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The Levin Lab
Peer-Reviewed Papers
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Peer-Reviewed Papers

Bioelectricity

Cervera, J., Levin, M., Mafe, S. (2024), Multicellular adaptation to electrophysiological perturbations analyzed by deterministic and stochastic bioelectrical models, Scientific Reports 2024, 27608 doi:10.1038/s41598-024-79087-7
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Pai, V., Zhang, G., Levin, M. (2024), “Bioelectricity in Development, Regeneration, and Cancers” Cell Bio 2023: A Joint Meeting of the American Society of Cell Biology and European Molecular Biology Organization, Bioelectricity 2024, 6:1 doi: 10.1089/bioe.2024.0006
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Djamgoz, M.B.A., Levin, M. (2024), Potential Shortcomings of Genomic Database: The Case of Nav1.5 Expression in Breast Cancer, Bioelectricity 2024, doi: 10.1089/bioe.2024.0033
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Cervera, J., Manzanares, J. A., Levin, M., Mafe, S. (2024), Oscillatory phenomena in electrophysiological networks: The coupling between cell bioelectricity and transcription, Computers in Biology and Medicine 2024, 80 doi: 10.1016/j.compbiomed.2024.108964
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Pio-Lopez, L., Levin, M. (2024), Aging as a loss of morphostatic information: a developmental bioelectricity perspective, Ageing Research Reviews, 2024, 97:102310
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Balasubramanian, S., Weston, D., Levin, M., Davidian, D. C. C. (2024), Electroceuticals: emerging applications beyond the nervous system and excitable tissues, Trends Pharmacol Sci, 2024, 2109
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Balasubramanian, S., Weston, D., Levin, M., Davidian, D. C. C. (2024), Charging Ahead: Examining the future therapeutic potential of electroceuticals, Advanced Therapeutics, 2024, 2300344
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O’Brien, T., Stremmel, J., Pio-Lopez, L., McMillen, P., Rasmussen-Ivey, C., and Levin, M. (2024), Machine Learning for Hypothesis Generation in Biology and Medicine: Exploring the latent space of neuroscience and developmental bioelectricity, Digital Discovery, 3(2): 249-263
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McMillen, P., and Levin, M. (2024), Optical Estimation of Bioelectric Patterns in Living Embryos, in M. Bizzarri (ed.), Systems Biology: Methods in Molecular Biology, 2745: 91-102, doi:10.1007/978-1-0716-3577-3_6
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Manicka, S., Pai, V. P, and Levin, M. (2023), Information integration during bioelectric regulation of morphogenesis of the embryonic frog brain, iScience, 26(12): 108398
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Blattner, M., and Levin, M. (2023) Long Range Communication via Gap Junctions and Stress in Planarian Morphogenesis: A Computational Study, Bioelectricity, 5(3): 196-209
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Cervera, J., Levin, M., and Mafé, S. (2023), Correcting instructive electric potential patterns in multicellular systems: External actions and endogenous processes, Biochimica et Biophysica Acta (BBA) - General Subjects, 1867(10): 130440
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Lagasse, E., and Levin, M. (2023), Future Medicine: from molecular pathways to the collective intelligence of the body, Trends in Molecular Medicine, 29(9): 687-710
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Masuelli, S., Real, S., McMillen, P., Oudin, M., Levin, M., and Roqué, M. (2023), The Yin and Yang of Breast Cancer: Ion Channels as Determinants of Left-Right Functional Differences, International Journal of Molecular Sciences, 24(13): 11121
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Cervera, J., Levin, M., and Mafé, S. (2023), Bioelectricidad en agregados multicelulares de células no excitables: modelos biofísicos, Revista Española de Física, 37(2): 24-29

Levin, M. (2023), Bioelectric networks: the cognitive glue enabling evolutionary scaling from physiology to mind, Animal Cognition, doi:10.1007/s10071-023-01780-3
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Pio-Lopez, L., and Levin, M. (2023), Morphoceuticals: Perspectives for discovery of drugs targeting anatomical control mechanisms in regenerative medicine, cancer and aging, Drug Discovery Today, 28(6): 103585
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Davies, J., and Levin, M. (2023), Synthetic morphology with agential materials, Nature Reviews Bioengineering, 1(1): 46-59
