Publications

Browse peer-reviewed publications by topic

All Peer-reviewed Publications

2025

Thoughts and thinkers: On the complementarity between objects and processes
Fields, C., Levin, M. (2025)
Physics of Life Reviews, 2025, doi: 10.1016/j.plrev.2025.01.008.

2024

Open problems in synthetic multicellularity
Solé, R., Conde–Pueyo, N., Pla–Mauri, J., Garcia–Ojalvo, J., Montserrat, N., Levin, M. (2024)
npj Systems Biology and Applications 10, 151 (2024) doi: 10.1038/s41540-024-00477-8

Discussions of machine versus living intelligence need more clarity
Rouleau, N., Levin, M. (2024) 
Nature Machine Intelligence, 2024, doi: 10.31219/osf.io/gz3km

The Multiscale Wisdom of the Body: Collective Intelligence as a Tractable Interface for Next-Generation Biomedicine
Levin, M. (2024) 
BioEssays 2024, doi:10.1002/bies.202400196

Multicellular adaptation to electrophysiological perturbations analyzed by deterministic and stochastic bioelectrical models
Cervera, J., Levin, M., Mafe, S. (2024) 
Sci Rep 14, 27608 (2024) doi:10.1038/s41598-024-79087-7

“Bioelectricity in Development, Regeneration, and Cancers” Cell Bio 2023: A Joint Meeting of the American Society of Cell Biology and European Molecular Biology Organization
Pai, V., Zhang, G., Levin, M. (2024) 
Bioelectricity 2024 6:1, 65-68 doi: 10.1089/bioe.2024.0006

Potential Shortcomings of Genomic Database: The Case of Nav1.5 Expression in Breast Cancer
Djamgoz, M.B.A., Levin, M. (2024)
Bioelectricity 2024, doi: 10.1089/bioe.2024.0033

Natural Induction: Spontaneous Adaptive Organisation without Natural Selection
Buckley, C.L.,  Lewens, T., Levin, M.,  Millidge, B.,  Tschantz, A., Watson, R.A. (2024)
Entropy 2024, 26, 765, doi: 10.3390/e26090765

Disgust as a primary emotional system and its clinical relevance
Tolchinsky, A., Ellis, G. F. R., Levin, M., Kaňková, Š., Burgdorf, J. S. (2024)
Frontiers in Psychology 2024, 15:1454774 doi: 10.3389/fpsyg.2024.1454774

Classical sorting algorithms as a model of morphogenesis: Self-sorting arrays reveal unexpected competencies in a minimal model of basal intelligence
Zhang, T., Goldstein, A., Levin M. (2024)
Adaptive Behavior 2024, doi: 10.1177/10597123241269740

Oscillatory phenomena in electrophysiological networks: The coupling between cell bioelectricity and transcription
Cervera, J., Manzanares, J. A., Levin, M., Mafe, S.  (2024)
Computers in Biology and Medicine 2024, 180 doi: 10.1016/j.compbiomed.2024.108964

Stress sharing as cognitive glue for collective intelligences: A computational model of stress as a coordinator for morphogenesis
Shreesha, L., Levin, M. (2024)
Biochemical and Biophysical Research Communications 2024, 731 doi: 10.1016/j.bbrc.2024.150396

Toward a nomenclature consensus for diverse intelligent systems: Call for collaboration
Kagan, B. J., Mahlis, M., Bhat, A., Bongard, J.,  Cole, V. M., Corlett, P., Gyngell,  C., Hartung, T.,  Jupp, B., Levin, M., Lysaght, T.,   Opie, N., Razi, A., Smirnova, L., Tennant,  Ian, Wade, P. T., Wang, G. (2024)
The Innovation 2024, 5(9) doi:10.1016/j.xinn.2024.100658

Evolutionary Implications of Self-Assembling Cybernetic Materials with Collective Problem-Solving Intelligence at Multiple Scales
Hartl, B., Risi, S., Levin, M. (2024)
Entropy 2024, 26(7), 532 doi:10.3390/e26070532

Self-Amputating and Interfusing Machines
Yang, B., Nasab, A. M., Woodman, S.J., Thomas, E., Tilton, L.G., Levin, M., Kramer-Bottiglio, R. (2024)
Advanced Materials 2024, 2400241, doi:10.1002/adma.202400241

Self-Improvising Memory: A Perspective on Memories as Agential, Dynamically Reinterpreting Cognitive Glue
Levin, M. (2024)
Entropy, 26(6), 481; doi: 10.3390/e26060481

A pro-reparative bioelectronic device for controlled delivery of ions and biomolecules
Asefifeyzabadi, N., Nguyen, T., Li, H., Zhu, K., Yang , H., Baniya, P., PhD, Lopez, A. M., Gallegos, A., Hsieh, H.  Dechiraju, H., Hernandez, C., Schorger, K., Recendez, C.,  Tebyani, M., Selberg, J., Luo, L., Muzzy, E., Hsieh, C., Barbee, A., Orozco, J.,  Alhamo, M.A., Levin, M., Aslankoohi, E., Gomez, M.,  Zhao, M., Teodorescu, M.,  Isseroff, R. R., Rolandi, M. (2024)
Wound Rep Reg, 1-11 (2024) doi:10.1111/wrr.13191 

Collective intelligence: A unifying concept for integrating biology across scales and substrates
McMillen, P., Levin, M. (2024)
Communications Biology, 2024, 7, 378 doi:10.1038/s42003-024-06037-4

Unraveling the Enigma of Organismal Death: Insights, Implications, and Unexplored Frontiers
Noble, P. A., Pozhitkov, A., Singh, K., Woods, E.,  Liu, C., Levin, M., Javan, G., Wan, J., Abouhashem, A. S., Mathew-Steiner, S. S., Sen, C. K. (2024)
ACS Physiology, 2024  doi:10.1152/physiol.00004.2024 

Snakes and ladders in the evolution of language
Field, C., and Levin, M. (2024)
Physics of Life Reviews, 2024 doi:10.1016/j.plrev.2024.04.007

Aging as a loss of morphostatic information: a developmental bioelectricity perspective
Pio-Lopez, L., Levin, M. (2024)
Ageing Research Reviews, 2024, 97:102310, doi:10.1016/j.arr.2024.102310

Programmable delivery of fluoxetine via wearable bioelectronics for wound healing in vivo
Li, H., Yang, H.-Y., Asefifeyzabadi, N., Baniya, P., Medina Lopez, A., Gallegos, A., Zhu, K., Hsieh, H.-C., Nguyen, T., Hernandez, C., Zlobina, K., Recendez, C., Tebyani, M., Carrión, H., Selberg, J., Luo, L., Alhamo, M. A., Soulika, A. M., Levin, M., Norouzi, N., Gomez, M., Zhao, M., Teodorescu, M., Isseroff, R. R., Rolandi, M. (2024)
Advanced Materials Technology, 9(7):2301115

Electroceuticals: emerging applications beyond the nervous system and excitable tissues
Balasubramanian, S., Weston, D., Levin, M., Davidian, D. C. C. (2024)
Trends Pharmacol Sci, 2109 doi.org/10.1016/j.tips.2024.03.001

Charging Ahead: Examining the future therapeutic potential of electroceuticals
Balasubramanian, S., Weston, D., Levin, M., Davidian, D. C. C. (2024)
Advanced Therapeutics, in press

Principled Limitations on Self-Representation for Generic Physical Systems
Fields, C., Glazebrook, J. F., and Levin, M. (2024)
Entropy, 26(3): 194

AI-driven Automated Discovery Tools Reveal Diverse Behavioral Competencies of Biological Networks
Etcheverry, M., Moulin-Frier, C., Oudeyer, P.-Y., and Levin, M. (2024)
eLife, 13: RP92683

Embryos assist morphogenesis of others through calcium and ATP signaling mechanisms in collective teratogen resistance 
Tung, A., Sperry, M., Clawson, W., Pavuluri, A., Bulatao, S., Yue, M., Flores, R. M., Pai, V. McMillen, P., Kuchling, F., and Levin, M. (2024) 
Nature Communications, 15(1): 535

Machine Learning for Hypothesis Generation in Biology and Medicine: Exploring the latent space of neuroscience and developmental bioelectricity 
O’Brien, T., Stremmel, J., Pio-Lopez, L., McMillen, P., Rasmussen-Ivey, C., and Levin, M. (2024) 
Digital Discovery, 3(2): 249-263

Optical Estimation of Bioelectric Patterns in Living Embryos  
McMillen, P., and Levin, M. (2024)  
in M. Bizzarri (Ed.), Systems Biology: Methods in Molecular Biology, 2745: 91-102, doi:10.1007/978-1-0716-3577-3_6

From reinforcement learning to agency: Frameworks for understanding basal cognition  
Seifert, G., Sealander, A., Marzen, S., and Levin, M. (2024)  
BioSystems, 235(1): 105107

2023

Nested Selves: Self-Organization and Shared Markov Blankets in Prenatal Development in Humans  
Ciaunica, A., Levin, M., Rosas, F. E., and Friston, K. (2023)  
Topics in Cognitive Science, in press, doi:10.1111/tops.12717

Motile Living Biobots Self-Construct from Adult Human Somatic Progenitor Seed Cells  
Gumuskaya, G., Srivastava, P., Cooper, B. G., Lesser, H., Semegran, B., Garnier, S., and Levin, M. (2023)  
Advanced Science, doi:10.1002/advs.202303575

Information integration during bioelectric regulation of morphogenesis of the embryonic frog brain  
Manicka, S., Pai, V. P, and Levin, M. (2023)  
iScience, 26(12): 108398

The Multiple Realizability of Sentience in Living Systems and Beyond  
Rouleau, N., and Levin, M. (2023)  
eNeuro, 10(11), doi:10.1523/eneuro.0375-23.2023

Long Range Communication via Gap Junctions and Stress in Planarian Morphogenesis: A Computational Study  
Blattner, M., and Levin, M. (2023)  
Bioelectricity, 5(3): 196-209

Cancer’s unique bioelectric properties: from cells to body-wide networks, Comment on: “The distinguishing electrical properties of cancer cells” by Elisabetta Di Gregorio, Simone Israel, Michael Staelens, Gabriella Tankel, Karthik Shankar, and Jack A. Tuszynski  
Mathews, J., and Levin, M. (2023)  
Physics of Life Reviews, 47: 113-115

Optimization of Oligomer Stamping Technique for Normally Closed Elastomeric Valves on Glass Substrate  
Dungan, J., Mathews, J., Levin, M., and Koomson, V. (2023)  
Micromachines, 14(9): 1659

Collective Intelligence of Morphogenesis as a Teleonomic Process  
Levin, M. (2023)  
in Corning, P. A., Kauffman, S. A., Noble, D., Shapiro, J. A., Vane-Wright, R. I. Pross, A. (Eds.), Evolution "On Purpose": Teleonomy in Living Systems, The MIT Press: Cambridge, MA, 175-197

Endless forms most beautiful 2.0: teleonomy and the bioengineering of chimaeric and synthetic organisms  
Clawson, W. P., and Levin, M. (2023)  
Biological Journal of the Linnean Society, 139(4): 457-486 [cover]

Closing the Loop on Morphogenesis: A Mathematical Model of Morphogenesis by Closed-Loop Reaction-Diffusion  
Grodstein, J., McMillen, P., and Levin, M. (2023)  
Frontiers in Cell and Developmental Biology, 11: 1087650

Correcting instructive electric potential patterns in multicellular systems: External actions and endogenous processes  
Cervera, J., Levin, M., and Mafé, S. (2023)  
Biochimica et Biophysica Acta (BBA) - General Subjects, 1867(10): 130440

Searching in the Dark: Evolving Biobot Swarm Compositions to Efficiently Explore Obstructed Environments  
Welch, P., Grasso, C., Gumuskaya, G., Levin, M., and Bongard, J. (2023)  
Proceedings of ALIFE 2023: Ghost in the Machine, article 70, doi:10.1162/isal_a_00683

Future Medicine: from molecular pathways to the collective intelligence of the body  
Lagasse, E., and Levin, M. (2023)  
Trends in Molecular Medicine, 29(9): 687-710 [cover]

Multiple ways to implement and infer sentience: Commentary on Segundo-Ortiz & Calvo on Plant Sentience  
Rouleau, N., and Levin, M. (2023)  
Animal Sentience, 8(33): 30

The Yin and Yang of Breast Cancer: Ion Channels as Determinants of Left-Right Functional Differences  
Masuelli, S., Real, S., McMillen, P., Oudin, M., Levin, M., and Roqué, M. (2023)  
International Journal of Molecular Sciences, 24(13): 11121

Towards an Ethics of Autopoietic Technology: Stress, Care, and Intelligence  
Witkowski, O., Doctor, T., Solomonova, E., Duane, B., and Levin, M. (2023)  
Biosystems, 231: 104964

Bioelectricidad en agregados multicelulares de células no excitables: modelos biofísicos  
Cervera, J., Levin, M., and Mafé, S. (2023)  
Revista Española de Física, 37(2), 24-29

