Seven properties of self-organization in the human brain

The principle of self-organization has acquired a fundamental significance in the newly emerging field of computational philosophy. Self-organizing systems have been described in various domains in science and philosophy including physics, neuroscience, biology and medicine, ecology, and sociology. While system architecture and their general purpose may depend on domain-specific concepts and definitions, there are (at least) seven key properties of self-organization clearly identified in brain systems: 1) modular connectivity, 2) unsupervised learning, 3) adaptive ability, 4) functional resiliency, 5) functional plasticity, 6) from-local-to-global functional organization, and 7) dynamic system growth. These are defined here in the light of insight from neurobiology, cognitive neuroscience and Adaptive Resonance Theory (ART), and physics to show that self-organization achieves stability and functional plasticity while minimizing structural system complexity. A specific example informed by empirical research is discussed to illustrate how modularity, adaptive learning, and dynamic network growth enable stable yet plastic somatosensory representation for human grip force control. Implications for the design of “strong” artificial intelligence in robotics are brought forward

The Laminar Organization of Visual Cortex: A Unified View of Development, Learning, and Grouping

Why are all sensory and cognitive neocortex organized into layered circuits? How do these layers organize circuits that form functional columns in cortical maps? How do bottom-up, top-down, and horizontal interactions within the cortical layers generate adaptive behaviors. This chapter summarizes an evolving neural model which suggests how these interactions help the visual cortex to realize: (1) the binding process whereby cortex groups distributed data into coherent object representations; (2) the attentional process whereby cortex selectively processes important events; and (3) the developmental and learning processes whereby cortex shapes its circuits to match environmental constraints. It is suggested that the mechanisms which achieve property (3) imply properties of (I) and (2). New computational ideas about feedback systems suggest how neocortex develops and learns in a stable way, and why top-down attention requires converging bottom-up inputs to fully activate cortical cells, whereas perceptual groupings do not.Defense Advanced Research Projects Agency and the Office of Naval Research (N00014-95-1-0409); National Science Foundation (IRI-97-20333); Office of Naval Research (N00014-95-1-0657

