Representative Publications

For a complete list of publications see https://orcid.org/0000-0002-9138-4833

Membrane & neuron excitability

A biophysical perspective on the resilience of neuronal excitability across timescales \ Marom, Marder \ Nature Rev. Neurosci. (2023):  Neuronal membrane excitability must be resilient to perturbations that can take place over timescales from milliseconds to months (or even years in long-lived animals). This paper offers a self-regulating ‘automatic’ mechanism that contributes to neuronal resilience by virtue of the kinetic properties of ion channels themselves.

Sodium channel slow inactivation normalizes firing in axons with uneven conductance distributions \ Zang, Marder, Marom \ Current Biology (2023): Here we model spike propagation in axons with uneven distributions of Na+ and K+ channels. Including slow inactivation of the Na+ channel enhances the reliability of spike propagation and can compensate for uneven channel distributions.

Dynamic clamp constructed phase diagram for the Hodgkin and Huxley model of excitability \ Ori, Hazan, Marder, Marom \ PNAS (2020): Here we study the relationship between sodium and potas- sium channel parameters and membrane excitability. We used the dynamic clamp, and established a real-time closed-loop interaction between a genetically controlled population of excitability-relevant ion channels and a low-dimensional mathematical description of excitability. The results provide insights into how robustness of excitability benefits from the variability of history-dependent time scales that ion channels display.

Emergence and maintenance of excitability: kinetics over structure \ Marom \ Current Opinion in Neurobiology (2016): Alongside studies that extend traditional focus on control-based regulation of structural parameters (channel densities), there is a budding interest in self-organization of kinetic parameters. In this picture, ionic channels are continually forced by activity in-and-out of a pool of states not available for the mechanism of excitability. The process, acting on expressed structure, provides a bed for generation of a spectrum of excitability modes.

Cellular function given parametric variation in the Hodgkin and Huxley model of excitability \ Ori, Marder, Marom \ PNAS (2018):  A theoretical study, showing that although the full Hodgkin–Huxley model is very sensitive to fluctuations that independently occur in its many parameters, the outcome is in fact determined by a simple combinations of these parameters along two physiological dimensions - structural and kinetic (denoted S and K, respectively). The impacts of parametric fluctuations on the dynamics of the system — seemingly complex in the high-dimensional representation of the Hodgkin–Huxley model — are tractable when examined within the S–K plane.

Entrainment of the intrinsic dynamics of single isolated neurons by natural-like input \ Gal, Marom \ J Neurosci. (2013): Neuronal dynamics is intrinsically unstable, producing activity fluctuations that are essentially scale free. Here we experimentally study single cortical neurons, and show that while these scale-free fluctuations are independent of temporal input statistics, they can be entrained by input variation. Response entrainment was found to be maximal when the input itself possesses natural-like, scale-free statistics.

Neural timescales or lack thereof \ Marom \ Prog. Neurobiol. (2010): This article aims at making readers, experimentalists and theorists, more aware of the abstractions made by an observer when measuring and reporting behavioral and neural timescales. These abstractions stow away the fact that, above lower boundaries that reflect fairly well understood physical constraints, observed and reported timescales are often not intrinsic to the biological system; rather, in most cases they reflect conditions that are imposed by the observer through the measuring procedure.

Self-organized criticality in single-neuron excitability \ Gal, Marom \ Phys. Rev. E (2013): Here, we suggest that neuronal response fluctuations reflect a process that positions the neuron near a transition point that separates excitable and unexcitable phases. This view is supported by the dynamical properties of the system as observed in experiments on isolated cultured cortical neurons, and by a theoretical mapping between the constructs of self-organized criticality and membrane excitability.

Interaction between duration of activity and time course of recovery from slow inactivation in mammalian brain Na+ channels \ Toib, Lyakhov, Marom \ J. Neurosci. (1998): The relationships between activity and availability for activation of voltage gated Na channels were examined using the Xenopus expression system. The main point of this work is that the time constant of recovery from the unavailable (inactivated) pool is related to the duration of previous activation by a power law. These relationships extend from tens of milliseconds to several minutes and are intrinsic to the channel protein.

Adaptive transition rates in excitable membranes \ Marom \ Frontiers Comp. Neurosci. (2009): This study shows that adaptation in excitable membranes is reducible to a simple Logistic-like equation in which the essential non-linearity is replaced by a feedback loop between the history of activation and an adaptive transition rate that is sensitive to a single dimension of the space of inactive states. This physiologically measurable dimension contributes to the stability of the system and serves as a powerful, intrinsic modulator of input–output relations that depends on the patterns of prior activity.

Modeling State-Dependent Inactivation of Membrane Currents \ Marom, Abbott \ Biophysical J. (1994): Inactivation of many ion channels occurs through largely voltage-independent traitions to an inactivated state from the open state or from other states in the pathway leaing to opening of the chanel. Because this form of inactvation is state-dependent rather than voltage-dendent, it cannot be described  by the standard Hodgkin-Huxley (HH) formalism. Using two examples ,we extend the standad HH formalism for modeling macroscopic currents to account for state-dependent inacvation.