The Brain is not Mental! Coupling Neuronal and Immune Cellular Processing in Human Organisms  
Ciaunica, A., Shmeleva, E. V., and Levin, M. (2023)  
Frontiers in Integrative Neuroscience, 17: 1057622

Regulative development as a model for origin of life and artificial life studies  
Fields, C., and Levin, M. (2023)  
Biosystems, 229: 104927

The collective intelligence of evolution and development  
Watson, R., and Levin, M. (2023)  
Collective Intelligence, 2(2), doi:10.1177/26339137231168355

Bioelectric Networks: the cognitive glue enabling evolutionary scaling from physiology to mind  
Levin, M. (2023)  
Animal Cognition, doi:10.1007/s10071-023-01780-3

Control flow in active inference systems Part II: Tensor networks as general models of control flow  
Fields, C., Fabrocini, F., Friston, K., Glazebrook, J. F., Hazan, H., Levin, M., and Marcianò, A. (2023)  
IEEE Transactions on Molecular, Biological, and Multi-Scale Communications, doi:10.1109/TMBMC.2023.3272158

Control flow in active inference systems Part I: Classical and quantum formulations of active inference  
Fields, C., Fabrocini, F., Friston, K., Glazebrook, J. F., Hazan, H., Levin, M., and Marcianò, A. (2023)  
IEEE Transactions on Molecular, Biological, and Multi-Scale Communications, doi:10.1109/TMBMC.2023.3272150

Darwin’s agential materials: evolutionary implications of multiscale competency in developmental biology  
Levin, M. (2023)  
Cellular and Molecular Life Sciences, 80: 142

Morphoceuticals: Perspectives for discovery of drugs targeting anatomical control mechanisms in regenerative medicine, cancer and aging  
Pio-Lopez, L., and Levin, M. (2023)  
Drug Discovery Today, 28(6): 103585

Cellular Signaling Pathways as Plastic, Proto-cognitive Systems: Implications for Biomedicine  
Mathews, J., Chang, J., Devlin, L., and Levin, M. (2023)  
Patterns, 4: 100737

Biological Robots: Perspectives on an Emerging Interdisciplinary Field  
Blackiston, D., Kriegman, S., Bongard, J., and Levin, M. (2023)  
Soft Robotics, 10(4): 694-686

The scaling of goals via homeostasis: an evolutionary simulation, experiment and analysis  
Pio-Lopez, L., Bischof, J., LaPalme, J. V., and Levin, M. (2023)  
Interface Focus, 13(3): 20220072 [cover]

The nonlinearity of regulation in biological networks  
Manicka, S., Johnson, K., Levin, M., and Murrugarra, D. (2023)  
npj Systems Biology and Applications, 9(1): 10

There’s Plenty of Room Right Here: Biological Systems as Evolved, Overloaded, Multi-Scale Machines  
Bongard, J., and Levin, M. (2023)  
Biomimetics, 8(1): 110

Synthetic morphology with agential materials  
Davies, J., and Levin, M. (2023)  
Nature Reviews Bioengineering, 1(1): 46-59

Cellular Competency during Development Alters Evolutionary Dynamics in an Artificial Embryogeny Model  
Shreesha, L., and Levin, M. (2023)  
Entropy, 25(1): 131

Bioelectricity of non-excitable cells and multicellular pattern memories: Biophysical modeling  
Cervera, J., Levin, M., and Mafe, S. (2023)  
Physics Reports, 1004: 1-31

Learning in Transcriptional Network Models: Computational Discovery of Pathway-level Memory and Effective Interventions  
Biswas, S., Clawson, W., and Levin, M. (2023)  
International Journal of Molecular Sciences, 24(1): 285

Transplantation of fragments from different planaria: a bioelectrical model for head regeneration  
Cervera, J., Manzanares, J. A., Levin, M.,  and Mafe, S. (2023)  
Journal of Theoretical Biology, 558: 111356

2022

Exploring the Behavior of Bioelectric Circuits Using Evolution Heuristic Search  
Hazan, H., and Levin, M. (2022)  
Bioelectricity, 4(4): 207-227

On-chip on-demand delivery of K+ for in vitro bioelectronics  
Dechiraju, H., Selberg, J., Jia, M., Pansodtee, P., Li, H., Hsieh, H-C., Hernandez, C.,  Asefifeyzabadi, N., Nguyen, T., Baniya, P., Marquez, G., Rasmussen-Ivey, C., Bradley, C., Teodorescu, M., Gomez, M., Levin, M., and Rolandi, M. (2022)  
AIP Advances, 12(12): 125205

Active Inference, Morphogenesis, and Computational Psychiatry  
Pio-Lopez, L., Kuchling, F., Tung, A., Pezzulo, G., and Levin, M. (2022)  
Frontiers in Computational Neuroscience, 16: 988977

The Free Energy Principle induces neuromorphic development  
Fields, C., Friston, K., Glazebrook, J. F., Levin, M., and Marcianò, A. (2022)  
Neuromorphic Computing and Engineering, 2: 042002

Information Theory as an experimental tool for integrating disparate biophysical signaling modules  
McMillen, P., Walker, S. I., and Levin, M. (2022)  
International Journal of Molecular Sciences, 23(17): 9580

Adversarial Takeover of Neural Cellular Automata  
Cavuoti, L., Sacco, F., Randazzo, E., and Levin, M. (2022)  
Proceedings of the 2022 Conference on Artificial Life (ALIFE 2022),  pp. 256-263

Competition for Finite Resources as Coordination Mechanism for Morphogenesis: an evolutionary algorithm study of digital embryogeny  
Smiley, P., and Levin, M. (2022)  
BioSystems, 221: 104672

Generalizing frameworks for sentience beyond natural species  
Levin, M. (2022)  
Animal Sentience, 7(32): 15

Screening Biophysical Sensors and Neurite Outgrowth Actuators in Human Induced-Pluripotent-Stem-Cell-Derived Neurons  
Pai, V. P., Cooper, B. G., and Levin, M. (2022)  
Cells, 11(16): 2470

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., & Ingber, D. E. (2022)  
Advanced Science, 9(26): 2200222 [cover]

HCN2 Channel-induced Rescue of Brain, Eye, Heart, and Gut Teratogenesis Caused by Nicotine, Ethanol, and Aberrant Notch Signaling  
Pai, V. P., and Levin, M. (2022)  
Wound Repair and Regeneration, 30(6): 6810706

Biology, Buddhism, and AI: Care as the Driver of Intelligence  
Doctor, T., Witkowski, O., Solomonova, E., Duane, B., and Levin, M. (2022)  
Entropy, 24(5): e24050710

Competency in Navigating Arbitrary Spaces as an Invariant for Analyzing Cognition in Diverse Embodiments  
Fields, C., and Levin, M. (2022)  
Entropy, 24(6): 819

Neurons as hierarchies of quantum reference frames  
Fields, C., Glazebrook, J. F., and Levin, M. (2022)  
Biosystems, 219: 104714

A free energy principle for generic quantum systems   
Fields, C., Friston, K., Glazebrook, J. F., and Levin, M. (2022)  
Progress in Biophysics and Molecular Biology, 173: 36-59

Metacognition as a Consequence of Competing Evolutionary Time Scales  
Kuchling, F., Fields, C., and Levin, M. (2022)  
Entropy, 24(5): 601

Design for an Individual: Connectionist approaches to the evolutionary transitions in individuality  
Watson, R. A., Levin, M., and Buckley, C. L. (2022)  
Frontiers in Ecology and Evolution, 10: 823588

Technological Approach to Mind Everywhere: an experimentally-grounded framework for understanding diverse bodies and minds  
Levin, M. (2022)  
Frontiers in Systems Neuroscience, 16: 768201

Biological underpinnings for lifelong learning machines  
Kudithipudi, D., Aguilar-Simon, M., Babb, J., Bazhenov, M., Blackiston, D., Bongard, J., Brna, A. P., Raja, S. C., Cheney, N., Clune, J., Daram, A., Fusi, S., Helfer, P., Kay, L., Ketz, N., Kira, Z., Kolouri, S., Krichmar, J. L., Kriegman, S., Levin, M., Madireddy, S., Manicka, S., Marjaninejad, A., McNaughton, B., Miikkulainen, R., Navratilova, Z., Pandit, T., Parker, A., Pilly, P. K., Risi, S., Sejnowski, T. J., Soltoggio, A., Soures, N., Tolias, A. S., Urbina-Meléndez, D., Valero-Cuevas, F. J., van de Ven, G. M., Vogelstein, J. T., Wang, F., Weiss, R., Yanguas-Gil, A., Zou, X., and Siegelmann, H. (2022)  
Nature Machine Intelligence, 4(3): 196-210

Studying Protista WBR and Repair Using Physarum polycephalum  
Sperry, M. M., Murugan, N. J., and Levin, M. (2022)  
in S. Blanchoud and B. Galliot (Eds.), Whole-Body Regeneration: Methods in Molecular Biology, 2450: 51-67

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. (2022)  
Cancers, 14(6): 1499

Modular cognition  
Levin, M., and Yuste, R. (2022)  
Aeon Essays [cover]

Multi-scale Chimerism: an experimental window on the algorithms of anatomical control  
Nanos, V., and Levin, M. (2022)  
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. (2022)  
Bioelectricity, 4(1): 18-30

Impact of Membrane Voltage on Formation and Stability of Human Renal Tubules in Vitro  
Adelfio, M., Bonzanni, M., Levin, M., and Kaplan, D. L. (2022)  
ACS Biomaterials Science & Engineering, 8(3): 1239-1246

When left does not seem right: epigenetic and bioelectric differences between left- and right-sided breast cancer  
Masuelli, S., Sebastian, R., Campoy, E., Branham, M. T., Marzese, D. M., Matthew, S., DeBlas, G., Rodolfo, A., Levin, M., and Roque, M. (2022)  
Molecular Medicine, 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. (2022)  
Science Advances, 8(4): abj2164 [cover]

Minimal developmental computation: a causal network approach to understand morphogenetic pattern formation  
Manicka, S., and Levin, M. (2022)  
Entropy, 24: 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. (2022)  
EBioMedicine, 75: 103767

2021

Morphology changes induced by intercellular gap junction blocking: a reaction-diffusion mechanism  
Cervera, J., Levin, M., and Mafe, S. (2021)  
BioSystems, 209: 104511

Evolution and emergence: higher order information structure in protein interactomes across the tree of life  
Klein, B., Hoel, E., Swain, A., Griebenow, R., and Levin, M. (2021)  
Integrative Biology, 13(12): 283-294 [cover]

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

Kinematic self-replication in reconfigurable organisms  
Kriegman, S., Blackiston, D., Levin, M., and Bongard, J. (2021)  
Proceedings of the National Academy of Science, 118(49): e2112672118

Behaviorist Approaches to Investigating Memory and Learning: A Primer for Synthetic Biology and Bioengineering  
Abramson, C., and Levin, M. (2021)  
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. (2021)  
Cancers, 13(21): 5300

Metabolic limits on classical information processing by biological cells  
Fields, C., and Levin, M. (2021)  
BioSystems, 209: 104513

Beyond Neurons: Long Distance Communication in Development and Cancer  
McMillen, P., Oudin, M. J, Levin, M., and Payne, S. L. (2021)  
Frontiers in Cell and Developmental Biology, 9: 739024

Stability and robustness properties of bioelectric networks: A computational approach  
Grodstein, J., and Levin, M. (2021)  
Biophysics Review, 2(3): 031305

Minimal physicalism as a scale-free substrate for cognition and consciousness  
Fields, C., Glazebrook, J. F., and Levin, M. (2021)  
Neuroscience of Consciousness, 7(2): niab013

Mechanosensation Mediates Long-Range Spatial Decision-Making in an Aneural Organism  
Murugan, N., Kaltman, D., Jin, H., Chien, M., Flores, R., Nguyen, C., Tuzoff, D., Minabutdinov, A., Kane, A., Novak, R., Ingber, D., and Levin, M. (2021)  
Advanced Materials, 33(34): 2008161 [cover]

Life, death, and self: Fundamental questions of primitive cognition viewed through the lens of body plasticity and synthetic organisms  
Levin, M. (2021)  
Biochemical and Biophysical Research Communications, 564: 114-133 [cover]

Scale invariant robot behavior with fractals  
Kriegman, S., Nasab, A-M., Blackiston, D., Steele, H., Levin, M., Kramer-Bottiglio, R., and Bongard, R. (2021)  
Robotics: Science and Systems (RSS), doi:10.15607/RSS.2021.XVII.059

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. (2021)  
Nanophotonics, 10(9): 2441-2450

Synthetic living machines: A new window on life  
Ebrahimkhani, M., and Levin, M. (2021)  
iScience, 24(5): 102505

Adversarial Reprogramming of Neural Cellular Automata: a robustness investigation  
Randazzo, E., Mordvintsev, A., Niklasson, E., and Levin, M. (2021)  
Distill, 6(5), doi:10.23915/distill.00027.004

Bioelectrical approaches to cancer as a problem of the scaling of the cellular self  
Levin, M. (2021)  
Progress in Biophysics and Molecular Biology, 165: 102-113