Prä- und postnatale Entwicklung topographischer Transformationen im Gehirn

This dissertation connects two independent fields of theoretical neuroscience: on the one hand, the self-organization of topographic connectivity patterns, and on the other hand, invariant object recognition, that is the recognition of objects independently of their various possible retinal representations (for example due to translations or scalings). The topographic representation is used in the presented approach, as a coordinate system, which then allows for the implementation of invariance transformations. Hence this study shows, that it is possible that the brain self-organizes before birth, so that it is able to invariantly recognize objects immediately after birth. Besides the core hypothesis that links prenatal work with object recognition, advancements in both fields themselves are also presented. In the beginning of the thesis, a novel analytically solvable probabilistic generative model for topographic maps is introduced. And at the end of the thesis, a model that integrates classical feature-based ideas with the normalization-based approach is presented. This bilinear model makes use of sparseness as well as slowness to implement "optimal" topographic representations. It is therefore a good candidate for hierarchical processing in the brain and for future research.Die vorliegende Arbeit verbindet zwei bisher unabhängig untersuchte Gebiete der theoretischen Neurowissenschaften: zum Einen die vorgeburtliche Selbstorganisation topographischer Verbindungsstrukturen und zum Anderen die invariante Objekterkennung, das heisst, die Erkennung von Objekten trotz ihrer mannigfaltigen retinalen Darstellungen (zum Beispiel durch Verschiebungen oder Skalierungen). Die topographische Repräsentierung wird hierbei während der Selbstorganisation als Koordinatensystem genutzt, um Invarianztransformationen zu implementieren. Dies zeigt die Möglichkeit auf, dass sich das Gehirn bereits vorgeburtlich detailliert selbstorganisieren kann, um nachgeburtlich sofort invariant Erkennen zu können. Im Detail führt Kapitel 2 in ein neues, probabilistisch generatives und analytisch lösbares Modell zur Ontogenese topographischer Transformationen ein. Dem Modell liegt die Annahme zugrunde, dass Ausgabezellen des Systems nicht völlig unkorreliert sind, sondern eine a priori gegebene Korrelation erreichen wollen. Da die Eingabezellen nachbarschaftskorreliert sind, hervorgerufen durch retinale Wellen, ergibt sich mit der Annahme rein erregender Verbindungen eine eindeutige topographische synaptische Verbindungsstruktur. Diese entspricht der bei vielen Spezies gefundenen topographischen Karten, z.B. der Retinotopie zwischen der Retina und dem LGN, oder zwischen dem LGN und dem Neokortex. Kapitel 3 nutzt eine abstraktere Formulierung des Retinotopiemechanismus, welche durch adiabitische Elimination der Aktivitätsvariablen erreicht wird, um den Effekt retinaler Wellen auf ein Modell höherer kortikaler Informationsverarbeitung zu untersuchen. Zu diesem Zweck wird der Kortex vereinfacht als bilineares Modell betrachtet, um einfache modulatorische Nichtlinearitäten mit in Betracht ziehen zu können. Zusätzlich zu den Ein- und Ausgabezellen kommen in diesem Modell Kontrolleinheiten zum Einsatz, welche den Informationsfluss aktiv steuern können und sich durch Wettbewerb und pränatalem Lernen auf verschiedene Muster retinaler Wellen spezialisieren. Die Ergebnisse zeigen, dass die entstehenden Verbindungsstrukturen affinen topographischen Abbildungen (insbesondere Translation, Skalierung und Orientierung) entsprechen, die nach Augenöffnen invariante Erkennung ermöglichen, da sie Objekte in der Eingabe in eine normalisierte Repräsentierung transformieren können. Das Modell wird für den eindimensionalen Fall ausführlich analysiert und die Funktionalität für den biologisch relevanteren zweidimensionalen Fall aufgezeigt. Kapitel 4 verallgemeinert das bilineare Modell des dritten Kapitels zu einem mehrschichtigen Modell, die shifter curcuits''. Diese ermöglichen eine logarithmisch in der Anzahl der Eingabezellen wachsende Anzahl an Synapsen, statt einer prohibitiv quadratischen Anzahl. Ausgenutzt wird die Orthogonalität von Translationen im Raum der Verbindungsstrukturen um diese durch harten Wettbewerb an einzelnen Synapsen zu organisieren. Neurobiologisch ist dieser Mechanismus durch Wettbewerb um einen wachstumsregulierenden Transmitter realisierbar. Kapitel 5 nutzt Methoden des probabilistischen Lernens, um das bilineare Modell auf das Lernen von optimalen Repräsentation der Eingabestatistiken zu optimieren. Da statistischen Methoden zweiter Ordnung, wie zum Beispiel das generative Modell aus Kapitel 2, keine lokalisierten rezeptiven Felder ermöglichen und somit keine (örtliche) Topographie möglich ist, wird sparseness'' verwendet um statistischen Abhängigkeiten höherer Ordnung zu lernen und gleichzeitig Topographie zu implementieren. Anwendungen des so formulierten Modells auf natürliche Bilder zeigen, dass lokalisierte, bandpass filternde rezeptive Felder entstehen, die primären kortikalen rezeptiven Feldern stark ähneln. Desweiteren entstehen durch die erzwungene Topographie Orientierungs- und Frequenzkarten, die ebenfalls kortikalen Karten ähneln. Eine Untersuchung des Modells mit zusätzlicher slowness'' der Ausgabezellen und in zeitlicher Nähe gezeigten transformierten natürlichen Eingabemustern zeigt, dass verschiedene Kontrolleinheiten konsistente und den Eingabetransformationen entsprechende rezeptive Felder entwickeln und somit invariante Darstellungen bezüglich der gezeigten Eingaben entwickeln

Towards a Unified Theory of Neocortex: Laminar Cortical Circuits for Vision and Cognition

A key goal of computational neuroscience is to link brain mechanisms to behavioral functions. The present article describes recent progress towards explaining how laminar neocortical circuits give rise to biological intelligence. These circuits embody two new and revolutionary computational paradigms: Complementary Computing and Laminar Computing. Circuit properties include a novel synthesis of feedforward and feedback processing, of digital and analog processing, and of pre-attentive and attentive processing. This synthesis clarifies the appeal of Bayesian approaches but has a far greater predictive range that naturally extends to self-organizing processes. Examples from vision and cognition are summarized. A LAMINART architecture unifies properties of visual development, learning, perceptual grouping, attention, and 3D vision. A key modeling theme is that the mechanisms which enable development and learning to occur in a stable way imply properties of adult behavior. It is noted how higher-order attentional constraints can influence multiple cortical regions, and how spatial and object attention work together to learn view-invariant object categories. In particular, a form-fitting spatial attentional shroud can allow an emerging view-invariant object category to remain active while multiple view categories are associated with it during sequences of saccadic eye movements. Finally, the chapter summarizes recent work on the LIST PARSE model of cognitive information processing by the laminar circuits of prefrontal cortex. LIST PARSE models the short-term storage of event sequences in working memory, their unitization through learning into sequence, or list, chunks, and their read-out in planned sequential performance that is under volitional control. LIST PARSE provides a laminar embodiment of Item and Order working memories, also called Competitive Queuing models, that have been supported by both psychophysical and neurobiological data. These examples show how variations of a common laminar cortical design can embody properties of visual and cognitive intelligence that seem, at least on the surface, to be mechanistically unrelated.National Science Foundation (SBE-0354378); Office of Naval Research (N00014-01-1-0624

