Foundational materials

Reading and Writing the Morphogenetic Code
Foundational White Paper of the Allen Discovery Center at Tufts University

2024

Neuromorphic responses of nanofluidic memristors in symmetric and asymmetric ionic solutions
Ramirez, P., Portillo, S., Cervera, J., Nasir, S., Ali, M., Ensinger, W., and Mafe, S.
The Journal of Chemical Physics, 160: 044701

Federated inference and belief sharing
Friston, K. J., Parr, T. Heins, C., Constant, A., Friedman, D., Isomura, T., Fields, C., Verbelen, T., Ramstead, M., Clippinger, J., and Frith, C. D.
Neuroscience and Biobehavioral Reviews, 156: 105500

2023

All quantum systems are strange: A commentary on “Path integrals, particular kinds, and strange things” by Friston, Da Costa, Sakthivadivel, Heins, Pavliotis, Ramstead, and Parr
Fields, C., and Glazebrook, J. F.
Physics of Life Reviews, 47: 145-147

Separability, Contextuality, and the Quantum Frame Problem
Fields, C., and Glazebrook, J. F.
International Journal of Theoretical Physics, 62: 159

A Variational Synthesis of Evolutionary and Developmental Dynamics
Friston, K., Friedman, D. A., Constant, A. Knight, V. B., Fields, C., Parr, T., and Campbell, J. O.
Entropy, 25(7): 964

Ordered ground state configurations of the asymmetric Wigner bilayer system—Revisited with unsupervised learning
Hartl, B., Mihalkovič, M., Šamaj, L., Mazars, M., Trizac, E., and Kahl, G.
Journal of Chemical Physics, 159(20): 204112

Does Deep Learning Have Epileptic Seizures? On the Modeling of the Brain
Depannemaecker, D., Pio-Lopez, L., and Gauld, C.
Cognitive Computation, doi: 10.1007/s12559-023-10113-y

Transplantation of fragments from different planaria: A bioelectrical model for head regeneration
Cervera, J., Manzanares, J. A., Levin, M., and Mafe, S.
Journal of Theoretical Biology, 558: 111356

Learning in Transcriptional Network Models: Computational Discovery of Pathway-Level Memory and Effective Interventions
Biswas, S., Clawson, W., and Levin, M.
International Journal of Molecular Sciences, 24(1): 285

2022

Active inference, morphogenesis, and computational psychiatry
Pio-Lopez, L., Kuchling, F., Tung, A., Pezzulo, G., and Levin, M.
Frontiers in Computational Neuroscience, 16: 988977

Competition for finite resources as coordination mechanism for morphogenesis: An evolutionary algorithm study of digital embryogeny
Smiley, P., and Levin, M.
Biosystems, 221: 104762

HCN2 channel-induced rescue of brain, eye, heart and gut teratogenesis caused by nicotine, ethanol and aberrant notch signalling
Pai, V. P., and Levin, M.
Wound Repair and Regeneration, 30(6): 681-706

Training spiking neuronal networks to perform motor control using reinforcement and evolutionary learning
Haşegan, D., Deible, M., Earl, C., D’Onofrio, D., Hazan, H., Anwar, H., and Neymotin, S. A.
Frontiers in Computational Neuroscience, 16: 1017284

Quantum Neural Networks and Topological Quantum Field Theories
Marciano, A., Chen, D., Fabrocini, F., Fields, C., Greco, E., Gresnigt, N., Jinklub, K., Lulli, M., Terzidis, K., and Zappala, E.
Neural Networks, 153: 164-178

Neurons as hierarchies of quantum reference frames
Fields, C., Glazebrook, J. F., and Levin, M.
Biosystems, 219: 104714

A free energy principle for generic quantum systems
Fields, C., Friston, K., Glazebrook, J. F., and Levin, M.
Progress in Biophysics and Molecular Biology, 173: 36-59

Enhancers of Host Immune Tolerance to Bacterial Infection Discovered Using Linked Computational and Experimental Approaches
Sperry, M. M., Novak, R., Keshari, V., Dinis, A. L. M., Cartwright, M. J., Camacho, D. M., Paré, J-F., Super, M., Levin, M., and Ingber, D. E.
Advanced Science (Weinheim), 9(26): e2200222 [cover]

Information Theory as an Experimental Tool for Integrating Disparate Biophysical Signaling Modules
McMillen, P., Walker, S. I., and Levin, M.
International Journal of Molecular Sciences, 23(17): 9580

Studying Protista WBR and Repair Using Physarum polycephalum
Sperry, M. M., Murugan, N. J., and Levin, M.
Methods in Molecular Biology, 2450: 51-67

Screening Biophysical Sensors and Neurite Outgrowth Actuators in Human Induced-Pluripotent-Stem-Cell-Derived Neurons
Pai, V. P., Cooper, B. G., and Levin, M.
Cells, 11(16): 2470

Emergence as the conversion of information: a unifying theory
Varley, T. F., and Hoel, E.
Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Sciences, 380(2227): 20210150

Competency in Navigating Arbitrary Spaces as an Invariant for Analyzing Cognition in Diverse Embodiments
Fields, C., and Levin, M.
Entropy (Basel), 24(6): 819

Screening neuroprotective compounds in herpes-induced Alzheimer’s disease cell and 3D tissue models
Silveira, I. A., Mullis, A. S., Cairns, D. M., Shevzov-Zebrun, A., Whalen, J., Galuppo, A., Walsh, K. G., and Kaplan, D. L.
Free Radical Biology and Medicine, 186: 76-92

Causal emergence is widespread across measures of causation
Comolatti, R., and Hoel, E.
ArXiv, 24: 107

Drug Repositioning Using Multiplex-Heterogeneous Network Embedding: A Case Study on SARS-CoV2
Pio-Lopez L.
In: Benito R.M., Cherifi C., Cherifi H., Moro E., Rocha L.M., Sales-Pardo M. (eds) Complex Networks & Their Applications X. COMPLEX NETWORKS 2021.
Studies in Computational Intelligence, Vol 1016

Inducing Vertebrate Limb Regeneration: A Review of Past Advances and Future Outlook
Davidian, D., and Levin, M.
Cold Spring Harbor Perspectives in Biology, 14(4): a040782

