Hebbian fast plasticity and working memory

Theories and models of working memory (WM) were at least since the mid-1990s dominated by the persistent activity hypothesis. The past decade has seen rising concerns about the shortcomings of sustained activity as the mechanism for short-term maintenance of WM information in the light of accumulating experimental evidence for so-called activity-silent WM and the fundamental difficulty in explaining robust multi-item WM. In consequence, alternative theories are now explored mostly in the direction of fast synaptic plasticity as the underlying mechanism.The question of non-Hebbian vs Hebbian synaptic plasticity emerges naturally in this context. In this review we focus on fast Hebbian plasticity and trace the origins of WM theories and models building on this form of associative learning.Comment: 12 pages, 2 figures, 1 box, submitte

The Performance of Associative Memory Models with Biologically Inspired Connectivity

This thesis is concerned with one important question in artificial neural networks, that is, how biologically inspired connectivity of a network affects its associative memory performance. In recent years, research on the mammalian cerebral cortex, which has the main responsibility for the associative memory function in the brains, suggests that the connectivity of this cortical network is far from fully connected, which is commonly assumed in traditional associative memory models. It is found to be a sparse network with interesting connectivity characteristics such as the “small world network” characteristics, represented by short Mean Path Length, high Clustering Coefficient, and high Global and Local Efficiency. Most of the networks in this thesis are therefore sparsely connected. There is, however, no conclusive evidence of how these different connectivity characteristics affect the associative memory performance of a network. This thesis addresses this question using networks with different types of connectivity, which are inspired from biological evidences. The findings of this programme are unexpected and important. Results show that the performance of a non-spiking associative memory model is found to be predicted by its linear correlation with the Clustering Coefficient of the network, regardless of the detailed connectivity patterns. This is particularly important because the Clustering Coefficient is a static measure of one aspect of connectivity, whilst the associative memory performance reflects the result of a complex dynamic process. On the other hand, this research reveals that improvements in the performance of a network do not necessarily directly rely on an increase in the network’s wiring cost. Therefore it is possible to construct networks with high associative memory performance but relatively low wiring cost. Particularly, Gaussian distributed connectivity in a network is found to achieve the best performance with the lowest wiring cost, in all examined connectivity models. Our results from this programme also suggest that a modular network with an appropriate configuration of Gaussian distributed connectivity, both internal to each module and across modules, can perform nearly as well as the Gaussian distributed non-modular network. Finally, a comparison between non-spiking and spiking associative memory models suggests that in terms of associative memory performance, the implication of connectivity seems to transcend the details of the actual neural models, that is, whether they are spiking or non-spiking neurons

A Cortical Attractor Network with Martinotti Cells Driven by Facilitating Synapses

The population of pyramidal cells significantly outnumbers the inhibitory interneurons in the neocortex, while at the same time the diversity of interneuron types is much more pronounced. One acknowledged key role of inhibition is to control the rate and patterning of pyramidal cell firing via negative feedback, but most likely the diversity of inhibitory pathways is matched by a corresponding diversity of functional roles. An important distinguishing feature of cortical interneurons is the variability of the short-term plasticity properties of synapses received from pyramidal cells. The Martinotti cell type has recently come under scrutiny due to the distinctly facilitating nature of the synapses they receive from pyramidal cells. This distinguishes these neurons from basket cells and other inhibitory interneurons typically targeted by depressing synapses. A key aspect of the work reported here has been to pinpoint the role of this variability. We first set out to reproduce quantitatively based on in vitro data the di-synaptic inhibitory microcircuit connecting two pyramidal cells via one or a few Martinotti cells. In a second step, we embedded this microcircuit in a previously developed attractor memory network model of neocortical layers 2/3. This model network demonstrated that basket cells with their characteristic depressing synapses are the first to discharge when the network enters an attractor state and that Martinotti cells respond with a delay, thereby shifting the excitation-inhibition balance and acting to terminate the attractor state. A parameter sensitivity analysis suggested that Martinotti cells might, in fact, play a dominant role in setting the attractor dwell time and thus cortical speed of processing, with cellular adaptation and synaptic depression having a less prominent role than previously thought

Hot coffee: associative memory with bump attractor cell assemblies of spiking neurons

