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The Intracranial Recordings Foundation was established in 2011 to support the organization of the Human Single Neuron conferences, the research, and the community of researchers in the field.

In the last decade we have had tremendous growth in interest and discoveries, we published the first research compilation in the field and held five big conferences.


For information and questions, email: or any of the Foundation Board of Directors: Moran CerfUeli RutishauserGabriel Kreiman,

The Intracranial Recordings Foundation is a 501(c)(3) organization.

In order to be informed on future meetings, data sharing opportunities, and important updates - please sign up below:

Welcome to the foundation.




November 10-11, 2011

New York University, NY, NY

Day 1:

Welcome address. Moran Cerf (NYU), Gabriel Kreiman (Harvard), Ueli Rutishauser (Cedars-Sinai)

Keynote address:

George Ojemann (Washington University). "50+ years of human single-neuron recording, a personal perspective."  

Itzhak Fried (Tel-Aviv University): "Neurons as will and representation"    

Session I: Perception and Memory I

Gabriel Kreiman (Harvard): "Selectivity, tolerance, and speed in visual object recognition"

Rodrigo Quiroga (Leicester University): "Concept cells"

Ueli Rutishauser (Cedars-Sinai): "Mechanisms of declarative memory formation in the human medial temporal lobe"

Moran Cerf (NYU): "Control of single neurons using thought"

Session II: Perception and Memory IO

Florian Mormann (Bonn): "Single-unit correlates of visual object recognition, working memory and attention in the human medial temporal lobe"

Hagar Gelbard-Sagiv (UCLA): "Neuronal correlates of memory formation and recollection in the human medial temporal lobe"  

Eli Nelken (Hebrew University): "Single neuron studies of auditory signals

Michael Kahana (University of Penssylvania): "Single unit studies of human spatial navigation, episodic memory, and decision making"

Session III: Clinical, technical, and surgical aspects

Adam Mamelak (Cedars-Sinai): "Setting up a human single-unit lab: ethical, clinical, surgical and practical aspects"

Rodrigo Quiroga (Leicester University): "Spike detection and sorting"

Rick Staba (UCLA): "Electrodes for subchronic human direct brain recording"

Moran Cerf (NYU): "Decoding real-time signals"

Raffi Malach (Weizmann Institute): "What can BOLD-fMRI tell us about single-unit responses in the human brain"


Session IV: Sponsors

Casey Stengel (Neuralynx)  

Andy Gotshalk (Blackrock)

Day 2:

Session IV: Motor and free will

Roy Mukamel (Tel-Aviv University): "Mirror neurons"

Gabriel Kreiman (Harvard): "Single neurons predict free will"

Wilson Truccolo (Brown): "Collective Dynamics at the level of Single Neurons in Human Neocortex"

George Ojemann (Washington University): "Language, Learning, Local Field Potentials prediction of spike timing"

Session V: Emotions and sleep

Ralph Adolphs (Caltech): "Amygdala responses to faces: Extensions to autism"

Moran Cerf (NYU): "Single neuron correlates of emotions regulation in humans"

Rick Staba (UCLA): "Human limbic single neuron firing patterns during sleep"

Session VI: Epilepsy

Sydney Cash (Harvard): Neuronalk Dynamics During Focal Seizures

Bradley Greger (University of Utah): "In Vivo extracellular recordings and in vitro intracellular recording from human epileptic neocortex: insights into the relations between inter-ictal spikes, action potentials, and high-frequency oscillations"

Session VII: The next 10 years

Keynote address: 

Christof Koch (Allen Institute): "Project Mindscope. Building cortical observatories."

Panel discussion: Itzhak Fried, Moran Cerf, Gabriel Kreiman, Ueli Rutishauser





November 13-14, 2014

Johns Hopkins, Baltimore, MD

Day 1:

Session I: Decision making

Sameer Sheth (Columbia): "The Physiology of Human Cognitive Control: Investigations in the Dorsal Anterior Cingulate Cortex"

Kareem Zaghoul (NIH): "Single unit activity in the human subthalamic nucleus during decision making"

Ziv Williams (MGH/Harvard): "Prefrontal recordings in relation to cognitive control"


Session II: Perception, recognition, faces

Katalin Gothard (U. Arizona): "Naturalistic social stimuli elicit eye-selective neural responses in the monkey amygdala"

Moran Cerf (Northwestern and NYU): "Single neuron correlates of emotion regulation in natural sensory content"


Flash poster presentation

Shaun Patel (MGH): "Rapid intermittent Deep Brain Stimulation biases behavior in financial decision-making task"

Juri Minxha (Caltech): "Fixation-aligned single cell responses in human and non-human primate amygdala"

Taufik Valiante (University of Toronto): "Laminar specific specialization in regular spiking neurons in superficial and deep cortical laminae of human cortex maintained in vitro."