Biology, Buddhism, and AI: Care as the Driver of Intelligence
Doctor, T., Witkowski, O., Solomonova, E., Duane, B., and Levin, M.
Entropy (Basel), 24(5): e24050710

The genetic control paradigm in biology: What we say, and what we are entitled to mean
McKenna, K. Z., Gawne, R., and Nijhout, H. F.
Progress in Biophysics and Molecular Biology, 169-170: 89-93

Design for an Individual: Connectionist approaches to the evolutionary transitions in individuality
Watson, R. A., Levin, M., and Buckley, C. L.
Frontiers in Ecology and Evolution, 10: 823588

Technological Approach to Mind Everywhere: An Experimentally-Grounded Framework for Understanding Diverse Bodies and Minds
Levin, M.
Frontiers in Systems Neuroscience, 16: 768201

Metacognition as a Consequence of Competing Evolutionary Time Scales
Kuchling, F., Fields, C., and Levin, M.
Entropy (Basel), 24(5): 601

Multi-scale Chimerism: an experimental window on the algorithms of anatomical control
Nanos, V., and Levin, M.
Cells and Development, 169: 203764 [cover]

A Computational Approach to Explaining Bioelectrically-induced Persistent, Stochastic Changes of Axial Polarity in Planarian Regeneration
Grodstein, J., and Levin, M.
Bioelectricity, 4(1): 18-30

Ion Channel Drugs Suppress Cancer Phenotype in NG108-15 and U87 Cells: Toward Novel Electroceuticals for Glioblastoma
Mathews, J., Kuchling, F., Baez-Nieto, D., Diberardinis, M., Pan, J. Q., and Levin, M.
Cancers, 14(6): 1499

Impact of Membrane Voltage on Formation and Stability of Human Renal Proximal Tubules in Vitro
Adelfio, M., Bonzanni, M., Levin, M., and Kaplan, D. L.
ACS Biomaterials Science & Engineering, 8(3): 1239-1246

Correction to: When left does not seem right: epigenetic and bioelectric differences between left and right‑sided breast cancer
Masuelli, S., Real, S., Campoy, E., Branham, M. T., Marzese, D. M., Salomon, M., De Blas, G., Arias, R., Levin, M., and Roque, M.
Molecular Medicine (Cambridge, Mass.), 28(1): 25

When left does not seem right: epigenetic and bioelectric differences between left- and right-sided breast cancer
Masuelli, S., Real, S., Campoy, E., Branham, M. T., Marzese, D. M., Salomon, M., De Blas, G., Arias, R., Levin, M., and Roque, M.
Molecular Medicine (Cambridge, Mass.), 28(1): 15

Acute multidrug delivery via a wearable bioreactor facilitates long-term limb regeneration and functional recovery in adult Xenopus laevis
Murugan, N. J., Vigran, H. J., Miller, K., Golding, A., Pham, Q. L., Sperry, M., Rasmussen-Ivey, C., Kane, A. W., Kaplan, D. L., and Levin, M.
Science Advances, 8(4): abj2164 [cover]

Minimal developmental computation: a causal network approach to understand morphogenetic pattern formation
Manicka, S., and Levin, M.
Entropy (Basel), 24(1): 107

Potassium channel-driven bioelectric signalling regulates metastasis in triple-negative breast cancer
Payne, S. L., Ram, P., Srinivasan, D. H., Le, T. T., Levin, M., and Oudin, M. J.
EBioMedicine, 75: 103767

2021

The rise of the biocyborg: synthetic biology, artificial chimerism and human enhancement
Pio-Lopez L.
New Genetics and Society, 40(4): 599-619

New Clues about the Origins of Biological Intelligence
Yuste, R., and Levin, M.
Scientific American, December 2021

Kinematic self-replication in reconfigurable organisms
Kriegman, S., Blackiston, D., Levin, M., and Bongard, J.
Proceedings of the National Academy of Sciences of the United States, 118(49): e2112672118

Behaviorist Approaches to Investigating Memory and Learning: A Primer for Synthetic Biology and Bioengineering
Abramson, C., and Levin, M.
Communicative & Integrative Biology, 14(1): 230-247

Cell Systems Bioelectricity: How Different Intercellular Gap Junctions Could Regionalize a Multicellular Aggregate
Riol, A., Cervera, J., Levin, M., and Mafe, S.
Cancers (Basel), 13(21): 5300

Stability and robustness properties of bioelectric networks: A computational approach
Grodstein, J., and Levin, M.
Biophysics Review, 2(3): 031305

Unmixing octopus camouflage by multispectral mapping of Octopus bimaculoides‘ chromatic elements
Guidetti, G., Levy, G., Matzeu, G., Finkelstein, J. M., Levin, M., and Omenetto, F. G.
Nanophotonics, 10(9): 2441-2450

Bioelectrical approaches to cancer as a problem of the scaling of the cellular self
Levin, M.
Progress in Biophysics and Molecular Biology, 165: 102-113

Morphology changes induced by intercellular gap junction blocking: a reaction-diffusion mechanism
Cervera, J., Levin, M., and Mafe, S.
BioSystems, 209: 104511

Metabolic limits on classical information processing by biological cells
Fields, C., and Levin, M.
BioSystems, 209: 104513

Beyond Neurons: Long Distance Communication in Development and Cancer
McMillen, P., Oudin, M. J, Levin, M., and Payne, S. L.
Frontiers in Cell and Developmental Biology, 9: 2637

Minimal physicalism as a scale-free substrate for cognition and consciousness
Fields, C., Glazebrook, J. F., and Levin, M.
Neuroscience of Consciousness, 2021(2): niab013

Mechanosensation Mediates Long-Range Spatial Decision-Making in an Aneural Organism
Murugan, N. J., Kaltman, D. H., Jin, P. H., Chien, M., Martinez, R., Nguyen, C. Q., Kane, A., Novak, R., Ingber, D. E., and Levin, M.
Advanced Materials, 33(34): 2008161 [cover]

Falsification and consciousness
Kleiner, J., and Hoel, E.
Neuroscience of Consciousness, 7(1): niab001