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Cervera, J., Levin, M., and Mafe, S. (2023), Bioelectricity of non-excitable cells and multicellular pattern memories: Biophysical modeling, Physics Reports, 1004: 1-31, doi:10.1016/j.physrep.2022.12.003
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Hazan, H., and Levin, M. (2022), Exploring the Behavior of Bioelectric Circuits Using Evolution Heuristic Search, Bioelectricity, 4(4): 207-227
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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), On-chip on-demand delivery of K+ for in vitro bioelectronics, AIP Advances, 12(12): 125205
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Cervera, J., Manzanares, J. A., Levin, M., and Mafe, S. (2022), Transplantation of fragments from different planaria: a bioelectrical model for head regeneration, Journal of Theoretical Biology, 558: 111356
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Pai, V. P., Cooper, B. G., and Levin, M. (2022), Screening Biophysical Sensors and Neurite Outgrowth Actuators in Human Induced-Pluripotent-Stem-Cell-Derived Neurons, Cells, 11(16): 2470
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Pai, V. P., and Levin, M. (2022), HCN2 Channel-induced Rescue of Brain, Eye, Heart, and Gut Teratogenesis Caused by Nicotine, Ethanol, and Aberrant Notch Signaling, Wound Repair and Regeneration, 30(6): 681-706
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Mathews, J., Kuchling, F., Baez-Nieto, D., Diberardinis, M., Pan, J. Q., and Levin, M. (2022), Ion Channel Drugs Suppress Cancer Phenotype in NG108-15 and U87 Cells: Toward Novel Electroceuticals for Glioblastoma, Cancers 14(6): 1499
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Grodstein, J., and Levin, M. (2022), A Computational Approach to Explaining Bioelectrically-induced Persistent, Stochastic Changes of Axial Polarity in Planarian Regeneration, Bioelectricity, 4(1): 18-30
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Adelfio, M., Bonzanni, M., Levin, M., and Kaplan, D. L. (2022), Impact of Membrane Voltage on Formation and Stability of Human Renal Tubules in Vitro, ACS Biomaterials Science & Engineering, 8(3): 1239-1246
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Masuelli, S., Sebastian, R., Campoy, E., Branham, M. T., Marzese, D. M., Matthew, S., DeBlas, G., Rodolfo, A., Levin, M., and Roque, M. (2022), When left does not seem right: epigenetic and bioelectric differences between left-and right-sided breast cancer, Molecular Medicine, 28(1): 15
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Manicka, S., and Levin, M. (2022), Minimal developmental computation: a causal network approach to understand morphogenetic pattern formation, Entropy, 24: 107
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Payne, S. L., Ram, P., Srinivasan, D. H., Le, T. T., Levin, M., and Oudin, M. J. (2021), Potassium channel-driven bioelectric signalling regulates metastasis in triple-negative breast cancer, EBioMedicine, 75: 103767
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Riol, A., Cervera, J., Levin, M., and Mafe, S. (2021), Cell Systems Bioelectricity: How Different Intercellular Gap Junctions Could Regionalize a Multicellular Aggregate, Cancers, 13(21): 5300
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Grodstein, J., and Levin, M. (2021), Stability and robustness properties of bioelectric networks: A computational approach, Biophysics Review, 2: 031305
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Levin, M. (2021), Bioelectrical approaches to cancer as a problem of the scaling of the cellular self, Progress in Biophysics and Molecular Biology, 165: 102-113
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Cervera, J., Levin, M., and Mafe, S. (2021), Morphology changes induced by intercellular gap junction blocking: a reaction-diffusion mechanism, BioSystems, 209: 104511
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Levin, M. (2021), Bioelectric Signaling: Reprogrammable Circuits Underlying Embryogenesis, Regeneration, and Cancer, Cell, 184(8): 1971-1989
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Nanos, V., and Levin, M. (2021), Rewiring Endogenous Bioelectric Circuits in the Xenopus laevis Embryo Model, in M. R. Ebrahimkhani and J. Hislop (Eds.), Programmed Morphogenesis: Methods in Molecular Biology, 2258: 93-103
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Srivastava, P., Kane, A., Harrison, C., and Levin, M. (2021), A Meta-Analysis of Bioelectric Data in Cancer, Embryogenesis, and Regeneration, Bioelectricity, 3(1): 42-67