Shaun Aibel Weiss (UCLA): "Searching for synchrony: a microelectrode study of neuronal spike firing during human seizures"




Session III: Clinical neuroscience (epilepsy, seizures, DBS)

Catherina Schevon (Columbia): "Unraveling the electrophysiology of human seizures"

Nathan Crone (Hopkins): "Neural population dynamics in human cortical function"

Stan Anderson (Hopkins): "Clinical and therapeutic implications of cortical neural network modeling"

Helen Mayberg (Emory): "Therapeutic Modulation of Cingulate-Cortical Oscillations in Major Depression: Groundwork for next-generation closed-loop technologies"


Conference dinner

Day 2:

Session IV: Motor and free will

Roy Mukamel (Tel-Aviv University): "Mirror neurons"

Gabriel Kreiman (Harvard): "Single neurons predict free will"

Day II. Friday, November 14, 2014

09:00 - 09:15: Welcome

Session IV: Memory

Elizabeth Buffalo (Seattle): "Neural activity related to visual exploration and navigation in primates"

Ueli Rutishauser (Cedars-Sinai/Caltech): "Neural population dynamics during declarative memory/retrieval"


Keynote address:

Larry Squire (UCSD): "Conscious and unconscious memory systems of the mammalian brain"

Session V: Memory II

John Wixted (UCSD): "Sparse and Distributed Coding of Episodic Memory in Neurons of the Human Hippocampus"

Florian Mormann (Bonn): "Single unit activity during perception and memory in the human medial temporal lobe"

Andreas Schulze-Bonhage (Freiburg) and Michael Kahana (U. Penn): "The spatial  context of retrieved memories is reflected by neural activity in the human hippocampal formation"

Session V: Memory II (Cont.)

Josh Jacobs (Drexel): "Entorhinal neuronal representations in human spatial navigation"

Session VI: Neural prosthesis

Richard Andersen (Caltech): "Decoding motor imagery from the Posterior Parietal Cortex of a tetraplegic human"

Carlos Vargas-Irwin (Brown): "Exploring the neural representation of attempted, imagined, and observed actions in human motor cortex using spike train similarity analysis"


Technology presentation

Designated by primary sponsor; Neuralynx


Session VII: Speical topics

Josef Parvizi (Stanford): "Intracranial EEG and Electrical Brain Stimulation"

Ed Lein and Jonathan Ting (Allen institute): "Multimodal interrogation of the cellular and local circuit architecture of the human neocortex"

Dion Khodagholy (NYU): "NeuroGrid. Recording action potentials from the surface of the brain"





November 10-11, 2016

Caltech, Pasadena, CA

Day 1:

Welcome address. Moran Cerf (NYU), Gabriel Kreiman (Harvard), Ueli Rutishauser (Cedars-Sinai)

Session I: Learning and memory

Rodrigo Quian-Quiroga (Leicester): "Concept cells and memory"

Josh Jacobs (Columbia): "Neuronal representations in spatial navigation and memory"

Gabriel Kreiman (Harvard): "Computational, behavioral and physiological mechanisms of episodic memory formation"

Nanthia Suthana (UCLA): "Neuronal characterization and modulation of human episodic memory"

Session II: Vision

Pieter Roelfsema (Amsterdam): "The activity of neurons in early visual cortex of humans and monkeys during perceptual organization"

James Bisley (UCLA): "Representations of stimulus similarity in parietal cortex"

Session III: Clinical neuroscience

William Anderson (Johns Hopkins): "Applied Computational Modeling of the Neocortex"

Nicholas Schiff (Cornell): "Cognitive motor dissociation: underlying mechanisms and challenges for establishing communication interfaces"

William Hutchison (U Toronto): "Beta activity in motor thalamus and STN during an inverted center out task"

Keynote address

Christof Koch (Allen Institute): "Characterizing neocortical mouse and human cell types"

Session IV: Funding initiatives

James Gnadt (NIH) and Kurt Thoroughman (NSF): "New and innovative federal funding opportunities"

Session V: Flash presentations


Session VI: Novel ECoG approaches

Florian Mormann (Bonn): "Single-neuron correlates of memory encoding and consolidation in the human MTL"

Ralph Adolhps (Caltech): "Concurrent electrical stimulation and fMRI to map out effective connectivity in the human brain"

Conference dinner

Day 2:

Session VII: Brain-Machine Interfaces

Richard Andersen (Caltech): "Brain-machine interfaces using the posterior parietal cortex"