The overfitted brain: Dreams evolved to assist generalization
Hoel, E.
Patterns, 2(5): 100244

Life, death, and self: Fundamental questions of primitive cognition viewed through the lens of body plasticity and synthetic organisms
Levin, M.
Biochemical and Biophysical Research Communications, 564: 114-133 [cover]

Synthetic living machines: A new window on life
Ebrahimkhani, M., and Levin, M.
iScience, 24(5): 102505

A cellular platform for the development of synthetic living machines
Blackiston, D., Lederer, E., Kriegman, S., Garnier, S., Bongard, J., and Levin, M.
Science Robotics, 6(52): eabf1571

Learning and synaptic plasticity in 3D bioengineered neural tissues
Cairns, D. M., Rusk, W., Levin, M., Rouleau, N., and Kaplan, D.
Neuroscience Letters, 750: 135799

Bioelectric Signaling: Reprogrammable Circuits Underlying Embryogenesis, Regeneration, and Cancer
Levin, M.
Cell, 184(8): 1971-1989

Bistability of Somatic Pattern Memories: Stochastic Outcomes in Bioelectric Circuits Underlying Regeneration
Pezzulo, G., Lapalme, J., Durant, F., and Levin, M.
Philosophical Transactions of the Royal Society B: Biological Sciences, 376(1821): 20190765

Living things are not (20th Century) machines: updating mechanism metaphors in light of the modern science of machine behavior
Bongard, J., and Levin, M.
Frontiers in Ecology and Evolution, 9: 650726

Uncovering cognitive similarities and differences, conservation and innovation
Levin, M., Keijzer, F., Lyon, P., and Arend, D.
B, 376(1821): 20200458 [cover]

Editorial: Interplay Between Ion Channels, the Nervous System, and Embryonic Development
Levin, M., and Ribera, A. B.
Frontiers in Molecular Neuroscience, 14: 618815

Valuing what happens: a biogenic approach to valence and (potentially) affect
Lyon, P., and Kuchling, F.
Philosophical Transactions of the Royal Society B: Biological Sciences, 376(1820): 201907525

Reframing cognition: getting down to biological basics
Lyon, P., Keijzer, F., Arendt, D., and Levin, M.
Philosophical Transactions of the Royal Society B: Biological Sciences, 376(1820): 20190750 [cover]

Shape Changing Robots: Bioinspiration, Simulation, and Physical Realization
Shah, D., Yang, B., Kriegman, S., Levin, M., Bongard, J., and Kramer-Bottiglio, R.
Advanced Materials, 33(19): e2002882

A Meta-Analysis of Bioelectric Data in Cancer, Embryogenesis, and Regeneration
Srivastava, P., Kane, A., Harrison, C., and Levin, M.
Bioelectricity, 3(1): 42-67

Editorial
Djamgoz, M. B. A., and Levin, M.
Bioelectricity, 3(1): 2

Epigenetic control of myeloid cells behavior by Histone Deacetylase activity (HDAC) during tissue and organ regeneration in Xenopus laevis
Pentagna, N., Pinheiro da Costa, T., Soares Dos Santos Cardoso, F., Martins de Almeida, F., Blanco Martinez, A. M., Abreu, J. G., Levin, M., and Carneiro, K.
Developmental and Comparative Immunology, 114: 103840

Rewiring Endogenous Bioelectric Circuits in the Xenopus laevis Embryo Model
Nanos, V., and Levin, M.
Methods in Molecular Biology, 2258: 93-103

Gene Regulatory Networks Exhibit Several Kinds of Memory: Quantification of Memory in Biological and Random Transcriptional Networks
Biswas, S., Manicka, S., Hoel, E., and Levin, M.
iScience, 24(3): 102131

A Comprehensive Conceptual and Computational Dynamics Framework for Autonomous Regeneration Systems
Minh-Thai, T. N., Samarasinghe, S., and Levin, M.
Artificial Life, 27(2): 80-104

2020

Causal Geometry
Chvykov, P., & Hoel, E.
Entropy (Basel), 23(1): 24

Examining the Causal Structures of Deep Neural Networks Using Information Theory
Marrow, S., Michaud, E. J., and Hoel, E.
Entropy (Basel), 22(12): 1429

Why isn’t sex optional? Stem-cell competition, loss of regenerative capacity, and cancer in metazoan evolution
Fields, C., and Levin, M.
Communicative & Integrative Biology, 13(1): 170-183

Community effects allow bioelectrical reprogramming of cell membrane potentials in multicellular aggregates: Model simulations
Cervera, J., Ramirez, P., Levin, M., and Mafe, S.
Physical Review E, 102(5-1): 052412

Effect of a Prostate Cancer Screening Decision Aid for African-American Men in Primary Care Settings
Allen, J. D., Filson, C. P., and Berry, D. L.
Cancer Epidemiology, Biomarkers & Prevention, 29(11): 2157-2164

Machine Learning-Driven Bioelectronics for Closed-Loop Control of Cells
Selberg, J., Jafari, M., Mathews, J., Jia, M., Pansodtee, P., Dechiraju, H., Wu, C., Cordero, S., Flora, A., Yonas, N., Jannetty, S., Diberardinis, M., Teodorescu, M., Levin, M., Gomez, M., and Rolandi, M.
Advanced Intelligent Systems, 2: 2000140 [cover]

Precise control of ion channel and gap junction expression is required for patterning of the regenerating axolotl limb
Sousounis, K., Erdogan, B., Levin, M., and Whited, J.
International Journal of Developmental Biology, 64: 485-494

How to understand cells, tissues and organisms as agents with agendas
Levin, M., and Dennett, D. C.
Aeon Essays

Shape changing robots: bioinspiration, simulation, and physical realization
Shah, D., Yang, B., Kriegman, S., Levin, M., Bongard, J., and Kramer-Bottiglio, R.
Advanced Materials, 33(19): 2002882

Does evolution have a target morphology?
Fields, C., and Levin, M.
Organisms, 4(1): 57-76