Networks of spiking neurons can have persistently firing stable bump attractors to represent continuous spaces (like temperature). This can be done with a topology with local excitatory synapses and local surround inhibitory synapses. Activating large ranges in the attractor can lead to multiple bumps, that show repeller and attractor dynamics; however, these bumps can be merged by overcoming the repeller dynamics. A simple associative memory can include these bump attractors, allowing the use of continuous variables in these memories, and these associations can be learned by Hebbian rules. These simulations are related to biological networks, showing that this is a step toward a more complete neural cognitive associative memory

Investigating the role of fast-spiking interneurons in neocortical dynamics

PhD ThesisFast-spiking interneurons are the largest interneuronal population in neocortex. It is well documented that this population is crucial in many functions of the neocortex by subserving all aspects of neural computation, like gain control, and by enabling dynamic phenomena, like the generation of high frequency oscillations. Fast-spiking interneurons, which represent mainly the parvalbumin-expressing, soma-targeting basket cells, are also implicated in pathological dynamics, like the propagation of seizures or the impaired coordination of activity in schizophrenia. In the present thesis, I investigate the role of fast-spiking interneurons in such dynamic phenomena by using computational and experimental techniques. First, I introduce a neural mass model of the neocortical microcircuit featuring divisive inhibition, a gain control mechanism, which is thought to be delivered mainly by the soma-targeting interneurons. Its dynamics were analysed at the onset of chaos and during the phenomena of entrainment and long-range synchronization. It is demonstrated that the mechanism of divisive inhibition reduces the sensitivity of the network to parameter changes and enhances the stability and exibility of oscillations. Next, in vitro electrophysiology was used to investigate the propagation of activity in the network of electrically coupled fast-spiking interneurons. Experimental evidence suggests that these interneurons and their gap junctions are involved in the propagation of seizures. Using multi-electrode array recordings and optogenetics, I investigated the possibility of such propagating activity under the conditions of raised extracellular K+ concentration which applies during seizures. Propagated activity was recorded and the involvement of gap junctions was con rmed by pharmacological manipulations. Finally, the interaction between two oscillations was investigated. Two oscillations with di erent frequencies were induced in cortical slices by directly activating the pyramidal cells using optogenetics. Their interaction suggested the possibility of a coincidence detection mechanism at the circuit level. Pharmacological manipulations were used to explore the role of the inhibitory interneurons during this phenomenon. The results, however, showed that the observed phenomenon was not a result of synaptic activity. Nevertheless, the experiments provided some insights about the excitability of the tissue through scattered light while using optogenetics. This investigation provides new insights into the role of fast-spiking interneurons in the neocortex. In particular, it is suggested that the gain control mechanism is important for the physiological oscillatory dynamics of the network and that the gap junctions between these interneurons can potentially contribute to the inhibitory restraint during a seizure.Wellcome Trust

Visual Cortex

The neurosciences have experienced tremendous and wonderful progress in many areas, and the spectrum encompassing the neurosciences is expansive. Suffice it to mention a few classical fields: electrophysiology, genetics, physics, computer sciences, and more recently, social and marketing neurosciences. Of course, this large growth resulted in the production of many books. Perhaps the visual system and the visual cortex were in the vanguard because most animals do not produce their own light and offer thus the invaluable advantage of allowing investigators to conduct experiments in full control of the stimulus. In addition, the fascinating evolution of scientific techniques, the immense productivity of recent research, and the ensuing literature make it virtually impossible to publish in a single volume all worthwhile work accomplished throughout the scientific world. The days when a single individual, as Diderot, could undertake the production of an encyclopedia are gone forever. Indeed most approaches to studying the nervous system are valid and neuroscientists produce an almost astronomical number of interesting data accompanied by extremely worthy hypotheses which in turn generate new ventures in search of brain functions. Yet, it is fully justified to make an encore and to publish a book dedicated to visual cortex and beyond. Many reasons validate a book assembling chapters written by active researchers. Each has the opportunity to bind together data and explore original ideas whose fate will not fall into the hands of uncompromising reviewers of traditional journals. This book focuses on the cerebral cortex with a large emphasis on vision. Yet it offers the reader diverse approaches employed to investigate the brain, for instance, computer simulation, cellular responses, or rivalry between various targets and goal directed actions. This volume thus covers a large spectrum of research even though it is impossible to include all topics in the extremely diverse field of neurosciences

Micro-, Meso- and Macro-Dynamics of the Brain

Neurosciences, Neurology, Psychiatr

Über die Selbstorganisation einer hierarchischen Gedächtnisstruktur für kompositionelle Objektrepräsentation im visuellen Kortex