Jaimie Henderson (Stanford): "Brain-Computer Interfaces for Communication and Generalized Computer Use"

Elizabeth Tyler-Kabala (U. Pittsburgh): "Chronic human single unit recordings: Balance of ethical considerations and potential benefits for future users of BCI technology"

Bolu Ajiboye (Case Western): "Re-thinking Paralysis: Brain-machine interfaces for movement restoration in persons with Chronic High Tetraplegia"

Session VIII: Executive function / Decision making

Michelle Basso (UCLA): "Impaired Decision-Making in Parkinson’s Disease"

Ueli Rutishauser (Cedars-Sinai): "Mechanisms of error monitoring in human medial frontal cortex"

Session IX: Learning and memory II

Itzhak Fried (Tel-Aviv University): "Present and future landscape of human single neuron recordings"

Karem Zaghoul (NIH): "Human cortical neurons reinstate spiking activity during episodic memory encoding and retrieval"

Kari Hoffman (York): "Sharp-wave ripples during memory-guided visual search"

Keynote address:

Wolfram Schultz (Cambridge): "Well-controlled risky gambles suitable for neuronal recording studies"

Session X: Ex-vivo human recordings

Taufik Valiante (University of Toronto): "Surgery"

Huib Mansveider (University of Amsterdam): "Unique properties of neuronal microcircuits of the human neocortex"

Gold sponsor talk

Casey Stengel, (Neuralynx, Inc.): "Research in a Clinical Setting: DC Recording and Stimulation"


Session XI: Epilepsy and sleep

Yuval Nir (Tel Aviv University): "Sleep, sleepiness, and anesthesia: a view from inside"





November 1-2, 2018

Caltech, Pasadena, CA

Day 1:

Session I: Large-scale brain interaction across primates

Bijan Pesaran (NYU): Studying the primate brain on a large-scale - imaging, electrophysiology and fmri.

Christopher Petkov (Newcastle University): Mental structures and the primate brain: From human ECoG to monkey neurons

Charles Schroeder/Idan Tal (Columbia): Temporal structure of transient events as a measure of cortical interactions.

Keynote address:

Richard Andersen (Caltech): "Developing a bidirectional human-machine interface that can read out intentions and write in sensations"

Session II: Decision making

Joshua Gold (University of Pennsylvania): "Basal ganglia recordings in human and non-human primates"

Ziv Williams (MGH): "Single neuronal correlates of social decisions in the human prefrontal cortex"

Sameer Sheth (Baylor): "Prominent temporal coding of decision variables in human prefrontal cortex"

Session III: Brain initiatives

Itzhak Fried (UCLA): "From human single neuron recordings to cognitive prediction and modulation"

Jim Gnadt (NIH/NINDS): "Research Opportunities in Humans with the NIH BRAIN Initiative"


Flash presentations

Alexander Unruh (U Bonn): "Decision confidence is represented at the single-unit level in the human medial temporal lobe"

Juri Minxha (Caltech): "Phase-locking of neurons in human medial frontal cortex to hippocampal theta is engaged by declarative memory-based decisions"

Marije Ter Wal (University of Birmingham): "Exploring the "what", "when" and "where" of memory reinstatement in human intracranial EEG recordings

Jan Kaminski (Cedars-Sinai): Evidence for domain-specific working memory buffers from human single-neuron recordings

Session IV: Ex-vivo study of human single cells

Jonathan Ting (Allen Institute): "Conserved and divergent features of human versus mouse neocortical cell types"

Mark Harnett (MIT): "Enhanced dendritic compartmentalization in human cortical neurons"

Viren Jain (Google AI): "High-throughput synapse-resolution connectomics in fly, bird, and human brains"

Taufik Valiante (University of Toronto): "Divergent electrophysiological properties of layer 2/3 and layer 5 pyramidal neurons in human temporal cortex"

Conference dinner

Day 2:

Session V: Language

Edward Chang (UCSF): "Encoding movement in speech motor cortex"

Mark Richardson (University of Pittsburgh): "Subthalamic nucleus unit activity during speech production"

Session VI: Epilepsy

Sydney Cash (MGH): "The role of inhibitory and excitatory neurons in sculpting seizure initiation and propagation"

Catherine Schevon (Columbia): "Cell-type specific activity at seizure onset in humans"

Keynote address:

Loren Frank (UCSF): "Neural substrates of prospection and new tools for understanding them"

Session V: New electrodes / technology

Shadi Dayeh (UCSD): "Scalable and Minimally Invasive Technologies for Recording Surface Units and Intracellular Potentials at Depth in Intact Brains"

Michael Roukes (Caltech): "Open Neurotech Alliance – Towards Next-Generation Neurotechnology in Humans"

Casey Stengel (Neuralynx): "Recording and Stimulation in Real-Time."