Bioelectronic Control of Chloride Ions and Concentration with Ag/AgCl Contacts
Jia, M., Dechiruji, H., Selberg J., Pansodtee P., Mathews, J., Wu, C., Levin, M., Teodorescu, M., and Rolandi, M.
APL Materials, 8(9): 091106</p

How Groups of Cells Cooperate to Build Organs and Organisms
Levin, M.
The Scientist, September 2020 [cover]

Nervous System and Tissue Polarity Dynamically Adapt to New Morphologies in Planaria
Bischof, J., Day, M., Miller, K., LaPalme, J., and Levin, M.
Developmental Biology, 467 (1-2): 51-65 [cover]

Emergence of Informative Higher Scales in Biological Systems: a computational toolkit for optimal prediction and control
Hoel, E., and Levin, M.
Communicative & Integrative Biology, 13(1): 108-118

A 3D tissue model of traumatic brain injury with excitotoxicity that is inhibited by chronic exposure to gabapentinoids
Rouleau, N., Bonzanni, M., Erndt-Marino, J. D., Sievert, K., Ramirez, C., Rusk, W., Levin, M., and Kaplan, D. L.
Biomolecules, 10(8): 1196

Matrix Deformation with Ectopic Cells Induced by Rotational Motion in Bioengineered Neural Tissues
Rouleau, N., Murugan, N. J., Rusk, W., Koester, C., and Kaplan, D. L.
Annals of Biomedical Engineering, 48(8): 2192-2203

Scale-Free Biology: Integrating Evolutionary and Developmental Thinking
Fields, C., and Levin, M.
Bioessays, 42(8): e1900228

Applications and ethics of computer-designed organisms
Levin, M., Bongard, J., and Lunshof, J. E.
Nature Reviews Molecular Cell Biology, 21: 655–656

Opinion: Use the Pandemic to Expand the Lab to the Home
Levin, M.
The Scientist

Regulation of axial and head patterning during planarian regeneration by a commensal bacterium
Williams, K., Bischof, J., Lee, F., Miller, K., LaPalme, J., Wolfe, B., and Levin, M.
Mechanisms of Development, 163: 103614 [cover]

The effects of membrane potential and extracellular matrix composition on vascular differentiation of cardiac progenitor cells
Daley, M. C., Bonzanni, M., MacKenzie, A. M., Kaplan, D. L., and Black, L. D., 3rd
Biochemical and Biophysical Research Communications, 530(1): 240-245

Bioelectricity: A Quick Reminder of a Fast-Advancing Discipline!
Djamgoz, M. B. A., and Levin, M.
Bioelectricity, 2(3): 208-209

Competitive and Coordinative Interactions between Body Parts Produce Adaptive Developmental Outcomes
Gawne, R., McKenna, K. Z., and Levin, M.
BioEssays, 42(8): 1900245 [cover]

HCN2 Channel-induced Rescue of Brain Teratogenesis via local and long-range bioelectric repair
Pai, V., Cervera, J., Mafe, S., Willocq, V., Lederer, E., and Levin, M.
Frontiers in Cellular Neuroscience, 14: 136

Bioelectrical Coupling of Single-Cell States in Multicellular Systems
Cervera, J., Levin, M., and Mafe, S.
The Journal of Physical Chemistry Letters, 11(9): 3234-3241

The Epithelial adhesin 1 tandem repeat region mediates protein display through multiple mechanisms
Raposo, C. J., McElroy, K. A., and Fuchs, S. M.
FEMS Yeast Research, 20(3): foaa018

von Willebrand factor D and EGF domains is an evolutionarily conserved and required feature of blastemas capable of multitissue appendage regeneration
Leigh, N. D., Sessa, S., Dragalzew, A. C., Payzin‐Dogru, D., Sousa, J. F., Aggouras, A. N., Johnson, K., Dunlap, G. S., Haas, B. J., Levin, M., Schneider, I., and Whited, J. L.
Evolution & Development, 22(4): 297-311 [cover]

Integrating variational approaches to pattern formation into a deeper physics: Reply to comments on “Morphogenesis as Bayesian inference: A variational approach to pattern formation and manipulation in complex biological systems”
Kuchling, F., Friston, K., Georgiev, G., and Levin, M.
Physics of Life Reviews, 33: 125-128

Morphogenesis as Bayesian inference: A variational approach to pattern formation and control in complex biological systems
Kuchling, F., Friston, K., Georgiev, G., and Levin, M.
Physics of Life Reviews, 33: 88-108

On the coupling of mechanics with bioelectricity and its role in morphogenesis
Leronni, A., Bardella, L., Dorfmann, L., Pietak, A., and Levin, M.
Journal of the Royal Society Interface, 17(167): 20200177

A 3D human brain-like tissue model of herpes-induced Alzheimer’s disease
Cairns, D. M., Rouleau, N., Parker, R. N., Walsh, K. G., Gehrke, L., and Kaplan, D. L.
Science Advances, 6(19): eaay8828

A Long-Living Bioengineered Neural Tissue Platform to Study Neurodegeneration
Rouleau, N., Cantley, W. L., Liaudanskaya, V., Berk, A., Du, C., Rusk, W., Peirent, E., Koester, C., Nieland, T. J. F., and Kaplan, D. L.
Macromolecular Bioscience, 20(3): e2000004

Sertraline induces DNA damage and cellular toxicity in Drosophila that can be ameliorated by antioxidants
Jajoo, A., Donlon, C., Shnayder, S., Levin, M., and McVey, M.
Scientific Reports, 10(1): 4512

Revisiting Burr and Northrop’s ‘The Electro-Dynamic Theory of Life’
Levin, M.
Biological Theory, 15: 83-90

Eya2 promotes cell cycle progression by regulating DNA damage response during vertebrate limb regeneration
Sousounis, K., Bryant, D. M., Martinez Fernandez, J., Eddy, S. S., Tsai, S. L., Gundberg, G. C., Han, J., Courtemanche, K., Levin, M., and Whited, J. L.
eLIFE, 9: e51217

Post-SSRI Sexual Dysfunction: A Bioelectric Mechanism?
Healy, D., LaPalme, J., and Levin, M.
Bioelectricity, 2(1): 7-13