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Pezzulo, G., Lapalme, J., Durant, F., and Levin, M. (2021), Bistability of Somatic Pattern Memories: Stochastic Outcomes in Bioelectric Circuits Underlying Regeneration, Philosophical Proceedings of the Royal Society B, 376(1821): 20190765
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Cervera, J., Ramirez, P., Levin, M., and Mafe, S. (2020), Community effects allow bioelectrical reprogramming of cell membrane potentials in multicellular aggregates: Model simulations, Physical Review E, 102(5): 052412
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Sousounis, K., Erdogan, B., Levin, M., and Whited, J. L. (2020), Precise control of ion channel and gap junction expression is required for patterning of the regenerating axolotl limb, International Journal of Developmental Biology, 64(10-11-12): 485-494
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Erndt-Marino, J., Yeisley, D. J., Chen, H., Levin, M., Kaplan, D. L., and Hahn, M. S. (2020), Interferon-Gamma Stimulated Murine Macrophages In Vitro: Impact of Ionic Composition and Osmolarity and Therapeutic Implications, Bioelectricity, 2(1): 48-58
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Leronni, A., Bardella, L., Dorfmann, L., Pietak, A., and Levin, M. (2020), On the coupling of mechanics with bioelectricity and its role in morphogenesis, Journal of the Royal Society Interface, 17(167): 20200177
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Pai, V., Cervera, J., Mafe, S., Willocq, V., Lederer, E., and Levin, M. (2020), HCN2 Channel-induced Rescue of Brain Teratogenesis via local and long-range bioelectric repair, Frontiers in Cellular Neuroscience, 14: 136
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Cervera, J., Levin, M., and Mafe, S. (2020), Bioelectrical Coupling of Single-Cell States in Multicellular Systems, Journal of Physical Chemistry Letters, 11(9): 3234-3241
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Levin, M. (2020), Revisiting Burr and Northrop's 'The Electro-Dynamic Theory of Life', Biological Theory, 15: 83-90
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McMillen, P., Novak, R., and Levin, M. (2020), Toward Decoding Bioelectric Events in Xenopus Embryogenesis: New Methodology for Tracking Interplay Between Calcium and Resting Potentials In Vivo, Journal of Molecular Biology, 432(2): 605-620
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Kamaldinov, T., Erndt-Marino, J., Levin, M., Kaplan, D. L., and Hahn, M. S. (2020), Assessment of Enrichment of Human Mesenchymal Stem Cells Based on Plasma and Mitochondrial Membrane Potentials, Bioelectricity, 2(1): 21-32
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Healy, D., Faber, C., Lapalme, J., and Levin, M. (2020), Post-SSRI Sexual Dysfunction: a bioelectric mechanism?, Bioelectricity, 2(1): 7-13
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Levin, M. (2020), The Biophysics of Regenerative Repair Suggests New Perspectives on Biological Causation, BioEssays, 42(2): 1900146
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Cervera, J., Meseguer, S., Levin, M., and Mafe, S. (2020), Bioelectrical model of head-tail patterning based on cell ion channels and intercellular gap junctions, Bioelectrochemistry, 132: 107410
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Levin, M., Selberg, J., and Rolandi, M. (2019), Endogenous Bioelectrics in Development, Cancer, and Regeneration: Drugs and Bioelectronic Devices as Electroceuticals for Regenerative Medicine, iScience, 22: 519-533
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Fields, C., Bischof, J., and Levin, M. (2020), Morphological Coordination: A Common Ancestral Function Unifying Neural and Non-Neural Signaling, Physiology, 35: 16-30
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Bonzanni, M., Payne, S. L., Adelfio, M., Kaplan, D. L., Levin, M., and Oudin, M. J. (2020), Defined extracellular ionic solutions to study and manipulate the cellular resting membrane potential, Biology Open, 9(1): bio048553
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Manicka, S., and Levin, M. (2019), Modeling somatic computation with non-neural bioelectric networks, Scientific Reports, 9: 18612
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Levin, M. (2019), The Computational Boundary of a "Self": developmental bioelectricity drives multicellularity and scale-free cognition, Frontiers in Psychology, 10:2688, doi: 10.3389/fpsyg.2019.02688