Session VI: Memory / spatial cognitioin

Robert Knight (UC Berkeley): "Neural Networks and Human Behavior"

Nanthia Suthana (UCLA): "Single neuron and oscillatory correlates of real-world spatial navigation in humans"

Simon Hanslmayr (University of Birmingham): "A synchronized hippocampus and a desynchronized neocortex underlie human episodic memory formation"

Zoltan Nadasdy (UT Austin/HCA): "Spike-phase grids in the human entorhinal cortex"

Dion Khodagholy (Columbia): "Learning-enhanced coupling between ripple oscillations"


Session VII: Faces and attention

Doris Tsao (Caltech): "The macaque face processing system"

Shuo Wang (WVU): "Single-neuron representation of goal-directed signals during visual search"

Thilo Womelsdorf (Vanderbilt): "Cross-level Interactions of Distinct Cell Types to Fronto-Striatal Network Activity During Attention and Learning"





November 13, 2020


Session I: Large-scale brain interaction across primates

Bijan Pesaran (NYU): Studying the primate brain on a large-scale - imaging, electrophysiology and fmri.


Keynote address:

Elizabeth Buffalo (UW). "Time, Space, and Memory in the Primate Hippocampus"

NIH BRAIN Initiative

John Ngai (BRAIN initiative), Karen David (NINDS), Jim Gnadt (NINDS)

Session I: Everything

Ashley Feinsinger (UCLA): "What's the value of participant engagement in basic human neuroscience?"

Kareem Zaghloul (NIH/NINDS): "Replay of cortical spiking sequences during episodic memory retrieval"

Christof Koch (Allen Institute): "Morphological, electrical and transcriptional characterization of human neocortical supragranular pyramidal neurons"

Yuval Nir (Tel-Aviv University): "Auditory processing in sleep and anesthesia: insights from single-neuron human studies"

Krishna Shenoy (Stanford University): "Cortical basis of speech and handwriting in humans for neural interfaces"

Session II: New technology

Nanthia Suthana (UCLA). "Recording single neurons in the human brain during naturalistic behaviors"

Mikhail Shapiro (Caltech): "Ultrasonic Imaging and Control of Neural Activity"


Session III: Memory and computational methods

Lukas Kunz (Freiburg): "A neural code for egocentric spatial maps in the human medial temporal lobe"

Salman Qasim (Columbia): "Phase precession in the human hippocampus and entorhinal cortex"

Jie Zheng (Harvard): "Event boundaries shape episodic memory"

Gray Umbach (UT Southwestern): "Time cells in the human medial temporal lobe aid episodic memory"

Kanaka Rajan (Mount Sinai): "Inferring brain-wide interactions using data-constrained neural network models"

Session IV: Brain-Macahine Interfaces

Sofia Sakellaridi (Caltech): "Understanding learning mechanisms in the human motor system using brain-machine interface in patients with tetraplegia"

Angelica Herrera (University of Pittsburgh): "Effect of object presence and grasp intention on M1 activity during planning and movement execution"

Session V: Decision making

Tomas Aquino (Caltech): "Correlating human vmPFC and amygdala neurons with value and uncertainty in the explore-exploit dilemma"

Habiba Azab (Baylor): "Algorithmic place cells in human hippocampus"

Clayton Mosher (Cedars-Sinai): "Distinct roles of dorsal and ventral subthalamic neurons in action selection and cancellation"

Session VI: Epilepsy and ex-vivo human slices

Meanhwan Kim (Allen Institute): "Molecular and genetic approaches for assaying human cell type synaptic connectivity"

Elliot Smith (University of Utah): "Human interictal epileptiform discharges are traveling waves reflecting ictal self-organization"




"Single Neuron Studies of the Human Brain"

Receive 30% discount on the book here with code MHSN30.


Open Letter to a Beginning Researcher in the Field of Human Single Neuron Investigations


Itzhak Fried, Ueli Rutishauser, Moran Cerf, and Gabriel Kreiman

Fifty-plus Years of Human Single Neuron Recordings: A Personal Perspective

George Ojemann

Methodological, Ethical, and Clinical Consideration

The Neurosurgical Theater of the Mind

Itzhak Fried

Ethical and Practical Considerations for Human Microelectrode Recording Studies

Adam N. Mamelak

Subchronic In Vivo Human Microelectrode Recording

Richard J. Staba, Tony A. Fields, Eric J. Behnke, and Charles L. Wilson

Data Analysis Techniques for Human Microwire Recordings: Spike Detection and Sorting, Decoding, Relation between Neurons and Local Field Potentials
Ueli Rutishauser, Moran Cerf, and Gabriel Kreiman