A 3-D approach to voltage-clamp data \ Marom \ J Theoreticl Biology (1992): This very early paper, which I am most proud of, was never cited, by no-one... It presents a simple transition from 2-D handling of voltage clamp data, to 3-D, thus uncovering some stimulating relationships between the graphic representation of the I–V–t space of electrical activity of the membrane, and the physiological and biophysical functions of the plotted ionic currents.

Neural network dynamics & function

Order-Based Representation in Random Networks of Cortical Neurons \ Shahaf, Eytan, Gal, Kermany, Lyakhov, Zrenner, Marom \ PLOS Computational Biology (2008):  We show that in spontaneously developing large-scale random networks of cortical neurons in vitro the order in which neurons are recruited following each stimulus is a naturally emerging representation primitive that is invariant to significant temporal changes in spike times. With a relatively small number of randomly sampled neurons, the information about stimulus position is fully retrievable from the recruitment order.

Dynamics and Effective Topology Underlying Synchronization in Networks of Cortical Neurons \ Eytan, Marom \ J. Neurosci. (2006): We characterize the process underlying the timescale of synchronization, its relationship to the effective topology of connectivity within an assembly, and its impact on propagation of activity within and between assemblies. We show that the basic mode of assembly activation, “network spike,” is a threshold-governed phenomenon, following the logistics of neuronal recruitment in an effectively scale-free connected network.

Learning in Networks of Cortical Neurons \ Shahaf, Marom \ J. Neurosci. (2001): The results presented here demonstrate selective learning in a network of real cortical neurons, without the involvement of a neural rewarding entity. The connectivity can be modulated by external focal stimulation in an activity-dependent manner. Most importantly, the networks explore the space of possible responses and stabilize at configurations that remove the stimuli.

On the precarious path of reverse neuro-engineering \ Marom, Meir, Braun, Gal, Kermany, Eytan \ Frontiers in Computational Neuroscience (2009): In this perspective we demonstrate that application of reverse engineering to the study of the design principle of a functional neuro-system with a known mechanism, may result in a perfectly valid but wrong induction of the system’s design principle. If in the very simple setup we bring here it is difficult to induce a design principle, what are our chances of exposing biological design principles when more realistic conditions are examined? [robot movie and description]

Selective Adaptation in Networks of Cortical Neurons \ Eytan, Brenner, Marom \ J. Neurosci. (2003): A key property of neural systems is their ability to adapt selectively to stimuli with different features. Using multisite electrical recordings from networks of cortical neurons developing ex vivo, we show that neurons adapt selectively to different stimuli invading the network. We focus on selective adaptation to frequent and rare stimuli; networks were stimulated at two sites with two different stimulus frequencies. When both stimuli were presented within the same period, neurons in the network attenuated their responsiveness to the more frequent input, whereas their responsiveness to the rarely delivered stimuli showed a marked average increase.

Dopamine-Induced Dispersion of Correlations Between Action Potentials in Networks of Cortical Neurons \ Eytan, Minerbi, Ziv, Marom \ J. Neurophysiol. (2004): The neuro- modulatory effects of dopamine at the synaptic and cellular levels are very rich, but it is difficult to extrapolate from these elementary levels what their effect might be at the behaviorally relevant level of neuronal ensembles. Using multi-site recordings from networks of cortical neurons developing ex vivo, we studied the effects of dopa- mine on connectivity within neuronal ensembles. We found that dopamine disperses correlations between individual neuronal activities while preserving the global distribution of correlations at the network level.

On depth psychology and physiology

Science, Psychoanalysis, and the Brain: Space for Dialogue \ Marom \ Cambridge University Press (2013) [originally published in English; translated to Hebrew and Italian]: This monograph is an invitation to a space for dialogue where reflections on neurophysiology are expressed with and guided by depth-psychology in mind; a space where neurophysiology resumes its traditional humbled stance towards matters of the psyche, and where the intellectual autonomy of depth psychology is acknowledged. The author leads the reader through the terrain of methodological errors that have plagued recent reductive approaches, paving the way for a dialogue that is based on an alternative, relational approach. 

Jeffrey Michael Harp / Victorian Surrealism

Dialogue Across Chasm: Are Psychology and Neurophysiology Incompatible? \ Marom \ Neuron (2020): To establish a genuine scientific discourse, we must accept a long due departure from the habit of neatly ar- ranging things in a hierarchy where ‘‘macroscopic’’ psychological mystery awaits explanation in terms of ‘‘microscopic’’ neural objects. Instead, a relational scientific methodology is wanted, accompanied by a dia- logic mode of conversation between the disciplines.

איור מחווה לאמן ג'פרי מישל הארפ / רות גווילי

הפיתוי הנצחי / מרום / אודיסאה 48 (2010): מדי כמה שנים, בענף מדעי זה או אחר נדמה לרגע, שאך כפסע בינינו ובין הבנה מלאה של התחום. יומרה שכזו מאפיינת כעת את מדעי המוח, כשחלק מהחוקרים מאמין שהנה, רק נצליח להסיר מדרכנו מכשול טכני אחד או שניים, כי אז נבין את רוח האדם במונחים ביולוגיים. המאמר מציע לנו להצטנע ולחזור להכיר במגבלותינו.