Interferon-Gamma Stimulated Murine Macrophages In Vitro: Impact of Ionic Composition and Osmolarity and Therapeutic Implications
HErndt-Marino, J., Yeisley, D. J., Chen, H., Levin, M., Kaplan, D. L., and Hahn, M. S.
Bioelectricity, 2(1): 48-58

Inaugural Issue
Djamgoz, M. B. A., and Levin, M.
Bioelectricity, 2(1): 1

Does regeneration recapitulate phylogeny?
Fields, C., and Levin, M.
Communicative & Integrative Biology, 13(1): 27-38

An in vivo brain–bacteria interface: the developing brain as a key regulator of innate immunity
Herrera-Rincon, C., Paré, J-F., Martyniuk, C. J., Jannetty, S. K., Harrison C., Fischer, A., Dinis, A., Keshari, V., Novak, R., and Levin, M.
NPJ Regenerative Medicine, 5: 2

Extra-genomic instructive influences in morphogenesis: A review of external signals that regulate growth and form
Tung, A., and Levin, M.
Developmental Biology, 461(1): 1-12 [cover]

The Biophysics of Regenerative Repair Suggests New Perspectives on Biological Causation
Levin, M.
BioEssays, 42(2): 1900146

A scalable pipeline for designing reconfigurable organisms
Kriegman, S., Blackiston, D., Levin, M., and Bongard, J.
Proceedings of the National Academy of Sciences of the United States, 117(4): 1853-1859

Assessment of Enrichment of Human Mesenchymal Stem Cells Based on Plasma and Mitochondrial Membrane Potentials
Kamaldinov, T., Erndt-Marino, J., Levin, M., Kaplan, D. L., and Hahn, M. S.
Bioelectricity, 2(1): 21-32

Bioelectrical model of head-tail patterning based on cell ion channels and intercellular gap junctions
Cervera, J., Meseguer, S., Levin, M., and Mafe, S.
Bioelectrochemistry, 132: 107410

Defined extracellular ionic solutions to study and manipulate the cellular resting membrane potential
Bonzanni, M., Payne, S. L., Adelfio, M., Kaplan, D. L., Levin, M., and Oudin, M. J.
Biology Open, 9(1): bio048553

Optogenetically induced cellular habituation in non-neuronal cells
Bonzanni, M., Rouleau, N., Levin, M., and Kaplan, D. L.
PLoS One, 15(1): e0227230

Toward Decoding Bioelectric Events in Xenopus Embryogenesis: New Methodology for Tracking Interplay Between Calcium and Resting Potentials In Vivo
McMillen, P., Novak, R., and Levin, M.
Journal of Molecular Biology, 432(2): 605-620 [cover]

Morphological Coordination: A Common Ancestral Function Unifying Neural and Non-Neural Signaling
Fields, C., Bischof, J., and Levin, M.
Physiology (Bethesda), 35(1): 16-30

2019

Planarian Regeneration as a Model of Anatomical Homeostasis: Recent Progress in Biophysical and Computational Approaches
Levin, M., Pietak, A. M., Bischof, J.
Seminars in Cell and Developmental Biology, 87: 125-144

Membrane potential depolarization alters calcium flux and phosphate signaling during osteogenic differentiation of human mesenchymal stem cells
Sundelacruz, S., Levin, M., and Kaplan, D.
Bioelectricity, 1(1): 56-66

The Bioelectricity Revolution: A Discussion Among the Founding Associate Editors
Adams, D. S., Djamgoz, M., Levin, M., Zhao, M., Rajnicek, A., Nuccitelli, R., Arcangeli, A., Kramer, R., and Bates, E. A.
Bioelectricity, 1(1): 8-15

Defined extracellular ionic solutions to study and manipulate the cellular resting membrane potential
Bonzanni, M., Payne, S. L., Adelfio, M., Kaplan, D. L., Levin, M., and Oudin, M. J.
Biology Open, 9(1): bio048553

Endogenous Bioelectrics in Development, Cancer, and Regeneration: Drugs and Bioelectronic Devices as Electroceuticals for Regenerative Medicine
Levin, M., Selberg, S., and Rolandi, M.
iScience, 22: 519–533

Modeling somatic computation with non-neural bioelectric networks
Manicka, S., and Levin, M.
Scientific Reports, 9(1): 18612

Regenerative Adaptation To Electrochemical Perturbation In Planaria: A Molecular Analysis Of Physiological Plasticity
Emmons-Bell, M., Durant, F., Tung, A., Pietak, A., Miller, K., Kane, A., Martyniuk, C. J., Davidian, D., Morokuma, J., and Levin, M.
iScience, 22: 147-165

Interdisciplinary Approach Needed to Crack Morphogenesis
Finkelstein J., McLaughlin K., and Levin M.
The Scientist, December 2019

The Computational Boundary of a “Self”: developmental bioelectricity drives multicellularity and scale-free cognition
Levin, M.
Frontiers in Psychology, 10: 2688

L-type voltage-gated Ca2+ channel CaV1.2 regulates chondrogenesis during limb development
Atsuta, Y., Tomizwa, R. R., Levin, M., and Tabin, C. J.
Proceedings of the National Academy of Sciences of the United States, 116(43): 21592-21601

Hyperosmolar potassium inhibits myofibroblast conversion and reduces scar tissue formation
Grasman, J., Williams, M., Razis, C., Bonzanni, M., Golding, A., Cairns, D. M., Levin, M., and Kaplan, D. L.
ACS Biomaterials Science & Engineering, 5(10): 5327-5336

Managing Ideas, People, and Projects: Organizational Tools and Strategies for Researchers
Levin, S. P., and Levin, M.
iScience, 20: 278-291

Bioelectric control of metastasis in solid tumors
Payne, S. L., Levin, M., and Oudin, M. J.
Bioelectricity, 1(3): 114-130

Somatic multicellularity as a satisficing solution to the prediction-error minimization problem
Fields, C., and Levin, M.
Communicative & Integrative Biology, 12(1): 119-132

Bioelectrical controls of morphogenesis: from ancient mechanisms of cell coordination to biomedical opportunities
Whited, J., and Levin, M.
Current Opinion in Genetics & Development, 57: 61-69 [cover]

