Eligibility Traces and Plasticity on Behavioral Time Scales: Experimental Support of neoHebbian Three-Factor Learning Rules

Most elementary behaviors such as moving the arm to grasp an object or walking into the next room to explore a museum evolve on the time scale of seconds; in contrast, neuronal action potentials occur on the time scale of a few milliseconds. Learning rules of the brain must therefore bridge the gap between these two different time scales. Modern theories of synaptic plasticity have postulated that the co-activation of pre- and postsynaptic neurons sets a flag at the synapse, called an eligibility trace, that leads to a weight change only if an additional factor is present while the flag is set. This third factor, signaling reward, punishment, surprise, or novelty, could be implemented by the phasic activity of neuromodulators or specific neuronal inputs signaling special events. While the theoretical framework has been developed over the last decades, experimental evidence in support of eligibility traces on the time scale of seconds has been collected only during the last few years. Here we review, in the context of three-factor rules of synaptic plasticity, four key experiments that support the role of synaptic eligibility traces in combination with a third factor as a biological implementation of neoHebbian three-factor learning rules

Emergence of Physiological Oscillation Frequencies in a Computer Model of Neocortex

Coordination of neocortical oscillations has been hypothesized to underlie the “binding” essential to cognitive function. However, the mechanisms that generate neocortical oscillations in physiological frequency bands remain unknown. We hypothesized that interlaminar relations in neocortex would provide multiple intermediate loops that would play particular roles in generating oscillations, adding different dynamics to the network. We simulated networks from sensory neocortex using nine columns of event-driven rule-based neurons wired according to anatomical data and driven with random white-noise synaptic inputs. We tuned the network to achieve realistic cell firing rates and to avoid population spikes. A physiological frequency spectrum appeared as an emergent property, displaying dominant frequencies that were not present in the inputs or in the intrinsic or activated frequencies of any of the cell groups. We monitored spectral changes while using minimal dynamical perturbation as a methodology through gradual introduction of hubs into individual layers. We found that hubs in layer 2/3 excitatory cells had the greatest influence on overall network activity, suggesting that this subpopulation was a primary generator of theta/beta strength in the network. Similarly, layer 2/3 interneurons appeared largely responsible for gamma activation through preferential attenuation of the rest of the spectrum. The network showed evidence of frequency homeostasis: increased activation of supragranular layers increased firing rates in the network without altering the spectral profile, and alteration in synaptic delays did not significantly shift spectral peaks. Direct comparison of the power spectra with experimentally recorded local field potentials from prefrontal cortex of awake rat showed substantial similarities, including comparable patterns of cross-frequency coupling

Clustering behaviour in networks with time delayed all-to-all coupling

Networks of coupled oscillators arise in a variety of areas. Clustering is a type of oscillatory network behavior where elements of a network segregate into groups. Elements within a group oscillate synchronously, while elements in different groups oscillate with a fixed phase difference. In this thesis, we study networks of N identical oscillators with time delayed, global circulant coupling with two approaches. We first use the theory of weakly coupled oscillators to reduce the system of delay differential equations to a phase model where the time delay enters as a phase shift. We use the phase model to determine model independent existence and stability results for symmetric cluster solutions. We show that the presence of the time delay can lead to the coexistence of multiple stable clustering solutions. We then perform stability and bifurcation analysis to the original system of delay differential equations with symmetry. We first study the existence of Hopf bifurcations induced by coupling time delay, and then use symmetric Hopf bifurcation theory to determine how these bifurcations lead to different patterns of symmetric cluster oscillations. We apply our results to two specfi c examples: a network of FitzHugh-Nagumo neurons with diffusive coupling and a network of Morris-Lecar neurons with synaptic coupling. In the case studies, we show how time delays in the coupling between neurons can give rise to switching between different stable cluster solutions, coexistence of multiple stable cluster solutions and solutions with multiple frequencies

Synchrony and bifurcations in coupled dynamical systems and effects of time delay

Dynamik auf Netzwerken ist ein mathematisches Feld, das in den letzten Jahrzehnten schnell gewachsen ist und Anwendungen in zahlreichen Disziplinen wie z.B. Physik, Biologie und Soziologie findet. Die Funktion vieler Netzwerke hängt von der Fähigkeit ab, die Elemente des Netzwerkes zu synchronisieren. Mit anderen Worten, die Existenz und die transversale Stabilität der synchronen Mannigfaltigkeit sind zentrale Eigenschaften. Erst seit einigen Jahren wird versucht, den verwickelten Zusammenhang zwischen der Kopplungsstruktur und den Stabilitätseigenschaften synchroner Zustände zu verstehen. Genau das ist das zentrale Thema dieser Arbeit. Zunächst präsentiere ich erste Ergebnisse zur Klassifizierung der Kanten eines gerichteten Netzwerks bezüglich ihrer Bedeutung für die Stabilität des synchronen Zustands. Folgend untersuche ich ein komplexes Verzweigungsszenario in einem gerichteten Ring von Stuart-Landau Oszillatoren und zeige, dass das Szenario persistent ist, wenn dem Netzwerk eine schwach gewichtete Kante hinzugefügt wird. Daraufhin untersuche ich synchrone Zustände in Ringen von Phasenoszillatoren die mit Zeitverzögerung gekoppelt sind. Ich bespreche die Koexistenz synchroner Lösungen und analysiere deren Stabilität und Verzweigungen. Weiter zeige ich, dass eine Zeitverschiebung genutzt werden kann, um Muster im Ring zu speichern und wiederzuerkennen. Diese Zeitverschiebung untersuche ich daraufhin für beliebige Kopplungsstrukturen. Ich zeige, dass invariante Mannigfaltigkeiten des Flusses sowie ihre Stabilität unter der Zeitverschiebung erhalten bleiben. Darüber hinaus bestimme ich die minimale Anzahl von Zeitverzögerungen, die gebraucht werden, um das System äquivalent zu beschreiben. Schließlich untersuche ich das auffällige Phänomen eines nichtstetigen Übergangs zu Synchronizität in Klassen großer Zufallsnetzwerke indem ich einen kürzlich eingeführten Zugang zur Beschreibung großer Zufallsnetzwerke auf den Fall zeitverzögerter Kopplungen verallgemeinere.Since a couple of decades, dynamics on networks is a rapidly growing branch of mathematics with applications in various disciplines such as physics, biology or sociology. The functioning of many networks heavily relies on the ability to synchronize the network’s nodes. More precisely, the existence and the transverse stability of the synchronous manifold are essential properties. It was only in the last few years that people tried to understand the entangled relation between the coupling structure of a network, given by a (di-)graph, and the stability properties of synchronous states. This is the central theme of this dissertation. I first present results towards a classification of the links in a directed, diffusive network according to their impact on the stability of synchronization. Then I investigate a complex bifurcation scenario observed in a directed ring of Stuart-Landau oscillators. I show that under the addition of a single weak link, this scenario is persistent. Subsequently, I investigate synchronous patterns in a directed ring of phase oscillators coupled with time delay. I discuss the coexistence of multiple of synchronous solutions and investigate their stability and bifurcations. I apply these results by showing that a certain time-shift transformation can be used in order to employ the ring as a pattern recognition device. Next, I investigate the same time-shift transformation for arbitrary coupling structures in a very general setting. I show that invariant manifolds of the flow together with their stability properties are conserved under the time-shift transformation. Furthermore, I determine the minimal number of delays needed to equivalently describe the system’s dynamics. Finally, I investigate a peculiar phenomenon of non-continuous transition to synchrony observed in certain classes of large random networks, generalizing a recently introduced approach for the description of large random networks to the case of delayed couplings