How does the Cerebral Cortex Work? Learning, Attention, and Grouping by the Laminar Circuits of Visual Cortex

The organization of neocortex into layers is one of its most salient anatomical features. These layers include circuits that form functional columns in cortical maps. A major unsolved problem concerns how bottom-up, top-down, and horizontal interactions are organized within cortical layers to generate adaptive behaviors. This article models how these interactions help visual co1tex to realize: (I) the binding process whereby cortex groups distributed data into coherent object representations; (2) the attentional process whereby cortex selectively processes important events; and (3) the developmental and learning processes whereby cortex shapes its circuits to match environmental constraints. New computational ideas about feedback systems suggest how neocortex develops and learns in a stable way, and why top-down attention requires converging bottom-up inputs to fully activate cortical cells, whereas perceptual groupings do not.Defense Advanced Research Projects Agency; National Science Foundation (IRI-97-20333); Office of Naval Research (N00014-95-1-0409, N00014-95-1-0657

Context-Sensitive Binding by the Laminar Circuits of V1 and V2: A Unified Model of Perceptual Grouping, Attention, and Orientation Contrast

A detailed neural model is presented of how the laminar circuits of visual cortical areas V1 and V2 implement context-sensitive binding processes such as perceptual grouping and attention. The model proposes how specific laminar circuits allow the responses of visual cortical neurons to be determined not only by the stimuli within their classical receptive fields, but also to be strongly influenced by stimuli in the extra-classical surround. This context-sensitive visual processing can greatly enhance the analysis of visual scenes, especially those containing targets that are low contrast, partially occluded, or crowded by distractors. We show how interactions of feedforward, feedback and horizontal circuitry can implement several types of contextual processing simultaneously, using shared laminar circuits. In particular, we present computer simulations which suggest how top-down attention and preattentive perceptual grouping, two processes that are fundamental for visual binding, can interact, with attentional enhancement selectively propagating along groupings of both real and illusory contours, thereby showing how attention can selectively enhance object representations. These simulations also illustrate how attention may have a stronger facilitatory effect on low contrast than on high contrast stimuli, and how pop-out from orientation contrast may occur. The specific functional roles which the model proposes for the cortical layers allow several testable neurophysiological predictions to be made. The results presented here simulate only the boundary grouping system of adult cortical architecture. However we also discuss how this model contributes to a larger neural theory of vision which suggests how intracortical and intercortical feedback help to stabilize development and learning within these cortical circuits. Although feedback plays a key role, fast feedforward processing is possible in response to unambiguous information. Model circuits are capable of synchronizing quickly, but context-sensitive persistence of previous events can influence how synchrony develops. Although these results focus on how the interblob cortical processing stream controls boundary grouping and attention, related modeling of the blob cortical processing stream suggests how visible surfaces are formed, and modeling of the motion stream suggests how transient responses to scenic changes can control long-range apparent motion and also attract spatial attention.Defense Advanced Research Projects agency and the Office of Naval Research (N00014-95-1-0409); National Science Foundation (IRI 94-01659, IRI 97-20333); ONR (N00014-92-J-1309, N00014-95-1-0657

Sparse Coding with a Somato-Dendritic Rule

© 2020 Elsevier Ltd. All rights reserved. This manuscript is licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International Licence http://creativecommons.org/licenses/by-nc-nd/4.0/.Cortical neurons are silent most of the time. This sparse activity is energy efficient, and the resulting neural code has favourable properties for associative learning. Most neural models of sparse coding use some form of homeostasis to ensure that each neuron fires infrequently. But homeostatic plasticity acting on a fast timescale may not be biologically plausible, and could lead to catastrophic forgetting in embodied agents that learn continuously. We set out to explore whether inhibitory plasticity could play that role instead, regulating both the population sparseness and the average firing rates. We put the idea to the test in a hybrid network where rate-based dendritic compartments integrate the feedforward input, while spiking somas compete through recurrent inhibition. A somato-dendritic learning rule allows somatic inhibition to modulate nonlinear Hebbian learning in the dendrites. Trained on MNIST digits and natural images, the network discovers independent components that form a sparse encoding of the input and support linear decoding. These findings con-firm that intrinsic plasticity is not strictly required for regulating sparseness: inhibitory plasticity can have the same effect, although that mechanism comes with its own stability-plasticity dilemma. Going beyond point neuron models, the network illustrates how a learning rule can make use of dendrites and compartmentalised inputs; it also suggests a functional interpretation for clustered somatic inhibition in cortical neurons.Peer reviewe