Reverse-engineering growth and form in Heidelberg
Levin, M., and Martinez Arias, A.
Development, 146(14): dev177261

A flow through device for simultaneous dielectrophoretic cell trapping and AC electroporation
Punjiya, M., Nejad, H. R., Mathews, J., Levin, M., and Sonkusale, S.
Scientific Reports, 9(1): 11988

From non-excitable single-cell to multicellular bioelectrical states supported by ion channels and gap junction proteins: electrical potentials as distributed controllers
Cervera, C., Pai, V. P., Levin, M., and Mafe, S.
Progress in Biophysics and Molecular Biology, 149: 39-53 [cover]

On the Generalization of Habituation: How Discrete Biological Systems Respond to Repetitive Stimuli: A Novel Model of Habituation That Is Independent of Any Biological System
Bonzanni, M., Rouleau, N., Levin, M., and Kaplan, D. L.
Bioessays, 41(7): e1900028

Adaptive correction of craniofacial defects in pre-metamorphic Xenopus laevis tadpoles involves thyroid hormone-independent tissue remodeling
Pinet, K., Deolankar, M., Leung, B., and McLaughlin, K. A.
Development, 146(14): dev175893

Mechanisms of physiological tissue remodeling in animals: Manipulating tissue, organ, and organism morphology
Pinet, K., and McLaughlin, K .A.
Developmental Biology, 451(2): 134-145 [cover]

Recovery of the Xenopus laevis heart from ROS-induced stress utilizes conserved pathways of cardiac regeneration
Jewhurst, K., and McLaughlin, K. A.
Development, Growth & Differentiation, 61(3): 212-227

Synchronization of Bioelectric Oscillations in Networks of Nonexcitable Cells: From Single-Cell to Multicellular States
Cervera, J., Manzanares, J. A., Mafe, S., and Levin, M.
Journal of Physical Chemistry B, 123(18): 3924-3934 [cover]

On “Isomerization as the secret Achilles’ heel of long-lived proteins”
Taylor, A.
Journal of Biological Chemistry, 294(25): 9689

The Cognitive Lens: a primer on conceptual tools for analysing information processing in developmental and regenerative morphogenesis
Manicka, S., and Levin, M.
Philosophical Transactions of the Royal Society B: Biological Sciences, 374(1774): 20180369

Neural control of body-plan axis in regenerating planaria
Pietak, A., Bischof, J., LaPalme, J., Morokuma, J., and Levin, M.
PLOS Computational Biology, 15(4): e1006904

The Role of Early Bioelectric Signals in the Regeneration of Planarian Anterior/Posterior Polarity
Durant, F., Bischof, J., Fields, C., Morokuma, J., LaPalme, J., Hoi, A., and Levin, M.
Biophysical Journal, 116(5): 948-961

Selective Serotonin Reuptake Inhibitor Use During Pregnancy and Major Malformations: The Importance of Serotonin for Embryonic Development and the Effect of Serotonin Inhibition on the Occurrence of Malformations
Bérard, A., Levin, M., Sadler, T., and Healy, D.
Bioelectricity, 1(1): 18-29

Inter-generational consequences for growing Caenorhabditis elegans in liquid
Lev, I., Bril, R., Liu, Y., Cere, L. I., and Rechavi, O.
Philosophical Transactions of the Royal Society B: Biological Sciences, 374(1770): 20180125

Publisher Correction: A call for a better understanding of causation in cell biology
Bizzarri, M., Brash, D. E., Briscoe, J., Grieneisen, V. A., Stern, C. D., and Levin, M.
Nature Reviews Molecular Cell Biology, 20(6): 384

A Call for a Better Understanding of Causation in Cell Biology
Bizzarri, M., Brash, D., Briscoe, J., Grieneisen, V., Stern, C. D., and Levin, M.
Nature Reviews Molecular Cell Biology, 20(5): 261-262

What Caused What? A Quantitative Account of Actual Causation Using Dynamical Causal Networks
Albantakis, L., Marshall, W., Hoel, E., and Tononi, G.
Entropy (Basel), 21(5): 459

Toward modeling regeneration via adaptable echo state networks
Hammelman, J., Siegelmann, H., Manicka, S., and Levin, M., in A. Adamatzky, S. Akl, and G. Sirakoulis (Eds.), From parallel to emergent computing, Chapter 6, pp. 117-133, CRC Press

Effects of ivermectin exposure on regeneration of D. dorotocephala planaria: exploiting human-approved ion channel drugs as morphoceuticals
Ferenc, N. N., and Levin, M.
Macromolecular Bioscience, 19(3): 1800237 [cover]

Functional Effects of a Neuromelanin Analogue on Dopaminergic Neurons in 3D Cell Culture
Collins, W., Rouleau, N., Bonzanni, M., Kapner, K., Jeremiah, A., Du, C., Pothos, E. N., and Kaplan, D. L.
ACS Biomaterials Science & Engineering, 5(1): 308-317

EDEn – Electroceutical Design Environment: An Ion Channel Database with Small Molecule Modulators and Tissue Expression Information
Churchill, C. D. M., Winter, P., Tuszynski, J. A., and Levin, M.
iScience, 11: 42-56

2018

Mechanistic targeting of advanced glycation end-products in age-related diseases
Rowan, S., Bejarano, E., and Taylor, A.
Biochimica et Biophysica Acta – Molecular Basis of Disease, 1864(12): 3631-3643

The bacterial metabolite indole inhibits regeneration of the planarian flatwork Dugesia japonica
Lee, F. J., Williams, K. B., Levin, M., and Wolfe, B. E.
iScience, 10: 135-148

Brief Local Application of Progesterone via a Wearable Bioreactor Induces Long-Term Regenerative Response in Adult Xenopus Hindlimb
Herrera-Rincon, C., Golding, A. S., Moran, Harrison, C., Martyniuk, C. J., Guay, J. A., Zaltsman, J., Carabello, H., Kaplan, D. L., and Levin, M.
Cell Reports, 25(6): 1593-1609

Entropic effects in cell lineage tree packings
Alsous, J. I., Villoutreix, P., Stoop, N., Shvartsman, S. Y., and Dunkel, J.
Nature Physics, 14: 1016–1021 [cover]