Learning and synaptic plasticity in 3D bioengineered neural tissues  
Rouleau, N., Cairns, D. M., Rusk, W., Levin, M., and Kaplan, D. (2021)  
Neuroscience Letters, 750: 135799

Bioelectric Signaling: Reprogrammable Circuits Underlying Embryogenesis, Regeneration, and Cancer  
Levin, M. (2021)  
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. (2021)  
Philosophical Proceedings of the Royal Society B, 376(1821): 20190765

Uncovering cognitive similarities and differences, conservation and innovation  
Levin, M., Keijzer, F., Lyon, P., and Arend, D. (2021)  
Philosophical Transactions of the Royal Society B, 376 (1821): 20200458 [cover]

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. (2021)  
iScience, 24(3): 102131

A cellular platform for the development of synthetic living machines  
Blackiston, D., Lederer, E., Kriegman, S., Garnier, S., Bongard, J., and Levin, M. (2021)  
Science Robotics, 6(52): eabf1571

Living things are not (20th Century) machines: updating mechanism metaphors in light of the modern science of machine behavior  
Bongard, J., and Levin, M. (2021)  
Frontiers in Ecology and Evolution, 9: 650726

Unlimited plasticity of embodied, cognitive subjects: a new playground for the UAL framework  
Levin, M. (2021)  
Biology and Philosophy, 36(2): 17

Self-organising Textures: Neural Cellular Automata Model of Pattern Formation  
Niklasson, E., Mordvintsev, A., Randazzo, E., and Levin, M. (2021)  
Distill, 6(2), doi:10.23915/distill.00027.003

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, F., Martins de Almeida F., Martinez, A. M. B., Abreu, J. G., Levin, M., and Carneiro, K. (2021)  
Developmental & Comparative Immunology, 114: 103840

Rewiring Endogenous Bioelectric Circuits in the Xenopus laevis Embryo Model  
Nanos, V., and Levin, M. (2021)  
in M. R. Ebrahimkhani and J. Hislop (Eds.), Programmed Morphogenesis: Methods in Molecular Biology, 2258: 93-103

Reframing cognition: getting down to biological basics  
Lyon, P., Keijzer, F., Arendt, D., and Levin, M. (2021)  
Philosophical Transactions of the Royal Society B, 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. (2021)  
Advanced Materials, 33(19): 2002882

Inducing Vertebrate Limb Regeneration: A Review of Past Advances and Future Outlook  
Davidian, D., and Levin, M. (2021)  
Cold Spring Harbor Perspectives in Biology, doi:10.1101/cshperspect.a040782

A Meta-Analysis of Bioelectric Data in Cancer, Embryogenesis, and Regeneration  
Srivastava, P., Kane, A., Harrison, C., and Levin, M. (2021)  
Bioelectricity, 3(1): 42-67

A Comprehensive Conceptual and Computational Dynamics Framework for Autonomous Regeneration Systems  
Minh-Thai, T. N., Samarasinghe, S., and Levin, M. (2021)  
Artificial Life, 27(2): 80-104

2020

Why isn't sex optional? Stem-cell competition, loss of regenerative capacity, and cancer in metazoan evolution  
Fields, C., and Levin, M. (2020)  
Communicative & Integrative Biology, 13(1): 170-183

How Do Living Systems Create Meaning?  
Fields, C., and Levin, M. (2020)  
Philosophies, 5(4): 36-59

Community effects allow bioelectrical reprogramming of cell membrane potentials in multicellular aggregates: Model simulations  
Cervera, J., Ramirez, P., Levin, M., and Mafe, S. (2020)  
Physical Review E, 102(5): 052412

Nervous System and Tissue Polarity Dynamically Adapt to New Morphologies in Planaria  
Bischof, J., Day, M., Miller, K., LaPalme, J., and Levin, M. (2020)  
Developmental Biology, 467(1-2): 51-65 [cover]

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. (2020)  
Mechanisms of Development, 163: 103614 [cover]

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. (2020)  
APL Materials, 8(9): 091106

How Groups of Cells Cooperate to Build Organs and Organisms  
Levin, M. (2020)  
The Scientist, 34(9): 38-45 [cover]

Self-classifying MNIST Digits: Achieving Distributed Coordination with Neural Cellular Automata  
Randazzo, E., Mordvintsev, A., Niklasson, E., Levin, M., and Greydanus, S. (2020)  
Distill, 5(8), doi:10.23915/distill.00027.002

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., Sievert, K., Ramirez, C., Rusk, W., Levin, M., and Kaplan, D. (2020)  
Biomolecules, 10(8): 1196

Emergence of Informative Higher Scales in Biological Systems: a computational toolkit for optimal prediction and control  
Hoel, E., and Levin, M.,(2020)  
Communicative & Integrative Biology, 13(1): 108-118

Growing Neural Cellular Automata  
Mordvintsev, A., Randazzo, E., Niklasson, E., and Levin, M. (2020)  
Distill, 5(2), doi:10.23915/distill.00027.002

Applications and ethics of computer-designed organisms  
Levin, M., Bongard, J., and Lunshof, J. E. (2020)  
Nature Reviews Molecular Cell Biology, 21(11): 655–656

Scale-Free Biology: Integrating Evolutionary and Developmental Thinking  
Fields, C., and Levin, M. (2020)  
BioEssays, 42(8): 1900228

Competitive and Coordinative Interactions between Body Parts Produce Adaptive Developmental Outcomes  
Gawne, R., McKenna, K., and Levin, M. (2020)  
BioEssays, 42(8): 1900245 [cover]

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. (2020)  
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. (2020)  
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. (2020)  
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. (2020)  
Journal of the Royal Society Interface, 17(167): 20200177

Richard Borgens, 1946-2019  
Levin, M., and Rajnicek, A. M. (2020)  
Bioelectricity, 2(2): 205

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. (2020)   
Frontiers in Cellular Neuroscience, 14: 136

Scalable sim-to-real transfer of soft robot designs  
Kriegman, S., Nasab, A. M., Shah, D., Steele, H., Branin, G., Levin, M., Bongard, J., and Kramer-Bottiglio, R. (2020)  
Proceedings of the 3rd IEEE International Conference on Soft Robotics (RoboSoft 2020), p. 359-366

Bioelectrical Coupling of Single-Cell States in Multicellular Systems  
Cervera, J., Levin, M., and Mafe, S. (2020),   
The Journal of Physical Chemistry Letters, 11(9): 3234-3241

Extra-genomic instructive influences in morphogenesis: A review of external signals that regulate growth and form  
Tung, A., and Levin, M. (2020)  
Developmental Biology, 461(1): 1-12 [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. (2020)  
International Journal of Developmental Biology, 64(10-11-12): 485-494

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. (2020)  
Advanced Intelligent Systems, 2(12): 2000140 [cover]

Does evolution have a target morphology?  
Fields, C., and Levin, M. (2020)  
Organisms. Journal of Biological Sciences, 4(1): 57-76

Interferon-Gamma Stimulated Murine Macrophages in Vitro: Impact of Ionic Composition and Osmolarity and Therapeutic Implications  
Erndt-Marino, J., Yeisley, D. J., Chen, H., Levin, M., Kaplan, D. L., and Hahn, M. S. (2020)  
Bioelectricity, 2(1): 48-58

Opinion: Use the Pandemic to Expand the Lab to the Home  
Levin, M. (2020)  
The Scientist

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. (2020)  
Scientific Reports, 10: 4512

Revisiting Burr and Northrop's 'The Electro-Dynamic Theory of Life'  
Levin, M. (2020)  
Biological Theory, 15: 83-90

Eya2 promotes cell cycle progression by regulating DNA damage response during vertebrate limb regeneration  
Sousounis, K., Bryant, D. M., Fernandez, J. M., Eddy, S. S., Tsai, S. L., Gundberg, G. C., Han, J., Courtemanche, K., Levin, M., and Whited, J. L. (2020)  
eLIFE, 9: e51217

Does regeneration recapitulate phylogeny?  
Fields, C., and Levin, M. (2020)  
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. (2020)  
npj Regenerative Medicine, 5: 2

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. (2020)  
Journal of Molecular Biology, 432(2): 605-620 [cover]

Formin, an opinion  
Davison, A., McDowell, G. S., Holden, J. M., Johnson, H. F., Wade, C. M., Chiba, S., Jackson, D. J., Levin, M., and Blaxter, M. L. (2020)  
Development, 147: dev187427

Optogenetically induced cellular habituation in non-neuronal cells  
Bonzanni, M., Rouleau, N., Levin, M., and Kaplan, D. L. (2020)  
PLOS One, 15(1): e0227230

A Scalable Pipeline for Designing Reconfigurable Organisms  
Kriegman, S., Blackiston, D., Levin, M., and Bongard, J. (2020)   
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. (2020)  
Bioelectricity, 2(1): 21-32

The Biophysics of Regenerative Repair Suggests New Perspectives on Biological Causation  
Levin, M. (2020)  
BioEssays, 42(2): 1900146

Bioelectrical model of head-tail patterning based on cell ion channels and intercellular gap junctions  
Cervera, J., Meseguer, S., Levin, M., and Mafe, S. (2020)  
Bioelectrochemistry, 132: 107410

Morphological Coordination: A Common Ancestral Function Unifying Neural and Non-Neural Signaling  
Fields, C., Bischof, J., and Levin, M. (2020)  
Physiology, 35: 16-30

Post-SSRI Sexual Dysfunction: a bioelectric mechanism?  
Healy, D., Faber, C., Lapalme, J., and Levin, M. (2020)  
Bioelectricity, 2(1): 7-13

How to understand cells, tissues and organisms as agents with agendas  
Levin, M., and Dennett, D. C. (2020)  
Aeon Essays

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. (2020)   
Biology Open, 9(1): bio048553

2019

Endogenous Bioelectrics in Development, Cancer, and Regeneration: Drugs and Bioelectronic Devices as Electroceuticals for Regenerative Medicine  
Levin, M., Selberg, J., and Rolandi, M. (2019)  
iScience, 22: 519–533

Modeling somatic computation with non-neural bioelectric networks  
Manicka, S., and Levin, M. (2019)  
Scientific Reports 9: 18612

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

The Computational Boundary of a 'Self': Developmental Bioelectricity Drives Multicellularity and Scale-Free Cognition  
Levin, M. (2019)  
Frontiers in Psychology, 10: 2688

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. (2019)  
iScience, 22: 147-165

L-type voltage-gated Ca2+ channel CaV1.2 regulates chondrogenesis during limb development  
Atsuta, Y., Tomizwa, R. R., Levin, M., and Tabin, C. J. (2019)  
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. (2019)  
ACS Biomaterials Science & Engineering, 5(10): 5327-5336

Managing Ideas, People, and Projects: Organizational Tools and Strategies for Researchers  
Levin, S., and Levin, M. (2019)  
iScience, 20: 278-291

Bioelectric control of metastasis in solid tumors  
Payne, S. L., Levin, M., and Oudin, M. J. (2019)  
Bioelectricity, 1(3): 114-130

Somatic Multicellularity as a Satisficing Solution to the Prediction-Error Minimization Problem  
Fields, C., and Levin, M. (2019)  
Communicative & Integrative Biology, 12(1): 119-132

A flow through device for simultaneous dielectrophoretic cell trapping and AC electroporation  
Punjiya, M., Nehad, H. R., Mathews, J., Levin, M., and Sonkusale, S. (2019)  
Scientific Reports, 9: 11988

Bioelectrical controls of morphogenesis: from ancient mechanisms of cell coordination to biomedical opportunities  
Whited, J., and Levin, M. (2019)  
Current Opinion in Genetics & Development, 57: 61-69 [cover]

Reverse-engineering growth and form in Heidelberg  
Levin, M., and Martinez-Arias, A. (2019)  
Development, 146(14): dev177261

Automated Shapeshifting for Function Recovery in Damaged Robots  
Kriegman, S., Walker, S., Shah, D. S., Kramer-Bottiglio, R., and Bongard, J. (2019)  
Proceedings of Robotics: Science and Systems XV: 28

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. (2019)  
Progress in Biophysics and Molecular Biology, 149: 39-53 [cover]

On the generalization of habituation: how discrete biological systems respond to repetitive stimuli  
Bonzanni, M., Rouleau, N., Levin, M., and Kaplan, D. (2019)  
BioEssays, 41(7): 1900028

Live Imaging of Intracellular pH in Planarians Using the Ratiometric Fluorescent Dye SNARF-5F-AM  
Beane, W. S., Adams, D. S., Morokuma, J., and Levin, M. (2019)  
Biology Methods and Protocols, 4(1): bpz005

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. (2019)  
Journal of Physical Chemistry B, 123(18): 3924-3934 [cover]

The Cognitive Lens: a primer on conceptual tools for analysing information processing in developmental and regenerative morphogenesis  
Manicka, S., and Levin, M. (2019)  
Philosophical Transactions of the Royal Society B, 374: 20180369

Neural control of body-plan axis in regenerating planaria  
Pietak, A., Bischof, J., LaPalme, J., Morokuma, J., and Levin, M. (2019)  
PLOS Computational Biology, 15(4): e1006904