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Emmons-Bell, M., Durant, F., Tung, A., Pietak, A., Miller, K., Kane, A., Martyniuk, C. J., Davidian, D., Morokuma, J., and Levin, M. (2019), Regenerative Adaptation To Electrochemical Perturbation In Planaria: A Molecular Analysis Of Physiological Plasticity, iScience, 22: 147-165
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Atsuta, Y., Tomizwa, R. R., Levin, M., and Tabin, C. J. (2019), L-type voltage-gated Ca2+ channel CaV1.2 regulates chondrogenesis during limb development, Proceedings of the National Academy of Sciences of the United States, 116(43): 21592-21601
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Payne, S. L., Levin, M., and Oudin, M. J. (2019), Bioelectric control of metastasis in solid tumors, Bioelectricity, 1(3): 114-130
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Whited, J., and Levin, M. (2019), Bioelectrical controls of morphogenesis: from ancient mechanisms of cell coordination to biomedical opportunities, Current Opinion in Genetics & Development, 57: 61-69
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Cervera, C., Pai, V. P., Levin, M., and Mafe, S. (2019), From non-excitable single-cell to multicellular bioelectrical states supported by ion channels and gap junction proteins: electrical potentials as distributed controllers, Progress in Biophysics and Molecular Biology, 149: 39-53
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Cervera, J., Manzanares, J. A., Mafe, S., and Levin, M. (2019), Synchronization of Bioelectric Oscillations in Networks of Nonexcitable Cells: From Single-Cell to Multicellular States, Journal of Physical Chemistry B, 123(18): 3924-3934
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Sundelacruz, S., Moody, A. T., Levin, M., and Kaplan, D. (2019), Membrane potential depolarization alters calcium flux and phosphate signaling during osteogenic differentiation of human mesenchymal stem cells, Bioelectricity, 1(1): 56-66
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Ferenc, N. N., and Levin, M. (2019), Effects of ivermectin exposure on regeneration of D. dorotocephala planaria: exploiting human-approved ion channel drugs as morphoceuticals, Macromolecular Bioscience, 19(3): 1800237
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Durant, F., Bischof, J., Fields, C., Morokuma, J., LaPalme, J., Hoi, A., and Levin, M. (2019), The role of early bioelectric signals in the regeneration of planarian anterior/posterior polarity, Biophysical Journal, 116(5): 948-961
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Churchill, C. D. M., Winter, P., Tuszynski, J. A., and Levin, M. (2019), EDEn - Electroceutical Design Environment: An Ion Channel Database with Small Molecule Modulators, iScience, 11: 42-56
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Cairns, D. M., Giordano, J. E., Conte, S., Levin, M., and Kaplan, D. L. (2018), Ivermectin promotes peripheral nerve regeneration during wound healing, ACS Omega, 3(10): 12392-12402
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Busse, S. M., McMillen, P. T., and Levin, M. (2018), Cross-limb Communication During Xenopus Hind-limb Regenerative Response: Non-Local Bioelectric Injury Signals, Development, 145(19): dev164210
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Ferreira, G. B. S., Scheutz, M., and Levin, M. (2018), Modeling Cell Migration in a Simulated Bioelectrical Signaling Network for Anatomical Regeneration, 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
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Brodsky, M., and Levin, M. (2018), From Physics to Pattern: Uncovering Pattern Formation in Tissue Electrophysiology, 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
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Cervera, J., Pietak, A., Levin, M., and Mafe, S. (2018), Bioelectrical coupling in multicellular domains regulated by gap junctions: a conceptual approach, Bioelectrochemistry, 123: 45-61
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Pietak, A., and Levin, M., (2018), Bioelectrical control of positional information in development and regeneration: a review of conceptual and computational advances, Progress in Biophysics and Molecular Biology, 137: 52-68
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Mathews, J., and Levin, M. (2018), The Body Electric 2.0: Recent Advances in Developmental Bioelectricity for Regenerative and Synthetic Bioengineering, Current Opinion in Biotechnology, 52: 134-144
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Pai, V. P., Pietak, A., Willocq, V., Ye, B., Shi, N-Q., and Levin, M. (2018), HCN2 Rescues brain defects by enforcing endogenous voltage pre-patterns, Nature Communications, 9(1): 998