Ivermectin promotes peripheral nerve regeneration during wound healing
Cairns, D. M., Giordano, J. E., Conte, S., Levin, M., and Kaplan, D. L.
ACS Omega, 3(10): 12392-12402

Activating PAX gene family paralogs to complement PAX5 leukemia driver mutations
Hart, M. R., Anderson, D. J., Porter, C. C., Neff, T., Levin, M., and Horwitz, M. S.
PLoS Genetics, 4(9): e1007642

Cross-limb Communication During Xenopus Hind-limb Regenerative Response: Non-Local Bioelectric Injury Signals
Busse, S. M., McMillen, P. T., and Levin, M.
Development, 145(19): dev164210

Inform: Efficient Information-Theoretic Analysis of Collective Behaviors
Moore, D. G., Valentini, G., Walker, S. I., and Michael Levin
Frontiers in Robotics and AI, 5: 60

Bioelectrical coupling in multicellular domains regulated by gap junctions: a conceptual approach
Cervera, J., Pietak, A., Levin, M., and Mafe, S.
Bioelectrochemistry, 123: 45-61 [cover]

Bioelectrical control of positional information in development and regeneration: a review of conceptual and computational advances
Pietak, A., and Levin, M.
Progress in Biophysics and Molecular Biology 137: 52-68

The Body Electric 2.0: Recent Advances in Developmental Bioelectricity for Regenerative and Synthetic Bioengineering
Mathews, J., and Levin, M.
Current Opinion in Biotechnology 52: 134-144

Are planaria individuals? What regenerative biology is telling us about the nature of multicellularity
Fields, C., and Levin, M.
Evolutionary Biology, 45: 237–247

Embodying Markov blankets: Comment on “Answering Schrödinger’s question: A free-energy formulation” by Maxwell James Désormeau Ramstead et al.
Pezzulo, G., and Levin, M.
Physics of Life Reviews, 24: 32-36

When the VEST Does Not Fit: Representations of Trial Results Deviating From Rigorous Data Interpretation
Allen, L. A., Adler, E. D., Bayes-Genis, A., Brisco-Bacik, M. A., Chirinos, J. A., Claggett, B., Cook, J. L., Fang, J. C., Gustafsson, F., Ho, C. Y., Kapur, N. K., Klewer, S. E., Kociol, R. D., Lanfear, D. E., Vardeny, O., and Sweitzer, N. K.
Circulation: Heart Failure, 11(4): e005116

Corrigendum to “Slime mould: The fundamental mechanisms of biological cognition” [BioSystems 165 (2018) 57-70]
Vallverdu, J., Castro, O., Mayne, R., Talanov, M., Levin, M., Baluska, F., Gunji, Y., Dussutour, A., Zenil, H., and Adamatzky, A.
Biosystems, 166: 66

Slime mould: The fundamental mechanisms of biological cognition
Vallverdu, J., Castro, O., Mayne, R., Talanov, M., Levin, M., Baluska, F., Gunji, Y., Dussutour, A., Zenil, H., and Adamatzky, A.
Biosystems, 165: 57-70

Multiscale memory and bioelectric error correction in the cytoplasm-cytoskeleton-membrane system
Fields, C., and Levin, M.
Wiley Interdisciplinary Reviews Systems Biology and Medicine, 10(2): e1410 [cover]

HCN2 Rescues brain defects by enforcing endogenous voltage pre-patterns
Pai, V. P., Pietak, A., Willocq, V., Ye, B., Shi, N. Q., and Levin, M.
Nature Communications, 9(1): 998

Inverse Drug Screening of Bioelectric Signaling and Neurotransmitter Roles: Illustrated Using a Xenopus Tail Regeneration Assay
Sullivan, K. G., and Levin, M.
Cold Spring Harbor Protocols, 2018(3): 154-163

Booting up the organism during development: Pre-behavioral functions of the vertebrate brain in guiding body morphogenesis
Herrera-Rincon, C., and Levin, M.
Communicative & Integrative Biology, 11(1): e1433440 [cover]

The bioelectric code: An ancient computational medium for dynamic control of growth and form
Levin, M., and Martyniuk, C. J.
Biosystems, 164: 76-93

Niclosamide rescues microcephaly in a humanized in vivo model of Zika infection using human induced neural stem cells
Cairns, D.M., Boorgu, D. S. S. K., Levin, M., and Kaplan, D. L.
Biology Open, 7(1): bio031807

Bioelectric signaling in regeneration: Mechanisms of ionic controls of growth and form
McLaughlin, K. A., and Levin, M.
Developmental Biology, 433(2): 177-189

2017

Identification of regenerative roadblocks via repeat deployment of limb regeneration in axolotls
Bryant, D. M., Sousounis, K., Payzin-Dogru, D., Bryant, S., Sandoval, A. G. W., Martinez Fernandez, J., Mariano, R., Oshiro, R., Wong, A. Y., Leigh, N. D., Johnson, K., and Whited, J. L.
NPJ Regenerative Medicine, 2: 30

HCN4 Ion Channel Function is Required for Early Events that Regulate Anatomical Left-Right Patterning in a Nodal- and Lefty-Independent Manner
Pai, V. P., Willocq, V., Pitcairn, E. J., Lemire, J. M., Paré, J-F., Shi, N. Q., McLaughlin, K. A., and Levin, M.
Biology Open, 6(10): 1445-1457

Bioelectric gene and reaction networks: computational modelling of genetic, biochemical and bioelectrical dynamics in pattern regulation
Pietak, A., and Levin, M.
Journal of the Royal Society Interface, 14(134): 20170425 [cover]

The brain is required for normal muscle and nerve patterning during early Xenopus development
Herrera-Rincon, C., Pai, V. P., Moran, K. M., Lemire, J. M., and Levin, M.
Nature Communications, 8(1): 587

Collective Growth in a Small Cell Network
Alsous, J. I., Villoutreix, P., Berezhkovskii, A. M., and Shvartsman, S. Y.
Current Biology, 27(17): 2670-2676

Space travel has effects on planarian regeneration that cannot be explained by a null hypothesis
Levin, M., Morokuma, J., and Finkelstein, J.
Regeneration (Oxf), 4(4): 156-158