At present, there is a huge lag between the artificial and the biological information processing systems in terms of their capability to learn. This lag could be certainly reduced by gaining more insight into the higher functions of the brain like learning and memory. For instance, primate visual cortex is thought to provide the long-term memory for the visual objects acquired by experience. The visual cortex handles effortlessly arbitrary complex objects by decomposing them rapidly into constituent components of much lower complexity along hierarchically organized visual pathways. How this processing architecture self-organizes into a memory domain that employs such compositional object representation by learning from experience remains to a large extent a riddle. The study presented here approaches this question by proposing a functional model of a self-organizing hierarchical memory network. The model is based on hypothetical neuronal mechanisms involved in cortical processing and adaptation. The network architecture comprises two consecutive layers of distributed, recurrently interconnected modules. Each module is identified with a localized cortical cluster of fine-scale excitatory subnetworks. A single module performs competitive unsupervised learning on the incoming afferent signals to form a suitable representation of the locally accessible input space. The network employs an operating scheme where ongoing processing is made of discrete successive fragments termed decision cycles, presumably identifiable with the fast gamma rhythms observed in the cortex. The cycles are synchronized across the distributed modules that produce highly sparse activity within each cycle by instantiating a local winner-take-all-like operation. Equipped with adaptive mechanisms of bidirectional synaptic plasticity and homeostatic activity regulation, the network is exposed to natural face images of different persons. The images are presented incrementally one per cycle to the lower network layer as a set of Gabor filter responses extracted from local facial landmarks. The images are presented without any person identity labels. In the course of unsupervised learning, the network creates simultaneously vocabularies of reusable local face appearance elements, captures relations between the elements by linking associatively those parts that encode the same face identity, develops the higher-order identity symbols for the memorized compositions and projects this information back onto the vocabularies in generative manner. This learning corresponds to the simultaneous formation of bottom-up, lateral and top-down synaptic connectivity within and between the network layers. In the mature connectivity state, the network holds thus full compositional description of the experienced faces in form of sparse memory traces that reside in the feed-forward and recurrent connectivity. Due to the generative nature of the established representation, the network is able to recreate the full compositional description of a memorized face in terms of all its constituent parts given only its higher-order identity symbol or a subset of its parts. In the test phase, the network successfully proves its ability to recognize identity and gender of the persons from alternative face views not shown before. An intriguing feature of the emerging memory network is its ability to self-generate activity spontaneously in absence of the external stimuli. In this sleep-like off-line mode, the network shows a self-sustaining replay of the memory content formed during the previous learning. Remarkably, the recognition performance is tremendously boosted after this off-line memory reprocessing. The performance boost is articulated stronger on those face views that deviate more from the original view shown during the learning. This indicates that the off-line memory reprocessing during the sleep-like state specifically improves the generalization capability of the memory network. The positive effect turns out to be surprisingly independent of synapse-specific plasticity, relying completely on the synapse-unspecific, homeostatic activity regulation across the memory network. The developed network demonstrates thus functionality not shown by any previous neuronal modeling approach. It forms and maintains a memory domain for compositional, generative object representation in unsupervised manner through experience with natural visual images, using both on- ("wake") and off-line ("sleep") learning regimes. This functionality offers a promising departure point for further studies, aiming for deeper insight into the learning mechanisms employed by the brain and their consequent implementation in the artificial adaptive systems for solving complex tasks not tractable so far.Gegenwärtig besteht immer noch ein enormer Abstand zwischen der Lernfähigkeit