A Call for a Better Understanding of Causation in Cell Biology  
Bizzari, M., Brash, D., Briscoe, J., Grieneisen, V., Stern, C. D., and Levin, M. (2019)  
Nature Reviews Molecular Cell Biology, 20(5): 261-262

Membrane potential depolarization alters calcium flux and phosphate signaling during osteogenic differentiation of human mesenchymal stem cells  
Sundelacruz, S., Moody, A. T., Levin, M., and Kaplan, D. (2019)  
Bioelectricity, 1(1): 56-66

Toward modeling regeneration via adaptable echo state networks  
Hammelman, J., Siegelmann, H., Manicka, S., and Levin, M. (2019)  
in A. Adamatzky, S. Akl, and G. Sirakoulis (Eds.), From parallel to emergent computing, Chapter 6, pages 117-133, CRC Press: Boca Raton, FL

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  
Berard, A., Levin, M., Sadler, T., and Healy, D. (2019)  
Bioelectricity, 1(1): 18-29

Effects of ivermectin exposure on regeneration of D. dorotocephala planaria: exploiting human-approved ion channel drugs as morphoceuticals  
Ferenc, N. N., and Levin, M. (2019)  
Macromolecular Bioscience, 19(3): 1800237 [cover]

Planarian Regeneration as a Model of Anatomical Homeostasis: Recent Progress in Biophysical and Computational Approaches  
Levin, M., Pietak, A., and Bischof, J. (2019),  
Seminars in Cell and Developmental Biology, 87: 125-144

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. (2019)  
Biophysical Journal, 116(5): 948-961

EDEn – Electroceutical Design Environment: Ion Channel Tissue Expression Database with Small Molecule Modulators  
Churchill, C. D. M., Winter, P., Tuszynski, J. A., and Levin, M. (2019)   
iScience, 11: 42-56

2018

Multiscale Memory and Bioelectric Error Correction in the Cytoplasm-Cytoskeleton-Membrane System  
Fields, C., and Levin, M. (2018)  
Wiley Interdisciplinary Reviews: Systems Biology and Medicine, 10(2): e1410 [cover]

A Computational Framework for Autonomous Self-repair Systems  
Minh-Thai, T. N., Aryal, J., Samarasinghe, S., and Levin, M. (2018)  
in Mitrovic, T., Xue, B., and Li, X. (Eds.), AI 2018: Advances in Artificial Intelligence, p153-159. Lecture Notes in Computer Science, vol 11320. Springer, Cham

Ahead of the Curve: Hidden Breakthroughs in the Biosciences, Vol 2  
Adams, D. S., and Levin, M. (2018)  
IOP Publishing Ltd: Bristol, UK [cover]

The bacterial metabolite indole inhibits regeneration of the planarian flatwork Dugesia japonica  
Lee, F. J., Williams, K. B., Levin, M., and Wolfe, B. E. (2018)  
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, K. M., Harrison, C., Martyniuk, C. J., Guay, J. A., Zaltsman, J., Carabello, H., Kaplan, D. L., and Levin, M. (2018)  
Cell Reports, 25(6): 1593-1609

Perspective: the promise of multi-cellular engineered living systems  
Kamm, R. D., Bashir, R., Arora, N., Dar, R. D., Gilette, M. U., Griffith, L. G., Kemp, M. L., Kinlaw, K., Levin, M., Martin, A. C., McDevitt, T. C., Nerem, R. M., Powers, M. J., Saif, T. A., Sharpe, J., Takayama, S., Takeuchi, S., Weiss, R., Ye, K., Yevick, H. G., and Zaman, M. H. (2018)  
APL Bioengineering, 2(4): 040901

Ivermectin promotes peripheral nerve regeneration during wound healing  
Cairns, D. M., Giordano, J. E., Conte, S., Levin, M., and Kaplan, D. L. (2018)  
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. (2018)  
PLoS Genetics, 14(9): 1007642

Cross-limb Communication During Xenopus Hind-limb Regenerative Response: Non-Local Bioelectric Injury Signals  
Busse, S. M., McMillen, P. T., and Levin, M. (2018)  
Development, 145(19): dev164210

Pattern Regeneration in Coupled Networks  
Moore, D. G., Walker, S. I., and Levin, M. (2018)  
in T. Ikegami, N. Virgo, O. Witkowski, M. Oka, R. Suzuki, and H. Iizuka (Eds.), ALIFE 2018: The 2018 Conference on Artificial Life. MIT Press: Tokyo, p. 204-205

Modeling Cell Migration in a Simulated Bioelectrical Signaling Network for Anatomical Regeneration  
Ferreira, G. B. S., Scheutz, M., and Levin, M. (2018)  
in T. Ikegami, N. Virgo, O. Witkowski, M. Oka, R. Suzuki, and H. Iizuka (Eds.), ALIFE 2018: The 2018 Conference on Artificial Life. MIT Press: Tokyo, p. 194-201

From Physics to Pattern: Uncovering Pattern Formation in Tissue Electrophysiology  
Brodsky, M., and Levin, M. (2018)  
in T. Ikegami, N. Virgo, O. Witkowski, M. Oka, R. Suzuki, and H. Iizuka (Eds.), ALIFE 2018: The 2018 Conference on Artificial Life. MIT Press: Tokyo, p. 351-358

Inform: Efficient Information-Theoretic Analysis of Collective Behaviors  
Moore, D. G., Valentini, G., Walker, S. I., and Levin, M. (2018)  
Frontiers in Robotics and AI, 5: 60

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. (2018) ​​​​  
Physics of Life Reviews, 24: 32-36

Bioelectrical coupling in multicellular domains regulated by gap junctions: a conceptual approach  
Cervera, J., Pietak, A., Levin, M., and Mafe, S. (2018)  
Bioelectrochemistry, 123: 45-61 [cover]  
PDF

Bioelectrical control of positional information in development and regeneration: a review of conceptual and computational advances  
Pietak, A., and Levin, M. (2018)  
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. (2018)  
Current Opinion in Biotechnology, 52: 134-144

Avian models and the study of invariant asymmetry: how the chicken and the egg taught us to tell right from left  
Monsoro-Burq, A. H., and Levin, M. (2018)  
International Journal of Developmental Biology, 62(1-3): 63-77

Are planaria individuals? What regenerative biology is telling us about the nature of multicellularity  
Fields, C., and Levin, M. (2018)  
Evolutionary Biology, 45(3): 237-247  

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. (2018)  
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. (2018)  
Cold Spring Harbor Protocols, 2018(3): 154-163

The bioelectric code: An ancient computational medium for dynamic control of growth and form  
Levin, M., and Martyniuk, C. J. (2018)  
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. (2018)  
Biology Open, 7(1): bio031807

Booting up the organism during development: pre-behavioral functions of the vertebrate brain in guiding body morphogenesis  
Herrera-Rincon, C., and Levin, M. (2018)  
Communicative & Integrative Biology, 11(1): e1433440 [cover]

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. (2018)  
BioSystems, 165: 57-70

Inform: a toolkit for information-theoretic analysis of complex systems  
Moore, D. G., Valentini, G., Walker, S. I., and Levin, M. (2018)  
Proceedings of the 2017 IEEE Symposium Series on Computational Intelligence (SSCI), 3258-3265

Introducing Simulated Stem Cells into a Bio-Inspired Cell-Cell Communication Mechanism for Structure Regeneration  
Ferreira, G. B. S., Scheutz, M., and Levin, M. (2018)  
Proceedings of the 2017 IEEE Symposium Series on Computational Intelligence (SSCI), 2778-2785

2017

Ion channel and neurotransmitter modulators as electroceutical approaches to the control of cancer  
Tuszynski, J., Tilli, T. M., and Levin, M. (2017)  
Current Pharmaceutical Design, 23(32): 4827-4841

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. (2017)  
Nature Communications, 8(1): 587

Coordinating heart morphogenesis: A novel role for Hyperpolarization-activated cyclic nucleotidegated (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. (2017)  
Communicative & Integrative Biology, 10(3): e1309488 [cover]

Bioelectric Gene and Reaction Networks: Computational Modeling of Genetic, Biochemical, and Bioelectrical Dynamics in Pattern Regulation  
Pietak, A. M., and Levin, M. (2017)  
Journal of the Royal Society Interface, 14(134): 20170425 [cover]

Bioelectric Signaling in Regeneration: Mechanisms of Ionic Controls of Growth and Form  
McLaughlin, K. A., and Levin, M. (2017)  
Developmental Biology, 433(2): 177-189

HCN4 Ion Channel Function is Required for Early Events that Regulate Anatomical Left-Right Patterning in a Nodal- and Lefty Asymmetric gene Expression-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. (2017)  
Biology Open, 6(10): 1445-1457

Cancer as a Disorder of Patterning Information: computational and biophysical perspectives on the cancer problem  
Moore, D., Walker, S., and Levin, M. (2017)  
Convergent Science Physical Oncology, 3(4): 043001

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. (2017)  
Developmental Biology, 424(1): 1-9 [cover]

Using optogenetics in vivo to stimulate regeneration  
Adams, D. S., Tseng. A-S., and Levin, M. (2017)  
in K. Appasani (Ed.), Optogenetics: From Neuronal Function to Mapping and Disease Biology, Cambridge University Press: Cambridge, UK, pp. 66-76

Investigating the Effects of Noise on a Cell-to-Cell Communication Mechanism for Structure Regeneration  
Ferreira, G. B., Scheutz, M., and Levin, M. (2017)  
in C. Knibbe, D. Parsons, D. Misevic, J. Rouzaud-Cornaba, N. Bredèche, S. Hassas, O. Simonin, and H. Soula (Eds.), Proceedings of the 14th European Conference on Artificial Life (ECAL 2017), Lyon, France, pp. 170-177

The Zahn Drawings: new illustrations of Xenopus embryo and tadpole stages for studies of craniofacial development  
Zahn, N., Levin, M., and Adams, D. S. (2017)  
Development, 144(15): 2708-2713 [cover]

Microfluidic Platform to Study Intercellular Connectivity Through on-Chip Electrical Impedance Measurement  
Dungan, J., Mathews, J., Levin, M., and Koomson, V. (2017)  
IEEE 60th International Midwest Symposium on Circuits and Systems, Boston, MA, p.56-69

Bioelectric regulation of innate immune system function in regenerating and intact Xenopus laevis  
Paré, J-F., Martyniuk, C. J., and Levin, M. (2017)  
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. (2017)  
Biophysical Journal, 112(10): 2231-2243 [cover]

Space travel has effects on planarian regeneration that cannot be explained by a null hypothesis  
Levin, M., Morokuma, J., and Finkelstein, J. (2017)  
Regeneration, 4(4): 156-158

Planarian regeneration in space: persistent anatomical, behavioral, and bacteriological changes induced by space travel  
Morokuma, J., Durant, F. R., Williams, K. B., Finkelstein, J. M., Blackiston, D. J., Clements, T., Reed, D. W., Roberts, M., Jain, M., Kimel, K., Trayger, S. A., Wolfe, B. E., and Levin, M. (2017)  
Regeneration, 4(2): 85-102 [cover]

Gap Junctional Signaling in Pattern Regulation: physiological network connectivity instructs growth and form  
Mathews, J., and Levin, M. (2017)  
Developmental Neurobiology, 77(5): 643-673

Endogenous Bioelectric Signaling Networks: Exploiting Voltage Gradients for Control of Growth and Form  
Levin, M., Pezzulo, G., and Finkelstein, J. M. (2017)  
Annual Review of Biomedical Engineering, 19: 353-87

IK channel activation increases tumor growth and induces differential behavioral responses in two breast epithelial cell lines  
Thurber, A. E., Nelson, M., Frost, C. L., Levin, M., Brackenbury, W. J., and Kaplan, D. L. (2017)  
Oncotarget, 8(26): 42382-42397

Serotonergic stimulation induces nerve growth and promotes visual learning via posterior eye grafts in a vertebrate model of induced sensory plasticity  
Blackiston, D., Vien, K., and Levin, M. (2017)  
npj Regenerative Medicine, 2: 8

Discovering novel phenotypes with automatically inferred dynamic models: partial melanocyte conversion in Xenopus  
Lobo, D., Lobikin, M., and Levin, M. (2017)  
Scientific Reports, 7: 41339

Reversals of bodies, brains, and behavior: quantitative analysis of laterality and its disturbance in model species  
Blackiston, D. J., and Levin, M. (2017)  
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. (2017)  
Information Sciences, 381: 229-249

Computing a worm: reverse-engineering planarian regeneration  
Lobo, D., and Levin, M. (2017)  
in Adamatzky, A. (Ed.), Advances in Unconventional Computing. Emergence, Complexity and Computation, vol 23, Springer: Cham, pp. 637-654

A Computational Model of Planarian Regeneration  
De, A., Chakravarthy, V. S., and Levin, M. (2017)  
International Journal of Parallel, Emergent, and Distributed Systems, 32(4): 331-347

