boly, M., Juel, B. E., et al. (2023) · PLoS Computational Biology · doi:10.1371/journal.pcbi.1011465 · Cited by 292
“Yet another is the study of the evolution of a substrate’s causal powers as conditional probabilities that update themselves [74].”
Can the macro beat the micro? Integrated information across spatiotemporal scales
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Metadata
- Authors
- Albantakis, L., Hintze, A., Koch, C., Adami, C., & Tononi, G.
- Affiliations
- University of Wisconsin–Madison (US)
- Michigan State University (US)
- Corresponding author
- Albantakis, Larissa
- Tononi, Giulio
- Venue
- PLoS Computational Biology
- Publisher
- Public Library of Science (PLoS)
- Published
- 2014-12-18
- Volume
- 10
- Issue
- 12
- Pages
- e1003966
- ISSN
- 1553-734X (ISSN-L)
- 1553-7358
- PMC
- PMC4270440
- OpenAlex
- W2070484080
- Cited by
- 101
- References
- 61
- Citations by year
- 2026: 8 · 2025: 7 · 2024: 10 · 2023: 5 · 2022: 8 · 2021: 12 · 2020: 10 · 2019: 12 · 2018: 10 · 2017: 8 · 2016: 6 · 2015: 5
- FWCI
- 6.31
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- PLoS Computational Biology (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- Diva portal (Dalarna University Library) (submittedVersion, other-oa)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Figshare (submittedVersion, cc-by)
- Funders
- Paul G. Allen Family Foundation
- Templeton World Charity Foundation
- Defense Advanced Research Projects Agency
- Topics
- Physical Sciences › Computer Science › Artificial Intelligence › Evolutionary Algorithms and Applications
- Life Sciences › Neuroscience › Cognitive Neuroscience › Embodied and Extended Cognition
- Physical Sciences › Computer Science › Artificial Intelligence › Psychiatry, Mental Health, Neuroscience
- Keywords
- animats, causal structure, information integration, conceptual information, environmental complexity, sequential memory, sensor integration, cognitive evolution, internal complexity
- MeSH terms
- Adaptation, Physiological, Algorithms, Computer Simulation, Biological Evolution, Models, Neurological, Selection, Genetic, Statistics, Nonparametric, Computational Biology, Feedback, Physiological, Genetic Fitness
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