Cognitive Neuroscience Findings and Insights

Single Neuron Correlates of Declarative Memory Formation and Retrieval in the Human Medial Temporal Lobe 
Ueli Rutishauser, Erin M. Schuman, and Adam N. Mamelak

Visual Cognitive Adventures of Single Neurons in the Human Medial Temporal Lobe

Florian Mormann, Matias J. Ison, Rodrigo Quian Quiroga, Christof Koch, Itzhak Fried, and Gabriel Kreiman 

Navigating Our Environment: Insights from Single Neuron Recordings in the Human Brain

Nanthia Suthana and Itzhak Fried

Microelectrode Studies of Human Sleep

Yuval Nir, Michel Le Van Quyen, Giulio Tononi, and Richard J. Staba

Studying Thoughts and Deliberations Using Single Neuron Recordings in Humans

Moran Cerf, Hagar Gelbard-Sagiv, and Itzhak Fried

Human Single Neuron Reward Processing in the Basal Ganglia and Anterior Cingulate

Shaun R. Patel, Demetrio Sierra-Mercado, Clarissa Martinez-Rubio, and Emad N. Eskandar

Electrophysiological Responses to Faces in the Human Amygdala

Ralph Adolphs, Hiroto Kawasaki, Oana Tudusciuc, Matthew Howard III, Chris Heller, William Sutherling, Linda Philpott, Ian Ross, Adam N. Mamelak, and Ueli Rutishauser

Human Lateral Temporal Cortical Single Neuron Activity during Language, Recent Memory, and Learning

George Ojemann

Clinical Neuroscience

Microelectrode Recordings in Deep Brain Stimulation Surgery

C. Rory Goodwin, Travis S. Tierney, Frederick A. Lenz, and William S. Anderson

Microstimulation Effects on Thalamic Neurons

Sanjay Patra, William D. Hutchison, Clement Hamani, Mojgan Hodaie, Andres M. Lozano, and Jonathan O. Dostrovsky

Human Single Neuron Activity for Reach and Grasp Motor Prostheses

Arjun K. Bansal

Human Single Neuron Recording as an Approach to Understand the Neurophysiology of Seizure Generation

Andreas Schulze-Bonhage and Rüdiger Köhling


The Next Ten Years and Beyond

Ueli Rutishauser, Itzhak Fried, Moran Cerf, and Gabriel Kreiman



Features research from members of the community


Time cells in the human hippocampus and entorhinal cortex support episodic memory
Gray Umbach, Pranish Kantak, Joshua Jacobs, Michael Kahana, Brad E Pfeiffer, Michael Sperling, Bradley Lega, 
PNAS 117 (45), 28463-28474.


The organization of temporal information is critical for the encoding and retrieval of episodic memories. In the rodent hippocampus and entorhinal cortex, evidence accumulated over the last decade suggests that populations of “time cells” in the hippocampus encode temporal information. We identify time cells in humans using intracranial microelectrode recordings obtained from 27 human epilepsy patients who performed an episodic memory task. We show that time cell activity predicts the temporal organization of retrieved memory items. We also uncover evidence of ramping cell activity in humans, which represents a complementary type of temporal information. These findings establish a cellular mechanism for the representation of temporal information in the human brain needed to form episodic memories.


Single-neuronal predictions of others’ beliefs in humans
Mohsen Jamali, Benjamin L. Grannan, Evelina Fedorenko, Rebecca Saxe, Raymundo Báez-Mendoza & Ziv M. Williams, NATURE 2021, 1-5.


Human social behaviour crucially depends on our ability to reason about others. This capacity for theory of mind has a vital role in social cognition because it enables us not only to form a detailed understanding of the hidden thoughts and beliefs of other individuals but also to understand that they may differ from our own1–3. Although a number of areas in the human brain have been linked to social reasoning4,5 and its disruption across a variety of psychosocial disorders6–8, the basic cellular mechanisms that underlie human theory of mind remain undefined. Here, using recordings from single cells in the human dorsomedial prefrontal cortex, we identify neurons that reliably encode information about others’ beliefs across richly varying scenarios and that distinguish self- from other-belief-related representations. By further following their encoding dynamics, we show how these cells represent the contents of the others’ beliefs and accurately predict whether they are true or false. We also show how they track inferred beliefs from another’s specific perspective and how their activities relate to behavioural performance. Together, these findings reveal a detailed cellular process in the human dorsomedial prefrontal cortex for representing another’s beliefs and identify candidate neurons that could support theory of mind.





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