2016

 
Ahead of the Curve: Hidden Breakthroughs in the Biosciences, Vol. 1  
Levin, M., and Adams, D. S. (2016)  
IOP Publishing: Bristol, UK [cover]

Introduction to provocative questions in Left-Right asymmetry  
Levin, M., Klar, A. J. S., and Ramsdell, A. (2016)  
Philosophical Transactions of the Royal Society B, 371(1710): 20150399

From cytoskeletal dynamics to organ asymmetry: a nonlinear, regulative pathway underlies left-right patterning  
McDowell, G., Rajadurai, S., and Levin, M. (2016)  
Philosophical Transactions of the Royal Society B, 371(1710): 20150409 [cover]

Top-down models in biology: explanation and control of complex living systems above the molecular level  
Pezzulo, G., and Levin, M. (2016)  
Journal of the Royal Society Interface, 13(124): 20160555

Bioelectrical coordination of cell activity toward anatomical target states: an engineering perspective on regeneration  
Herrera-Rincon, C., Guay, J., and Levin, M. (2016)  
in D. Gardiner (Ed.), Regenerative Engineering and Developmental Biology: Principles and Applications, CRC Press: Boca Raton FL, Chapter 4, pp. 55-112

Vertically and horizontally-transmitted memories – the fading boundaries between regeneration and inheritance in planaria  
Neuhof, M., Levin, M., and Rechavi, O. (2016)  
Biology Open, 5(9): 1177-1188 [cover]

Dynamic structure discovery and repair for 3D cell assemblages  
Ferreira, G., Smiley, M., Scheutz, M., and Levin, M. (2016)  
Proceedings of the Fifteenth International Conference on the Synthesis and Simulation of Living Systems (ALIFEXV), pp. 352-359

A Level Set Approach to Simulating Xenopus laevis Tail Regeneration  
Serlin, Z., Rife, J., and Levin, M. (2016)  
Proceedings of the Fifteenth International Conference on the Synthesis and Simulation of Living Systems (ALIFEXV), pp. 528-535

Exploring Instructive Physiological Signaling with the Bioelectric Tissue Simulation Engine (BETSE)  
Pietak, A., and Levin, M. (2016)  
Frontiers in Bioengineering and Biotechnology, 4: 55

On Having No Head: Cognition throughout Biological Systems  
Baluska, F., and Levin, M. (2016)  
Frontiers in Psychology, 7: 902

Genome-wide analysis reveals conserved transcriptional responses downstream of resting potential change in Xenopus embryos, axolotl regeneration, and human mesenchymal cell differentiation  
Pai, V. P., Martyniuk, C. J., Echeverri, K., Sundelacruz, S., Kaplan, D. L., and Levin, M. (2016)  
Regeneration, 3(1): 3-25

A Tunable Silk Hydrogel Device for Studying Limb Regeneration in Adult Xenopus laevis  
Golding, A., Guay, J. A., Herrera-Rincon, C., Levin, M., and Kaplan, D. L. (2016)  
PLoS One, 11(6): e0155618

Physiological Inputs Regulate Species-Specific Anatomy During Embryogenesis and Regeneration  
Sullivan, K. G., Emmons-Bell, M., and Levin, M. (2016)  
Communicative & Integrative Biology, 9(4): e1192733 [cover]

Computational discovery and in vivo validation of hnf4 as a regulatory gene in planarian re-generation  
Lobo, D., Morokuma, J., and Levin, M. (2016)  
Bioinformatics, 32(17): 2681-2685

Color and intensity discrimination in Xenopus laevis tadpoles  
Rothman, G., Blackiston, D., and Levin, M. (2016)  
Animal Cognition, 19(5): 911-919

Telocytes in their context with other intercellular communication agents  
Edelstein, L., Fuxe, K., Levin, M., Popescu, B., and Smythies, J. (2016)  
Seminars in Cell and Developmental Biology, 55: 9-13

Formin Is Associated with Left-Right Asymmetry in the Pond Snail and the Frog  
Davison, A., McDowell, G. S., Holden, J. M., Johnson, H. F., Koutsovoulos, G. D., Liu, M. M., Hulpiau, P., Van Roy, F., Wade, C. W., Banerjee, R., Yang, F., Chiba, S., Davey, J. W., Jackson, D. J., Levin, M., and Blaxter, M. L. (2016)  
Current Biology, 26(5): 654-660 [cover]

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. (2016)  
Regeneration, 3(2): 78-102 [cover]

Neurotransmitter signaling pathways required for normal development in Xenopus laevis embryos: a pharmacological survey screen  
Sullivan, K. G., and Levin, M. (2016)  
Journal of Anatomy, 229(4): 483-502 [cover]

Use of genetically encoded, light-gated ion translocators to control tumorigenesis  
Chernet, B. T., Adams, D. S., Lobikin, M., and Levin, M. (2016)  
Oncotarget, 7(15): 19575-19588

Bioelectric signaling via potassium channels: a mechanism for craniofacial dysmorphogenesis in KCNJ2-associated Andersen-Tawil Syndrome  
Adams, D. S., Uzel, S. G. M., Akagi, J., Wlodkowic, D., Andreeva, V., Yelick, P. C., Devitt-Lee, A., Paré, J-F., and Levin, M. (2016)  
Journal of Physiology, 594(12): 3245-3270 [cover]

Conserved roles for cytoskeletal components in determining laterality  
McDowell, G. S., Lemire, J. M., Paré, J-F., Cammarata, G., Lowery, L. A., and Levin, M. (2016)  
Integrative Biology, 8(3): 267-286 [cover]

Bioelectric modulation of macrophage polarization  
Li, C., Levin, M., and Kaplan, D. (2016)  
Scientific Reports, 6: 21044

On chirality of slime mould  
Dimonte, A., Adamatzky, A., Erokhin, V., and Levin, M. (2016)  
BioSystems, 140: 23-27

MoCha: molecular characterization of unknown pathways  
Lobo, D., Hammelman, J., and Levin, M. (2016)  
Journal of Computational Biology, 23(4): 291-297

Towards a physarum learning chip  
Whiting, J. G. H., Jones, J., Bull, L., Levin, M., and Adamatzky, A. (2016)  
Scientific Reports, 6: 19948

Artificial neural networks as models of robustness in development and regeneration: stability of memory during morphological remodeling  
Hammelman, J., Lobo, D., and Levin, M. (2016)  
in S. Shanmuganathan and S. Samarasinghe (Eds.), Artificial Neural Network (ANN) Modeling, vol. 628, pp. 45-65

2015

Comparison of the depolarization response of human mesenchymal stem cells from different donors  
Sundelacruz, S., Levin M., and Kaplan, D. L. (2015)  
Scientific Reports, 5: 18279

Gap junctional blockade stochastically induces different species-specific head anatomies in genetically wild-type Girardia dorotocephala flatworms  
Emmons-Bell, M., Durant, F., Hammelman, J., Bessonov, N., Volpert, V., Morokuma, J., Pinet, K., Adams, D. S., Pietak, A., Lobo, D., and Levin, M. (2015)  
International Journal of Molecular Sciences, 16(11): 27865-27896 [cover]

Re-membering the body: applications of computational neuroscience to the top-down control of regeneration of limbs and other complex organs  
Pezzulo, G., and Levin, M. (2015)  
Integrative Biology, 7(12): 1487-1517 [cover]

Automatic neuron segmentation and neural network analysis method for phase contrast microscopy images  
Pang, J., Özkucur, N., Ren, M., Kaplan, D. L., Levin, M., and Miller, E. L. (2015)  
Biomedical Optics Express, 6(11): 4395-4416

Serotonergic regulation of melanocyte conversion: A bioelectrically regulated network for stochastic all-or-none hyperpigmentation  
Lobikin, M., Lobo, D., Blackiston, D. J., Martyniuk, C. J., Tkachenko, E., and Levin, M. (2015)  
Science Signaling, 8(397): ra99

Bioelectric memory: modeling resting potential bistability in amphibian embryos and mammalian cells  
Law, R., and Levin, M. (2015)  
Theoretical Biology and Medical Modelling, 12(1): 22

H+/K+-ATPase activity is required for biomineralization in sea urchin embryos  
Schatzberg, D., Lawton, M., Hadyniak, S. E., Ross, E. J., Carney, T., Beane, W. S., Levin, M., and Bradham, C. A. (2015)  
Developmental Biology, 406(2): 259-270

Target morphology and cell memory: a model of regenerative pattern formation  
Bessonov, N., Levin, M., Morozova, N., Reinberg, N., Tosenberger, A., and Volpert, V. (2015)  
Neural Regeneration Research, 10(12): 1901-1905 [cover]

The stability of memories during brain remodeling: A perspective  
Blackiston, D. J., Shomrat, T., and Levin, M. (2015)  
Communicative & Integrative Biology, 8(5): e1073424 [cover]

Local and long-range endogenous resting potential gradients antagonistically regulate apoptosis and proliferation in the embryonic CNS  
Pai, V. P., Lemire, J. M., Chen, Y., Lin G., and Levin, M. (2015)  
International Journal of Developmental Biology, 59(7-9): 327-340

Selective depolarization of transmembrane potential alters muscle patterning and muscle cell localization in embryonic Xenopus Laevis  
Lobikin, M., Paré, J-F., Kaplan, D. L., and Levin, M. (2015)  
International Journal of Developmental Biology, 59(7-9): 303-311 [cover]

Inferring regulatory networks from experimental morphological phenotypes: a computational method reverse-engineers planarian regeneration  
Lobo, D., and Levin, M. (2015)  
PLoS Computational Biology, 11(6): e1004295

A conceptual model of morphogenesis and regeneration  
Tosenberger, A., Bessonov, N., Levin, M., Reinberg, N., Volpert, V., and Morozova, N. (2015)  
Acta Biotheoretica, 63(3): 283-294

Gap junctions: versatile mediators of long-range developmental signals  
Levin, M. (2015)  
The Node

Nerves read the electrical typography of their microenvironment in making growth decisions  
Levin, M. (2015)  
The Node

Knowing one's place: a free energy approach to pattern regulation  
Friston, K., Levin, M., Sengupta B., and Pezzulo, G. (2015)  
Journal of the Royal Society Interface, 12(105): 20141383

Endogenous gradients of resting potential instructively pattern embryonic neural tissue via Notch signaling and regulation of proliferation  
Pai, V., Lemire, J. M., Paré, J-F., Lin, G., Chen, Y., and Levin, M. (2015)  
Journal of Neuroscience, 35(10): 4366-4385

Membrane potential depolarization causes alterations in neuron arrangement and connectivity in cocultures  
Özkucur, N., Quinn, K. P., Pang, J. C., Du, C., Georgakoudi, I., Miller, E., Levin, M., and Kaplan, D. L. (2015)  
Brain and Behavior, 5(1): 24-38

On a model of pattern recognition based on cell memory  
Bessonov, N., Levin, M., Morozova, N., Reinberg, N., Tosenberger, A., and Volpert, V. (2015)  
PLoS One, 10(2): e0118091

A novel method for inducing nerve growth via modulation of host resting potential  
Blackiston, D., Anderson, G. M., Rahman, N., Bieck, C., and Levin, M. (2015)  
Neurotherapeutics, 12(1): 170-184

Long-range gap junctional signaling controls oncogene-mediated tumorigenesis in Xenopus laevis embryos  
Chernet, B. T., Fields, C., and Levin, M. (2015)  
Frontiers in Physiology, 5: 519

2014

Molecular bioelectricity: how endogenous voltage potentials control cell behavior and instruct pattern regulation in vivo  
Levin, M. (2014)  
Molecular Biology of the Cell, 25(24): 3835-3850

Optogenetic Control of Apoptosis in Targeted Tissues of Xenopus laevis Embryos  
Jewhurst, K., Levin, M., and McLaughlin, K. A. (2014)  
Journal of Cell Death, 7: 25-31

Optogenetics in developmental biology: using light to control ion flux-dependent signals in Xenopus embryos  
Adams, D.S., Lemire, J. M., Kramer, R. H., and Levin, M. (2014)  
International Journal of Developmental Biology, 58(10-12): 851-861

Left-right patterning in Xenopus conjoined twin embryos requires serotonin signaling and gap junctions  
Vandenberg, L. N., Blackiston, D. J., Rea, A. C., Dore, T. M., and Levin, M. (2014)  
International Journal of Developmental Biology, 58(10-12): 799-809

Fishing on chips: up-and-coming technological advances in analysis of zebrafish and Xenopus embryos  
Zhu, F., Skommer, J., Huang, Y., Akagi, J., Adams, D. S., Levin, M., Hall, C. J., Crosier, P. S., and Wlodkowic, D. (2014)  
Cytometry A, 85(11): 921-932

Limbform: a functional ontology-based database of limb regeneration experiments  
Lobo, D., Feldman, E. B., Shah, M., Malone, T. J., and Levin, M. (2014)  
Bioinformatics, 30(24): 3598-3600

Depolarization of Cellular Resting Membrane Potential Promotes Neonatal Cardiomyocyte Proliferation In Vitro  
Lan, J. Y., Williams, C., Levin, M., and Black, L. D. III (2014)  
Cellular and Molecular Bioengineering, 7(3): 432-445