Linking the Laminar Circuits of Visual Cortex to Visual Perception

A detailed neural model is being developed of how the laminar circuits of visual cortical areas V1 and V2 implement context-sensitive binding processes such as perceptual grouping and attention, and develop and learn in a stable way. The model clarifies how preattentive and attentive perceptual mechanisms are linked within these laminar circuits, notably how bottom-up, top-down, and horizontal cortical connections interact. Laminar circuits allow the responses of visual cortical neurons to be influenced, not only by the stimuli within their classical receptive fields, but also by stimuli in the extra-classical surround. Such context-sensitive visual processing can greatly enhance the analysis of visual scenes, especially those containing targets that are low contrast, partially occluded, or crowded by distractors. Attentional enhancement can selectively propagate along groupings of both real and illusory contours, thereby showing how attention can selectively enhance object representations. Model mechanisms clarify how intracortical and intercortical feedback help to stabilize cortical development and learning. Although feedback plays a key role, fast feedforward processing is possible in response to unambiguous information.Defense Advanced Research Projects Agency and the Office of Naval Research (N00014-95-1-0409); National Science Foundation (IRI-97-20333); Office of Naval Research (N00014-95-1-0657

Refinement and Pattern Formation in Neural Circuits by the Interaction of Traveling Waves with Spike-Timing Dependent Plasticity

Traveling waves in the developing brain are a prominent source of highly correlated spiking activity that may instruct the refinement of neural circuits. A candidate mechanism for mediating such refinement is spike-timing dependent plasticity (STDP), which translates correlated activity patterns into changes in synaptic strength. To assess the potential of these phenomena to build useful structure in developing neural circuits, we examined the interaction of wave activity with STDP rules in simple, biologically plausible models of spiking neurons. We derive an expression for the synaptic strength dynamics showing that, by mapping the time dependence of STDP into spatial interactions, traveling waves can build periodic synaptic connectivity patterns into feedforward circuits with a broad class of experimentally observed STDP rules. The spatial scale of the connectivity patterns increases with wave speed and STDP time constants. We verify these results with simulations and demonstrate their robustness to likely sources of noise. We show how this pattern formation ability, which is analogous to solutions of reaction-diffusion systems that have been widely applied to biological pattern formation, can be harnessed to instruct the refinement of postsynaptic receptive fields. Our results hold for rich, complex wave patterns in two dimensions and over several orders of magnitude in wave speeds and STDP time constants, and they provide predictions that can be tested under existing experimental paradigms. Our model generalizes across brain areas and STDP rules, allowing broad application to the ubiquitous occurrence of traveling waves and to wave-like activity patterns induced by moving stimuli

Molecular mechanisms of synaptic plasticity in adult mammalian sensory cortex

Experience-dependent changes in synaptic composition and function (synaptic plasticity) underlie many brain functions including learning and memory, formation of sensory maps, as well as the capability to recover from injury. Most of these functions decline with age, supporting the observation that synaptic plasticity is greater in juveniles than in adults. However, it has been known for some time that peripheral deafferentation in adult animals can induce large-scale reorganization of sensory cortex, which suggests that the adult cortex retains a level of synaptic plasticity that is typically masked by normal activity inputs. Here I present a series of experiments in the adult mammalian sensory cortex that examine the mechanisms and methods to enhance experience-dependent synaptic plasticity. The results showed: 1. Peripheral deafferentation of a single digit in the adult raccoon induces progressive modulation of excitation and inhibition in the deafferented somatosensory cortex that might be needed for the reorganization of receptive fields; 2. Visual deprivation in adult rats reverses three molecular mechanisms that have been correlated with the decrease in synaptic plasticity in adult cortex. These include the developmental increase in the level of inhibition relative to excitation, the development switch in NMDA receptor subunit composition from NR2b to NR2a, and the developmental decrease in tPA activity in visual cortex; 3.Visual deprivation rejuvenates ocular dominance plasticity in the adult visual cortex, and promotes the functional recovery of an eye deprived of vision from birth in adulthood. This work provides further understanding of the molecular mechanisms underlying experience-dependent cortical plasticity. In addition, it demonstrates that ocular dominance plasticity can be regulated throughout life, and proposes a non-invasive method to enhance synaptic plasticity in adult cortex