The Zahn Drawings: New Illustrations of Xenopus Embryo and Tadpole Stages for Studies of Craniofacial Development
Zahn, N., Levin, M., and Adams, D. S.
Development, 144(15): 2708-2713 [cover]

Endogenous Bioelectric Signaling Networks: Exploiting Voltage Gradients for Control of Growth and Form
Levin, M., Pezzulo, G., and Finkelstein, J. M.
Annual Review of Biomedical Engineering, 19: 353-387

Planarian regeneration in space: persistent anatomical, behavioral, and bacteriological changes induced by space travel
Morokuma, J., Durant, F., Williams, K. B., Finkelstein, J. M., Blackiston, D. J., Clements, T., Reed, D. W., Roberts, M., Jain, M., Kimel, K., Trauger, S. A., Wolfe, B. E., and Levin, M.
Regeneration (Oxf), 4(2): 85-102 [cover]

Bioelectric regulation of innate immune system function in regenerating and intact Xenopus laevis
Paré, J-F., Martyniuk, C. J., and Levin, M.
NPJ Regenerative Medicine, 2: 15

Long-term, stochastic editing of regenerative anatomy via targeting endogenous bioelectric gradients
Durant, F., Morokuma, J., Fields, C., Williams, K., Adams, D. S., and Levin, M.
Biophysical Journal, 2112(10): 2231-2243 [cover]

Coordinating heart morphogenesis: a novel role for Hyperpolarization-activated cyclic nucleotide-gated (HCN) channels during cardiogenesis in Xenopus laevis
Pitcairn, E., Harris, H., Epiney, J., Pai, V. P., Lemire, J. M., Ye, B., Shi, N. Q., Levin, M., and McLaughlin, K. A.
Communicative & Integrative Biology, 10(3): e1309488 [cover]

Ion channel and neurotransmitter modulators as electroceutical approaches to the control of cancer
Tuszynski, J., Tilli, T. M., and Levin, M.
Current Pharmaceutical Design, 23(32): 4827-4841

Repeated removal of developing limb buds permanently reduces appendage size in the highly-regenerative axolotl
Bryant, D. M., Sousounis, K., Farkas, J. E., Bryant, S., Thao, N., Guzikowski, A. R., Monaghan, J. R., Levin, M., and Whited, J. L.
Developmental Biology, 424(1): 1-9 [cover]

Serotonergic stimulation induces nerve growth and promotes visual learning via posterior eye grafts in a vertebrate model of induced sensory plasticity
Blackiston, D.J., Vien, K., and Levin, M.
NPJ Regenerative Medicine, 2: 8

Reversals of bodies, brains, and behavior: quantitative analysis of laterality and its disturbance in model species,
Blackiston, D. J., and Levin, M.
in L. J. Rogers and G. Vallortigara (Eds.), Lateralized Brain Functions: Methods in Human and Non-human Species, “Neuromethods”, W. Walz (Ed.), 122: 667-694

Modeling regenerative processes with Membrane Computing
Garcia-Quismondo, M., Levin, M., and Lobo, D.
Information Sciences, 381: 229-2497

Discovering novel phenotypes with automatically inferred dynamic models: a partial melanocyte conversion in Xenopus
Lobo, D., Lobikin, M., and Levin, M.
Scientific Reports, 7: 41339

A Matter of Time: Small RNAs Regulate the Duration of Epigenetic Inheritance
Houri-Zeevi, L., and Rechavi, O.
Trend in Genetics, 33(1): 46-57

2016

Gene-Diet Interactions in Age-Related Macular Degeneration
Rowan, S., and Taylor, A.
Advances in Experimental Medicine and Biology, 854: 95-101

Introduction to provocative questions in left-right asymmetry
Levin, M., Klar, A. J., and Ramsdell, A. F.
Philosophical Transactions of the Royal Society B: Biological Sciences, 371(1710): 20150399

From cytoskeletal dynamics to organ asymmetry: a nonlinear, regulative pathway underlies left-right patterning
McDowell, G., Rajadurai, S., and Levin, M.
Philosophical Transactions of the Royal Society B: Biological Sciences, 371(1710): 20150409 [cover]

Unfolded-protein response-associated stabilization of p27(Cdkn1b) interferes with lens fiber cell denucleation, leading to cataract
Lyu, L., Whitcomb, E. A., Jiang, S., Chang, M. L., Gu, Y., Duncan, M. K., Cvekl, A., Wang, W. L., Limi, S., Reneker, L. W., Shang, F., Du, L., and Taylor, A.
The FASEB Journal, 30(3): 1087-1095

Top-down models in biology: explanation and control of complex living systems above the molecular level
Pezzulo, G., and Levin, M.
Journal of the Royal Society Interface, 13(124): 20160555

Hospital Phenotypes in the Management of Patients Admitted for Acute Myocardial Infarction
Xu, X., Li, S. X., Lin, H., Normand, S. L., Lagu, T., Desai, N., Duan, M., Kroch, E. A., and Krumholz, H. M.
Medical Care, 54(10): 929-936

Vertically- and horizontally-transmitted memories – the fading boundaries between regeneration and inheritance in planaria
Neuhof, M., Levin, M., and Rechavi, O.
Biology Open, 5(9): 1177-1188 [cover]

Physiological inputs regulate species-specific anatomy during embryogenesis and regeneration
Sullivan, K. G., Emmons-Bell, M., and Levin, M.
Communicative & Integrative Biology, 29(4): e1192733 [cover]

Exploring Instructive Physiological Signaling with the Bioelectric Tissue Simulation Engine
Pietak, A., and Levin, M.
Frontiers in Bioengineering and Biotechnology, 4: 55

A computational model of planarian regeneration
De. A., Chakravarthy, V. S., and Levin, M.
International Journal of Parallel, Emergent and Distributed Systems, 32(4): 331-347

Physiological controls of large-scale patterning in planarian regeneration: a molecular and computational perspective on growth and form
Durant, F., Lobo, D., Hammelman, J., and Levin, M.
Regeneration (Oxf), 3(2): 78-102 [cover]