Bioelectrical mechanisms for programming growth and form: taming physiological networks for soft body robotics  
Mustard, J., and Levin, M. (2014)  
Soft Robotics, 1(3): 169-191

A bioinformatics expert system linking functional data to anatomical outcomes in limb regeneration  
Lobo, D., Feldman, E. B., Shah, M., Malone, T., and Levin, M. (2014)  
Regeneration, 1(2): 37-56

Design of a flexible component gathering algorithm for converting cell-based models to graph representations for use in evolutionary search  
Budnikova, M., Habig, J. W., Cornia, N., Levin. M., Lobo, D., and Andersen, T. (2014)  
BMC Bioinformatics, 15(1): 178

Transmembrane voltage potential of somatic cells controls oncogene-mediated tumorigenesis at long-range  
Chernet, B., and Levin, M. (2014)  
Oncotarget, 5(10): 3287-3306 [cover]

Alteration of bioelectrically controlled processes in the embryo: a teratogenic mechanism for anticonvulsants  
Hernandez-Diaz S., and Levin, M. (2014)  
Reproductive Toxicology, 47: 111-114

Bioelectric controls of stem cell function  
Pai, V., and Levin, M. (2014)  
in F. Calegari and C. Waskow (Eds.), Stem Cells: From Basic Research to Therapy, Volume 1, CRC Press: Boca Raton FL, Chapter 5, pp. 106-145  

Endogenous bioelectrical networks store non-genetic patterning information during development and regeneration  
Levin, M. (2014)  
Journal of Physiology, 592(11): 2295-2305

A linear-encoding model explains the variability of the target morphology in regeneration  
Lobo, D., Solano, M., Bubenik, G. A., and Levin, M. (2014)  
Journal of the Royal Society Interface, 11(92): 20130918

Endogenous bioelectric cues as morphogenetic signals in vivo  
Lobikin, M., and Levin, M. (2014)  
in D. Fels, M. Cifra, and F. Scholkmann (Eds.), Fields of the Cell, Research Signpost: Kerala, India, Chapter 15, pp. 283-302

V-ATPase maintains neural stem cells in the developing mouse cortex  
Lange, C., Prenninger, S., Knuckles, P., Taylor, V., Levin, M., and Calegari, F. (2014)  
in S. Nakamuri (Ed.), Handbook of H+ ATPases, Pan Stanford Pte. Ltd: Singapore,  pp. 413-430

2013

It's Never Too Early to Get It Right: A Conserved Role for the Cytoskeleton in Left-Right Asymmetry  
Vandenberg, L. N., Lemire, J. M, and Levin, M. (2013)  
Communicative & Integrative Biology, 6(6): e27155

Remembrance of Brains Past  
Levin, M. (2013)  
The Node 

Light-activated serotonin for exploring its action in biological systems  
Rea, A. C., Vandenberg, L. N., Ball, R. E., Snouffer, A. A., Hudson, A. G., Zhu, Y., McLain, D. E., Johnston, L. L., Lauderdale, J. D., Levin, M., and Dore, T. M. (2013)  
Chemistry & Biology, 20(12): 1536-1546

Bioelectric modulation of wound healing in a 3D in vitro model of tissue-engineered bone  
Sundelacruz, S., Li, C., Choi, Y. J., Levin, M., and Kaplan, D. L. (2013)  
Biomaterials, 34(28): 6695-6705

Depolarization Alters Phenotype, Maintains Plasticity of Predifferentiated Mesenchymal Stem Cells  
Sundelacruz, S., Levin, M., and Kaplan, D. L. (2013)  
Tissue Engineering Part A, 19(17-18): 1889-1908

Endogenous voltage potentials and the microenvironment: bioelectric signals that reveal, induce, and normalize cancer  
Chernet, B., and Levin, M. (2013)  
Journal of Clinical and Experimental Oncology, Suppl 1: S1-002  [cover]

Reprogramming cells and tissue patterning via bioelectrical pathways: molecular mechanisms and biomedical opportunities  
Levin, M. (2013)  
Wiley Interdisciplinary Reviews: Systems Biology and Medicine, 5(6): 657-676

An automated training paradigm reveals long-term memory in planaria and its persistence through head regeneration  
Shomrat, T., and Levin, M. (2013)  
Journal of Experimental Biology, 216(20): 3799-3810  
Overview

Bioelectrical signaling has rich history  
Levin, M. (2013)  
Physics Today, 66(9): 11

Inversion of left-right asymmetry alters performance of Xenopus tadpoles in nonlateralized cognitive tasks  
Blackiston, D. J., and Levin, M. (2013)  
Animal Behaviour, 86(2): 459-466

A unified model for left-right asymmetry? Comparison and synthesis of molecular models of laterality  
Vandenberg, L. N., and Levin, M. (2013)  
Developmental Biology, 379(1): 1-15  [cover]

Towards a bioinformatics of patterning: a computational approach to understanding regulative morphogenesis  
Lobo, D., Malone, T. J., and Levin, M. (2013)  
Biology Open, 2(2): 156-169 [cover]

Rab GTPases are required for early orientation of the left-right axis in Xenopus  
Vandenberg, L. N., Morrie, R. D., Seebohm, G., Lemire, J. M., and Levin, M. (2013)  
Mechanisms of Development, 130(4-5): 254-271  [cover]

Endogenous Voltage Gradients as Mediators of Cell-Cell Communication: Strategies for Investigating Bioelectrical Signals During Pattern Formation  
Adams, D. S., and Levin, M. (2013)  
Cell and Tissue Research, 352(1): 95-122  [cover]

Ectopic eyes outside the head in Xenopus tadpoles provide sensory data for light-mediated learning  
Blackiston, D. J., and Levin, M. (2013)  
Journal of Experimental Biology, 216(6): 1031-1040

Planform: an application and database of graph-encoded planarian regenerative experiments  
Lobo, D., Malone, T. J., and Levin, M. (2013)  
Bioinformatics, 29(8): 1098-1100

Humane anesthesia and pain management in amphibian limb surgery of Rana pipiens  
Koustubhan, P., Kaplan, D., and Levin, M. (2013)  
Cold Spring Harbor Protocols, 2013(2): 149-155  [cover]

Transmembrane voltage potential is an essential cellular parameter for the detection and control of tumor development  
Chernet, B. T., and Levin, M. (2013)  
Disease Models & Mechanisms, 6(3): 595-607

Cracking the bioelectric code: probing endogenous ionic controls of pattern formation  
Tseng, A-S., and Levin, M. (2013)  
Communicative & Integrative Biology, 6(1): e22595   [cover]

Light-activation of the Archaerhodopsin H+ pump reverses age-dependent loss of vertebrate regeneration: sparking system-level controls in vivo  
Adams, D. S., Tseng, A-S. and Levin, M. (2013)  
Biology Open, 2(3): 306-313

Serotonin has early, cilia-independent roles in Xenopus left-right patterning  
Vandenberg, L. N., Lemire, J. M., and Levin, M. (2013)  
Disease Models and Mechanisms, 6(1): 261-268  [cover]

Bioelectric signaling regulates head and organ size during planarian regeneration  
Beane, W. S., Morokuma, J., Lemire, J. M., and Levin, M. (2013)  
Development, 140(2): 313-322

2012

Low frequency vibrations induce malformations in two aquatic species in a frequency-, waveform-, and direction-specific manner  
Vandenberg, L. N., Stevenson, C., and Levin, M. (2012)  
PLoS One, 7(12): e51473

Neurally derived tissues in Xenopus laevis embryos exhibit a consistent bioelectrical left-right asymmetry  
Pai, V. P., Vandenberg, L. N., Blackiston, D. J., and Levin, M. (2012)  
Stem Cells International, 2012: 353491

Resting potential, oncogene-induced tumorigenesis, and metastasis: the bioelectric basis of cancer in vivo  
Lobikin, M., Chernet, B., Lobo, D., and Levin, M. (2012)  
Physical Biology, 9(6): 065002  [cover]

Early nonciliary role for microtubule proteins in left–right patterning is conserved across kingdoms  
Lobikin, M., Wang, G., Xu, J.-S., Hsieh, Y.-W., Chuang, C.-F., Lemire, J. M., and Levin, M. (2012)  
Proceedings of the National Academy of Sciences of the United States, 109(31): 12586-12591  

Transducing bioelectrical signals into epigenetic pathways during tadpole tail regeneration  
Tseng, A.-S., and Levin, M. (2012)  
Anatomical Record, 295(10): 1541-1551

Morphogenetic fields in embryogenesis, regeneration, and cancer: non-local control of complex patterning  
Levin, M. (2012)  
BioSystems, 109(3): 243­-261

Normalized shape and location of perturbed craniofacial structures in the Xenopus tadpole reveal an innate ability to achieve correct morphology  
Vandenberg, L. N., Adams, D. S., and Levin, M. (2012)  
Developmental Dynamics, 241(5): 863-878 [cover]

Aversive training methods in Xenopus laevis: general principles  
Blackiston, D. J., and Levin, M. (2012)  
Cold Spring Harbor Protocols, 2012(5): 521-527

Inhibition of planar cell polarity extends neural growth during regeneration, homeostasis, and development  
Beane, W. S., Tseng, A.-S., Morokuma, J. M., Lemire, J. M., and Levin, M. (2012)  
Stem Cells and Development, 21(12): 2085-2094 [cover]

Photoconversion for Tracking the Dynamics of Cell Movement in Xenopus laevis Embryos  
Chernet, B. T., Adams, D. S., and Levin, M. (2012)  
Cold Spring Harbor Protocols, 2016(6): 683-690 [cover]

A versatile protocol for mRNA electroporation of Xenopus laevis embryos  
Chernet, B. T., and Levin, M. (2012)  
Cold Spring Harbor Protocols, 2012(4): 447-452

General principles for measuring resting membrane potential and ion concentration using fluorescent bioelectricity reporters  
Adams, D. S., and Levin, M. (2012)  
Cold Spring Harbor Protocols, 2012(4): 385-397 [cover]

Measuring resting membrane potential using the fluorescent voltage reporters DiBAC4(3) and CC2-DMPE  
Adams, D. S., and Levin, M. (2012)  
Cold Spring Harbor Protocols, 2012(4): 459-464

Transmembrane voltage potential controls embryonic eye patterning in Xenopus laevis  
Pai, V., Aw, S., Shomrat, T., Lemire, J. M., and Levin, M. (2012)  
Development, 139(2): 313-323

Molecular bioelectricity in developmental biology: new tools and recent discoveries  
Levin, M. (2012)  
BioEssays, 34(3): 205-217  [cover]

Polarity proteins are required for left-right axis orientation and twin-twin instruction  
Vandenberg, L. N., and Levin, M. (2012)  
Genesis, 50(3): 219-234

Regulation of Cell Behavior and Tissue Patterning by Bioelectrical Signals: challenges and opportunities for biomedical engineering  
Levin, M., and Stevenson, C. (2012)  
Annual Reviews in Biomedical Engineering, 14: 295-323

Modeling planarian regeneration: a primer for reverse-engineering the worm  
Lobo, D., Beane, W., and Levin, M. (2012)  
PLoS Computational Biology, 8(4): e1002481  [cover]

2011

Transmembrane potential of GlyCl-expressing instructor cells induces a neoplastic-like conversion of melanocytes via a serotonergic pathway  
Blackiston, D., Adams, D. S., Lemire, J. M., Lobikin, M., and Levin, M. (2011)  
Disease Models and Mechanisms, 4(1): 67-85  [cover]

A chemical genetics approach reveals H,K-ATPase-mediated membrane voltage is required for planarian head regeneration  
Beane, W. S., Morokuma, J., Adams, D. S., and Levin, M. (2011) ​​​​​  
Chemistry & Biology, 18(1): 77-89

The H(+) vacuolar ATPase maintains neural stem cells in the developing mouse cortex  
Lange, C., Prenninger, S., Knuckles, P., Taylor, V., Levin, M., and Calegari, F. (2011)  
Stem Cells and Development, 20(5): 843-850

Histone deacetylase activity is necessary for left-right patterning during vertebrate development  
Carneiro, K., Donnet, C., Rejtar, T., Karger, B. L., Díaz, E., Kortagere, S., Lemire, J. M., and Levin, M. (2011)  
BMC Developmental Biology, 11: 29

Low-frequency vibrations disrupt left-right patterning in the Xenopus embryo  
Vandenberg, L. N., Pennarola, B. W., and Levin, M. (2011)  
PLoS One, 6(8): e23306

Patterned femtosecond-laser ablation of Xenopus laevis melanocytes for studies of cell migration, wound repair, and developmental processes  
Mondia, J. P., Adams, D. S., Orendorff, R. D., Levin, M., and Omenetto, F. (2011)  
Biomedical Optics Express, 2(8): 2383-2391

Long-distance signals are required for morphogenesis of the regenerating Xenopus tadpole tail, as shown by femtosecond-laser ablation  
Mondia, J. P., Levin, M., Omenetto, F. G., Orendorff, R. D., Branch, M. R., and Adams, D. S. (2011)  
PLoS One, 6(9): e24953