von künstlichen und biologischen Informationsverarbeitungssystemen. Dieser Abstand ließe sich durch eine bessere Einsicht in die höheren Funktionen des Gehirns wie Lernen und Gedächtnis verringern. Im visuellen Kortex etwa werden die Objekte innerhalb kürzester Zeit entlang der hierarchischen Verarbeitungspfade in ihre Bestandteile zerlegt und so durch eine Komposition von Elementen niedrigerer Komplexität dargestellt. Bereits bekannte Objekte werden so aus dem Langzeitgedächtnis abgerufen und wiedererkannt. Wie eine derartige kompositionell-hierarchische Gedächtnisstruktur durch die visuelle Erfahrung zustande kommen kann, ist noch weitgehend ungeklärt. Um dieser Frage nachzugehen, wird hier ein funktionelles Modell eines lernfähigen rekurrenten neuronalen Netzwerkes vorgestellt. Im Netzwerk werden neuronale Mechanismen implementiert, die der kortikalen Verarbeitung und Plastizität zugrunde liegen. Die hierarchische Architektur des Netzwerkes besteht aus zwei nacheinander geschalteten Schichten, die jede eine Anzahl von verteilten, rekurrent vernetzten Modulen beherbergen. Ein Modul umfasst dabei mehrere funktionell separate Subnetzwerke. Jedes solches Modul ist imstande, aus den eintreffenden Signalen eine geeignete Repräsentation für den lokalen Eingaberaum unüberwacht zu lernen. Die fortlaufende Verarbeitung im Netzwerk setzt sich zusammen aus diskreten Fragmenten, genannt Entscheidungszyklen, die man mit den schnellen kortikalen Rhythmen im gamma-Frequenzbereich in Verbindung setzen kann. Die Zyklen sind synchronisiert zwischen den verteilten Modulen. Innerhalb eines Zyklus wird eine lokal umgrenzte winner-take-all-ähnliche Operation in Modulen durchgeführt. Die Kompetitionsstärke wächst im Laufe des Zyklus an. Diese Operation aktiviert in Abhängigkeit von den Eingabesignalen eine sehr kleine Anzahl von Einheiten und verstärkt sie auf Kosten der anderen, um den dargebotenen Reiz in der Netzwerkaktivität abzubilden. Ausgestattet mit adaptiven Mechanismen der bidirektionalen synaptischen Plastizität und der homöostatischen Aktivitätsregulierung, erhält das Netzwerk natürliche Gesichtsbilder von verschiedenen Personen dargeboten. Die Bilder werden der unteren Netzwerkschicht, je ein Bild pro Zyklus, als Ansammlung von Gaborfilterantworten aus lokalen Gesichtslandmarken zugeführt, ohne Information über die Personenidentität zur Verfügung zu stellen. Im Laufe der unüberwachten Lernprozedur formt das Netzwerk die Verbindungsstruktur derart, dass die Gesichter aller dargebotenen Personen im Netzwerk in Form von dünn besiedelten Gedächtnisspuren abgelegt werden. Hierzu werden gleichzeitig vorwärtsgerichtete (bottom-up) und rekurrente (lateral, top-down) synaptische Verbindungen innerhalb und zwischen den Schichten gelernt. Im reifen Verbindungszustand werden infolge dieses Lernens die einzelnen Gesichter als Komposition ihrer Bestandteile auf generative Art gespeichert. Dank der generativen Art der gelernten Struktur reichen schon allein das höhere Identitätssymbol oder eine kleine Teilmenge von zugehörigen Gesichtselementen, um alle Bestandteile der gespeicherten Gesichter aus dem Gedächtnis abzurufen. In der Testphase kann das Netzwerk erfolgreich sowohl die Identität als auch das Geschlecht von Personen aus vorher nicht gezeigten Gesichtsansichten erkennen. Eine bemerkenswerte Eigenschaft der entstandenen Gedächtnisarchitektur ist ihre Fähigkeit, ohne Darbietung von externen Stimuli spontan Aktivitätsmuster zu generieren und die im Gedächtnis abgelegten Inhalte in diesem schlafähnlichen "off-line" Regime wiederzugeben. Interessanterweise ergibt sich aus der Schlafphase ein direkter Vorteil für die Gedächtnisfunktion. Dieser Vorteil macht sich durch eine drastisch verbesserte Erkennungsrate nach der Schlafphase bemerkbar, wenn das Netwerk mit den zuvor nicht dargebotenen Ansichten von den bereits bekannten Personen konfrontiert wird. Die Leistungsverbesserung nach der Schlafphase ist umso deutlicher, je stärker die Alternativansichten vom Original abweichen. Dieser positive Effekt ist zudem komplett unabhängig von der synapsenspezifischen Plastizität und kann allein durch die synapsenunspezifische, homöostatische Regulation der Aktivität im Netzwerk erklärt werden. Das entwickelte Netzwerk demonstriert so eine im Bereich der neuronalen Modellierung bisher nicht gezeigte Funktionalität. Es kann unüberwacht eine Gedächtnisdomäne für kompositionelle, generative Objektrepräsentation durch die Erfahrung mit natürlichen Bildern sowohl im reizgetriebenen, wachähnlichen Zustand als auch im reizabgekoppelten, schlafähnlichen Zustand formen und verwalten. Diese Funktionalität bietet einen vielversprechenden Ausgangspunkt für weitere Studien, die die neuronalen Lernmechanismen des Gehirns ins Visier nehmen und letztendlich deren konsequente Umsetzung in technischen, adaptiven Systemen anstreben