HDAC activity is required during Xenopus tail regeneration  
Tseng, A-S., Carneiro, K., Lemire, J. M., and Levin, M. (2011)  
PLoS One, 6(10): e26382

Endogenous Bioelectric Signals as Morphogenetic Controls of Development, Regeneration, and Neoplasm  
Levin, M. (2011)  
in C. Pullar (Ed.), The Physiology of Bioelectricity in Development, Tissue Regeneration, and Cancer, CRC Press: Boca Raton FL, pp. 39-89

The wisdom of the body: future techniques and approaches to morphogenetic fields in regenerative medicine, developmental biology, and cancer  
Levin, M. (2011) ​​​​  
Regenerative Medicine, 6(6): 667-673

2010

A second-generation device for automated training and quantitative behavior analyses of molecularly tractable model organisms  
Blackiston, D., Shomrat, T., Nicolas, C. L., Granata, C., and Levin, M. (2010)  
PLoS One, 5(12): e14370

Long-range Neural and Gap Junction Protein-mediated Cues Control Polarity During Planarian Regeneration  
Oviedo, N. J., Morokuma, J., Walentek, P., Kema, I. P., Gu, M. B., Ahn, J. M., Hwang, J. S., Gojobori, T., and Levin, M. (2010)  
Developmental Biology, 339(1): 188-199 

Consistent left-right asymmetry cannot be Established by late organizers in Xenopus unless the late organizer is a conjoined twin  
Vandenberg, L. N., and Levin, M. (2010)  
Development, 137(7): 1095-1105 [cover]

The ATP-sensitive K+-channel (KATP) controls early left-right patterning in Xenopus and chick embryos  
Aw, S., Koster, J., Pearson, W., Nicols, C., Shi, N. Q., Carneiro, K., and Levin, M. (2010)  
Developmental Biology, 346(1): 39-53

Induction of vertebrate regeneration by a transient sodium current  
Tseng, A-S., Beane, W. S., Lemire, J. M., Masi, A., and Levin, M. (2010)  
Journal of Neuroscience, 30(39): 13192-13200 [cover]

BioDome regenerative sleeve for biochemical and biophysical stimulation of tissue regeneration  
Hechavarria, D., Dewilde, A., Braunhut, S., Levin, M., and Kaplan, D. K. (2010)  
Medical Engineering and Physics, 32(9): 1065-1073

High-throughput Xenopus laevis immunohistochemistry using agarose sections  
Blackiston, D., Vandenberg, L. N., and Levin, M. (2010) ​​​​  
Cold Spring Harbor Protocols, 2010(12): 1364-1371

Far from solved: a perspective on what we know about early mechanisms of left-right asymmetry  
Vandenberg, L. N., and Levin, M. (2010) ​​​​  
Developmental Dynamics, 239(12): 3131-3146 [cover]

2009

Molecular mechanisms establishing consistent left-right asymmetry during vertebrate embryogenesis  
Aw, S., and Levin, M. (2009)  
in I. E. C Sommer and R. S, Kahn (Eds.), Language Lateralization and Psychosis, Cambridge University Press: Cambridge UK, pp. 3-18

Is left-right asymmetry a form of planar cell polarity?  
Aw, S., and Levin, M. (2009)  
Development, 136(3): 355-366

Perspectives and open problems in the early phases of left-right patterning  
Vandenberg, L., and Levin, M. (2009)  
Seminars in Cell and Developmental Biology, 20(4): 456-463  [cover]

Particle tracking model of electrophoretic morphogen movement reveals stochastic dynamics of embryonic gradient  
Zhang, Y., and Levin, M. (2009)  
Developmental Dynamics, 238(8): 1923-1935

Left-right asymmetry in the chick embryo requires core planar cell polarity protein Vangl2  
Zhang, Y., and Levin, M. (2009)  
Genesis, 47(11): 719-728

Bioelectric mechanisms in regeneration: Unique aspects and future perspectives  
Levin, M. (2009)  
Seminars in Cell and Developmental Biology, 20(5): 543-556

Errors of Geometry: regeneration in a broader perspective  
Levin, M. (2009)  
Seminars in Cell and Developmental Biology, 20(6): 643-645

Regeneration: recent advances, major puzzles, and biomedical opportunities  
Levin, M. (2009) ​​​​​  
Seminars in Cell and Developmental Biology, 20(5): 515-516

Role of membrane potential in the regulation of cell proliferation and differentiation  
Sundelacruz, S., Levin, M., Levin M., and Kaplan, D. L. (2009)  
Stem Cell Reviews, 5(3): 231-46

Bioelectric controls of cell proliferation: ion channels, membrane voltage, and the cell cycle  
Blackiston, D. J., K. McLaughlin, and Levin, M. (2009)  
Cell Cycle, 8(21): 3527-3536 [cover]

2008

KCNQ1 and KCNE1 K+ channel components are involved in early left-right patterning in Xenopus embryos  
Morokuma, J., Blackiston, D., and Levin, M. (2008)  
Cellular Physiology and Biochemistry, 21(5-6): 357-372

Planarian PTEN homologs regulate stem cells and regeneration through TOR signaling  
Oviedo, N. J., Pearson, B. J., Levin, M., and Alvarado, A. S. (2008)  
Disease Models and Mechanisms, 1(2-3): 131-143

Modulation of potassium channel confers a hyper-proliferative invasive phenotype on embryonic stem cells  
Morokuma, J., Blackiston, D., Adams, D. S., Seebohm, G., Trimmer, B., and Levin, M. (2008)  
Proceedings of the National Academy of Sciences of the United States, 105(43): 16608-16613

Membrane potential controls adipogenic and osteogenic differentiation of mesenchymal stem cells  
Sundelacruz, S., Levin, M., and Kaplan, D. L. (2008)  
PLoS One, 3(11): e3737

Live imaging of planarian membrane potential using DiBAC4(3)  
Oviedo, N. J., Nicolas, C. L., Adams, D. S., and Levin, M. (2008)  
Cold Spring Harbor Protocols, 3(10): pdb.prot5055

Gene knockdown in planarians using RNA interference  
Oviedo, N. J., Nicolas, C. L., Adams, D. S., and Levin, M. (2008)  
Cold Spring Harbor Protocols, 3(10): pdb.prot5054

Establishing and maintaining a colony of planarians  
Oviedo, N. J., Nicolas, C. L., Adams, D. S., and Levin, M. (2008) ​​​​  
Cold Spring Harbor Protocols, 3(10): pdb.prot5053

The planarian regeneration model as a context for the study of drug effects and mechanisms  
Oviedo, N., and Levin, M. (2008)  
in R. B. Raffa & S. M. Rawls (Eds.), Planaria: A Model for Drug Action and Abuse, RG Landes Co.: Austin, pp. 95-104

Analysis of behavior in the planarian model  
Nicolas, C. L., Abramson, C. I., and Levin, M. (2008)  
in R. B. Raffa & S. M. Rawls (Eds.), Planaria: A Model for Drug Action and Abuse, RG Landes Co.: Austin, pp. 83-94

What's Left in Asymmetry?  
Aw, S., and Levin, M. (2008)  
Developmental Dynamics, 237(12): 3453-3464

Planarians: a versatile and powerful model system for molecular studies of regeneration, adult stem cell regulation, aging, and behavior  
Oviedo, N., Nicolas, C. L., Adams, D. S., and Levin, M. (2008)  
Cold Spring Harbor Protocols, 3(10): pdb.em0101

Planarians: a versatile and powerful model system for molecular studies of regeneration, adult stem cell regulation, aging, and behavior  
Oviedo, N., Nicolas, C. L., Adams, D. S., and Levin, M. (2008)  
in Emerging Model Organisms: A Laboratory Manual, Volume 1, Cold Spring Harbor Press: Cold Spring Harbor NY, Chapter 8

Tail regeneration in Xenopus laevis as a model for understanding tissue repair  
Tseng, A-S., and Levin, M. (2008)  
Journal of Dental Research, 87(9): 806-816

2007

Apoptosis is required during early stages of tail regeneration in Xenopus laevis  
Tseng, A-S., Adams, D. S., Qiu, D., Koustubhan, P., and Levin, M. (2007)  
Developmental Biology, 301(1): 62-69

H+ Pump-dependent changes in membrane voltage are an early mechanism necessary and sufficient to induce Xenopus tail regeneration  
Adams, D. S., Masi, A., and Levin, M. (2007) ​​​  
Development, 134(7): 1323-1335 [cover]

smedxin-11 is a Planarian Stem Cell Gap Junction Gene Required for Regeneration and Homeostasis  
Oviedo, N. J., and Levin, M. (2007) ​​​​  
Development, 134(17): 3121-3131

H,K-ATPase protein Localization and Kir4.1 function reveal concordance of three axes during early determination of left-right asymmetry  
Aw, S., Adams, D. S., Qiu, D., and Levin, M. (2007)  
Mechanisms of Development, 125(3-4): 353-372

Establishing and maintaining a Xenopus laevis colony for research laboratories  
Koustubhan, P., Sorocco, D., and Levin, M. (2007)  
in M. Conn (Ed.), Source Book of Models for Biomedical Research, Humana Press, pp. 139-160

Left-right patterning from the inside out: widespread evidence for intracellular control  
Levin, M., and Palmer, R. (2007)  
BioEssays, 29(3): 271-287

Gap junctional communication in morphogenesis  
Levin, M. (2007)  
Progress in Biophysics and Molecular Biology, 94(1-2): 186-206

Large-Scale Biophysics: Ion Flows and Regeneration  
Levin, M. (2007)  
Trends in Cell Biology, 17(6): 261-270 [cover]

What lies at the interface of regenerative medicine and developmental biology?  
Ingber, D., and Levin, M. (2007)  
Development, 134(14): 2541-2547 

Gap junctions provide new links in Left-Right patterning  
Oviedo, N., and Levin, M. (2007)  
Cell, 129(4): 645-647

2006

Ion flow regulates left-right asymmetry in sea urchin development  
Hibino, T., Ishii, Y., Levin, M., and Nishino, A. (2006) ​​​​​  
Development Genes and Evolution, 216(5): 265-276

Evidence for the regulation of left-right asymmetry in Ciona intestinalis by ion flux  
Shimeld, S. M., and Levin, M. (2006) ​​​​​  
Developmental Dynamics, 235(6): 1543-1553

Early, H+-V-ATPase-dependent proton flux is necessary for consistent left-right patterning of non-mammalian vertebrates  
Adams, D. S., Robinson, K. R., Fukumoto, T., Yuan, S., Albertson, R. C., Yelick, P., Kuo, L., McSweeney, M., and Levin, M. (2006) ​​​​​  
Development, 133(9): 1657-1671

Automated Analysis of Behavior: A Computer-Controlled System for Drug Screening and the Investigation of Learning  
Hicks, C., Sorocco, D., and Levin, M. (2006)  
Journal of Neurobiology, 66(9): 977-990

Mathematical Model of Morphogen Electrophoresis through Gap Junctions  
Esser, A. T., Smith, K. C., Weaver, J. C., and Levin, M. (2006)  
Developmental Dynamics, 235(8): 2144-2159

Inverse Drug Screens: a rapid and inexpensive method for implicating molecular targets  
Adams, D. S., and Levin, M. (2006) ​​​​  
Genesis, 44(11): 530-540

Strategies and techniques for investigation of biophysical signals in patterning  
Adams, D., and Levin, M. (2006)  
in M. Whitman and A. K. Sater (Eds.), Analysis of Growth Factor signaling in Embryos. Methods in Signal Transduction Series, CRC Press: Boca Raton FL, pp. 177-264

Of Minds and Embryos: Left-Right Asymmetry and the Serotonergic Controls of Pre-Neural Morphogenesis  
Levin, M., Buznikov, G. A., and Lauder, J. M. (2006) ​​​​​  
Developmental Neuroscience, 28(3): 171-185 [cover]

Is the Early Left-Right Axis like a Plant, a Kidney, or a Neuron? The Integration of Physiological Signals in Left-Right Asymmetry  
Levin, M. (2006) ​​​​  
Birth Defects Research (Part C), 78(3): 191-223

2005

Asymmetric expression of Syndecan-2 in early chick embryogenesis  
Fukumoto, T., and Levin, M. (2005)  
Gene Expression Patterns, 5(4): 525-528

Serotonin signaling is a very early step in patterning of the left-right axis in chick and frog embryos  
Fukumoto, T., Kema, I. P., and Levin, M. (2005)  
Current Biology, 15(9): 794-803

Eye regeneration assay reveals an invariant functional left-right asymmetry in the early bilaterian, Dugesia japonica  
Nogi, T., Yuan, Y. E., Sorocco, D., Perez-Tomas, R., and Levin, M. (2005)  
Laterality, 10(3): 193-205

Localization and loss-of-function implicates ciliary proteins in early, cytoplasmic roles in left-right asymmetry  
Qiu, D., Cheng, S. M., Wozniak, L., McSweeney, M., Perrone, E., and Levin, M. (2005)  
Developmental Dynamics, 234(1): 176-189

Xenopus TRPN1 (NOMPC) localizes to microtubule-based cilia in epithelial cells, including inner-ear hair cells  
Shin, J-B., Adams, D., Paukert, M., Siba, M., Sidi, S., Levin, M., Gillespie, P. G., and Grunder, S. (2005)  
Proceedings of the National Academy of Sciences of the United States, 102(35): 12572-12577

BMP-3 is a novel inhibitor of both activin and BMP-4 signaling in Xenopus embryos  
Gamer, L. W., Nove, J., Levin, M., and Rosen, V. (2005)  
Developmental Biology, 285(1): 156-168

Characterization of innexin gene expression and functional roles of gap-junctional communication in planarian regeneration  
Nogi, T., and Levin, M. (2005)  
Developmental Biology, 287(2): 314-335

Serotonin transporter function is an early step in left-right patterning in chick and frog embryos  
Fukumoto, T., Blakely, R., and Levin, M. (2005) ​​​​​  
Developmental Neuroscience, 27(6): 349-363

Left-right asymmetry in embryonic development: a comprehensive review  
Levin, M. (2005) ​​​​​  
Mechanisms of Development, 122(1): 3-25 [cover]

2004

A novel immunohistochemical method for evaluation of antibody specificity and detection of labile targets in biological tissue  
Levin, M. (2004)  
Journal of Biophysical and Biochemical Methods, 58(1): 85-96  

The embryonic origins of left-right asymmetry  
Levin, M. (2004) ​​​​​  
Critical Reviews in Oral Biology and Medicine, 15(4): 197-206  

Early Patterning of the Left/Right Axis  
Adams, D. S., and Levin, M. (2004)  
in C. D. Stern (Ed.), Gastrulation: from cells to embryo, Cold Spring Harbor Laboratory Press: Long Island  NY, pp. 403-417

2003

Fusicoccin signaling reveals 14-3-3 protein function as a novel step in left-right patterning during amphibian embryogenesis  
Bunney, T. D., De Boer, A. H., and Levin, M. (2003)  
Development, 130(20): 4847-4858

Bioelectromagnetics in morphogenesis  
Levin, M. (2003) ​​​​​  
Bioelectromagnetics, 24(5): 295-315

Motor protein control of ion flux is an early step in embryonic left-right asymmetry  
Levin, M. (2003)  
BioEssays, 25(10): 1002-1010

2002

Asymmetries in H+/K+-ATPase and cell membrane potentials comprise a very early step in left-right patterning  
Levin, M., Thorlin, T., Robinson, K., Nogi, T., and Mercola, M. (2002)  
Cell, 111(1): 77-89

Early embryonic expression of ion channels and pumps in chick and Xenopus development  
Rutenberg, J., Cheng, S. M., and Levin, M. (2002) ​​​​​  
Developmental Dynamics, 225(4): 469-484

Katp channel activity is required for hatching in Xenopus  
Cheng, S. M., Chen, I., and Levin, M. (2002)  
Developmental Dynamics, 225(4): 588-591

Isolation and community: A review of the role of gap-junctional communication in embryonic patterning  
Levin, M. (2002)  
Journal of Membrane Biology, 185(3): 177-192

2001

Left-Right asymmetry determination in vertebrates  
Mercola, M., and Levin, M. (2001) ​​​​​  
Annual Review of Cell and Developmental Biology, 17: 779-805  

Asymmetry of Body and Brain: Embryological and Twin Studies  
Levin, M. (2001)  
in N. Smelser and P. Baltes (Eds.), International Encyclopedia of the Social and Behavioral Sciences, Elsevier: Oxford UK, pp. 853-859

2000

Expression of connexin 30 in Xenopus embryos and its involvement in hatching gland function  
Levin, M., and Mercola, M. (2000)  
Developmental Dynamics, 219(1): 96-101

1999

Matrix-based GA representations in a model of evolving animal communication  
Levin, M. (1999)  
in L. Chambers (Ed.), The Practical Handbook of Genetic Algorithms: Complex Coding Systems, Vol. 3, CRC Press: Boca Raton FL, Chapter 5, pp. 103-117  

Twinning and embryonic left-right asymmetry  
Levin, M. (1999) ​​​​​  
Laterality, 4(3): 197-208

Cerberus regulates left-right asymmetry of the embryonic head and heart  
Zhu, L., Marvin, M. J., Gardiner, A., Lassar, A. B., Mercola, M., Stern, C. D., and Levin, M. (1999)  
Current Biology, 9(17): 931-938  

Gap Junction-Mediated Transfer of Left-Right Patterning Signals in the Early Chick Blastoderm is Upstream of Shh Asymmetry in the Node  
Levin, M., and Mercola, M. (1999)  
Development, 126(21): 4703-4714

Left-right asymmetry in animal embryogenesis  
Levin, M. (1999)  
in G. Palyi, C. Zucchi, and L. Caglioti (Eds.), Advances in Biochirality, Elsevier Science Ltd: Oxford UK, Chapter 12, pp. 137-152  

Endogenous electromagnetic fields and radiations in regeneration, development, and neoplasm  
Levin, M. (1999)  
Proceedings of the First World Congress on the Effects of Electricity and Magnetism in the Natural World, Madeira, Portugal

1998

The roles of activin and follistatin signaling in chick gastrulation  
Levin, M. (1998) ​​​​​  
International Journal of Developmental Biology, 42(4): 553-559

Gap junctions are involved in the early generation of left right asymmetry  
Levin, M., and Mercola, M. (1998)  
Developmental Biology, 203(1): 90-105

Evolutionary conservation of mechanisms upstream of asymmetric nodal expression: Reconciling chick and Xenopus  
Levin, M., and Mercola, M. (1998)  
Developmental Genetics, 23(3): 185-193

Left-Right asymmetry and the chick embryo  
Levin, M. (1998)  
Seminars in Cell & Developmental Biology, 9(1): 67-76

The compulsion of chirality: toward an understanding of left-right asymmetry  
Levin, M., and Mercola, M. (1998)  
Genes & Development, 12(6): 763-769

1997

Applied DC magnetic fields cause alterations in the time of cell divisions and developmental abnormalities in early sea urchin embryos  
Levin, M., and Ernst, S. G. (1997)  
Bioelectromagnetics, 18(3): 255-263

Left/Right Patterning Signals and the Independent Regulation of Different Aspects of Situs in the Chick Embryo  
Levin, M., Pagan, S., Roberts, D. J., Cooke, J., Kuehn, M. R., and Tabin, C. J. (1997)  
Developmental Biology, 189(1): 57-67

Two molecular models of initial left-right asymmetry generation  
Levin, M., and Nascone, N. (1997)  
Medical Hypotheses, 49(5): 429-435

Left-right asymmetry in vertebrate embryogenesis  
Levin, M. (1997)  
BioEssays, 19(4): 287-296 [cover]

1996

Laterality defects in conjoined twins  
Levin, M., Roberts, D. J., Holmes, L. B., and Tabin, C. (1996)  
Nature, 384(6607): 321

1995

Applied AC and DC Magnetic Fields Cause Alterations in the Mitotic Cycle of Early Sea Urchin Embryos  
Levin, M., and Ernst, S. G. (1995)  
Bioelectromagnetics, 16(4): 231-240

A molecular pathway determining left-right asymmetry in chick embryogenesis  
Levin, M., Johnson, R. L., Stern, C. D., Kuehn, M., and Tabin, C. (1995)  
Cell, 82(5): 803-814 [cover]

Use of Genetic Algorithms to Solve Biomedical Problems  
Levin, M. (1995)  
M.D. Computing, 12(3): 193-198

The evolution of understanding: A genetic algorithm model of the evolution of animal communication  
Levin, M. (1995)  
BioSystems, 36(3): 167-178

Locating putative protein signal sequences  
Levin, M. (1995)  
in L. Chambers (Ed.), The Practical Handbook of Genetic Algorithms: New Frontiers, Vol. 2, CRC Press: Boca Raton FL, Chapter 2, pp. 53-66

1994

A Julia set model of field-directed morphogenesis: developmental biology and artificial life  
Levin, M. (1994)  
Computer Applications in the Biosciences, 10(2): 85-103

Discontinuous and alternate q-system fractals  
Levin, M. (1994)  
Computers and Graphics, 18(6): 873-884

1993

Current and potential applications of bioelectromagnetics in medicine  
Levin, M. (1993)  
Subtle Energies and Energetic Medicine Journal, 4(1): 77-87

Conferences, Abstracts, and Poster Sessions

SBMLtoODEjax: Efficient Simulation and Optimization of Biological Network Models in JAX  
Etcheverry, M., Levin, M., Moulin-Frier, C., and Oudeyer, P.-Y. (2023)  
NeurIPS 2023 AI for Science Workshop, New Orleans, LA

Brief multi-drug treatment via a wearable silk-hydrogel bioreactor induces long-term limb re-patterning and regeneration in adult Xenopus Laevis  
Murugan, N., Vigran, H., Miller, K., Pham, Q., Golding, A., Kane, A., Kaplan, D., and Levin, M. (2021)  
The FASEB Journal, 35(S1), doi:10.1096/fasebj.2021.35.s1.03874

Xenopticon: Microfluidic platform for time-resolved analysis of Xenopus embryo development  
Novak, R., Choe, Y., Dinis, A., Ingram, M., Keshari, V., Nestor, B. A., Super, M., Levin, M., and Ingber, D. E. (2020)  
Proceedings of 21st International Conference on Miniaturized Systems for Chemistry and Life Sciences (MicroTAS 2017), 1639-1640

Transmembrane voltage gradient in GlyR-expressing niche cells controls behavior of neural crest derivatives in vivo  
Blackiston, D., Adams, D., Lemire, J., and Levin, M. (2010)  
Developmental Biology, 344(1): 526

Long-range neural and gap junction protein-mediated cues control polarity during planarian regeneration  
Oviedo, N. J., Morokuma, J., Walentek, P., Kema, I. P., Gu, M. B., Ahn, J. M., Hwang, J. S., Gojobori, T., Levin, M. (2010)  
Developmental Biology, 344(1): 522

NaV-mediated sodium transport is required for vertebrate appendage regeneration  
Tseng, A. S., Beane, W. S., Lemire, J. M., Masi, A., and Levin, M. (2010)  
Developmental Biology, 344(1): 519

H,K-ATPase-mediated ion transport regulates anterior patterning in regenerating planaria  
Beane W. S., Morokuma J., and Levin, M. (2010)  
Developmental Biology, 344(1): 522

The Par6 complex is required for both early and late orientation of the left-right axis in Xenopus  
Vandenberg, L., and Levin, M. (2010)  
Developmental Biology, 344(1): 426-427

H,K-ATPase-mediated ion transport regulates anterior patterning in regenerating planaria  
Beane W. S., Oviedo N.J., Adams D. S., and Levin, M. (2009)  
Mechanisms of Development, 126 (Sup. 1): S294-S295

Role of KvLQT-1/minK K+ channel in early embryonic patterning  
Morokuma, J., and Levin, M. (2006)  
Developmental Biology, 295(1): 451

A leftward fluid flow precedes nodal induction in Xenopus  
Blum, M., Schweickert, A., Beyer, T., Weber, T., Andre, P., Fischer, A., Viebahn, C., and Levin, M. (2006)  
Developmental Biology, 295(1): 443

Xenopus tadpole tail regeneration requires the activity of the proton pump V- ATPase, and proton pumping is sufficient to partially rescue the loss of function phenotype  
Adams, D. S., Masi, A., and Levin, M. (2006)  
Developmental Biology, 295(1): 355-356

Bioelectrical controls of morphogenesis  
Levin, M., Adams, D. S., Aw, S., Morokuma, J., and Oviedo, N. J. (2006)  
Developmental Biology, 295(1): 331-332

The Role of KATP Channels in Development of Left-Right Asymmetry in Xenopus  
Chen, I., and Levin, M. (2004)  
Journal of Dental Research, 83: A1340

Electrical Controls of Regeneration in Planaria  
Nogi, T. and Levin, M. (2003)  
Developmental Biology, 259: 586

Electric Embryos: Endogenous Ion Fluxes and Voltage Gradients in Left—Right Asymmetry  
Levin, M. (2003)  
Developmental Biology, 259: 482

Elements of Left—Right Patterning: Gap Junctions, pH, and Membrane Voltage  
Adams, D., and Levin, M. (2003)  
Developmental Biology, 259: 482-483

Serotonin Is a Novel Very Early Signaling Mechanism in Left—Right Asymmetry  
Fukumoto, T., and Levin, M. (2003)  
Developmental Biology, 259: 490

Gap junction-mediated transfer of left-right patterning signals  
Levin, M., and Mercola, M. (1999)  
Molecular Biology of the Cell, 39: 1

From left-right asymmetric gene expression to its morphological manifestations  
Pagan, S. M., Levin, M., and Tabin, C. (1997)  
Journal of Morphology, 232(3): 306

DC and AC magnetic field effects on early sea-urchin development  
Levin, M., Gonzalez, J. L., and Ernst, S. G. (1991)  
Biological Bulletin, 181: 511