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Levin, M., and Martyniuk, C. J. (2018), The bioelectric code: An ancient computational medium for dynamic control of growth and form, BioSystems, 164: 76-93
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Herrera-Rincon, C., and Levin, M. (2018), Booting up the organism during development: pre-behavioral functions of the vertebrate brain in guiding body morphogenesis, Communicative & Integrative Biology, 11(1): e1433440
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Pitcairn, E., Harris, H., Epiney, J., Pai, V. P., Lemire, J. M., Ye, B., Shi, N-Q., Levin, M., and McLaughlin, K. A. (2017), Coordinating heart morphogenesis: A novel role for Hyperpolarization-activated cyclic nucleotidegated (HCN) channels during cardiogenesis in Xenopus laevis, Communicative & Integrative Biology, 10(3): e1309488
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Pietak, A. M., and Levin, M. (2017), Bioelectric Gene and Reaction Networks: Computational Modeling of Genetic, Biochemical, and Bioelectrical Dynamics in Pattern Regulation, Journal of the Royal Society Interface, 14(134): 20170425
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McLaughlin, K. A., and Levin, M. (2017), Bioelectric Signaling in Regeneration: Mechanisms of Ionic Controls of Growth and Form, Developmental Biology, 433(2): 177-189
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Pai, V. P., Willocq, V., Pitcairn, E. J., Lemire, J. M., Paré, J-F., Shi, N-Q., McLaughlin, K. A., and Levin, M. (2017), 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, Biology Open, 6(10): 1445-1457
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Adams, D. S., Tseng, A-S., and Levin, M. (2017), Using optogenetics in vivo to stimulate regeneration, in K. Appasani (Ed.), Optogenetics: From Neuronal Function to Mapping and Disease Biology. Cambridge University Press: Cambridge, UK, p. 66-76

Dungan, J., Mathews, J., Levin, M., and Koomson, V. (2017), Microfluidic Platform to Study Intercellular Connectivity Through on-Chip Electrical Impedance Measurement, IEEE 60th International Midwest Symposium on Circuits and Systems, Boston, MA, p. 56-69
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Paré, J-F., Martyniuk, C. J., and Levin, M. (2017), Bioelectric regulation of innate immune system function in regenerating and intact Xenopus laevis, npj Regenerative Medicine, 2: 15
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Durant, F., Morokuma, J., Fields, C., Williams, K., Adams, D. S., and Levin, M. (2017), Long-Term, Stochastic Editing of Regenerative Anatomy via Targeting Endogenous Bioelectric Gradients, Biophysical Journal, 112(10): 2231-2243
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Mathews, J., and Levin, M. (2017), Gap Junctional Signaling in Pattern Regulation: physiological network connectivity instructs growth and form, Developmental Neurobiology, 77(5): 643-673
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Levin, M., Pezzulo, G., and Finkelstein, J. M. (2017), Endogenous Bioelectric Signaling Networks: Exploiting Voltage Gradients for Control of Growth and Form, Annual Review of Biomedical Engineering, 19: 353-387
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Thurber, A. E., Nelson, M., Frost, C. L., Levin, M., Brackenbury, W. J., and Kaplan D. L. (2017), IK channel activation increases tumor growth and induces differential behavioral responses in two breast epithelial cell lines, Oncotarget, 8(26): 42382-42397
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Lobo, D., Lobikin, M., and Levin, M. (2017), Discovering novel phenotypes with automatically inferred dynamic models: partial melanocyte conversion in Xenopus, Scientific Reports, 7: 41339
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Pezzulo, G., and Levin, M. (2016), Top-down models in biology: explanation and control of complex living systems above the molecular level, Journal of the Royal Society Interface, 13(124): 20160555
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Herrera-Rincon, C., Guay, J., and Levin, M. (2016), Bioelectrical coordination of cell activity toward anatomical target states: an engineering perspective on regeneration, Chapter 4 in D. Gardiner (Ed.), Regenerative Engineering and Developmental Biology: Principles and Applications, CRC Press: Boca Raton, FL, p. 55-112
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Pietak, A., and Levin, M. (2016), Exploring Instructive Physiological Signaling with the Bioelectric Tissue Simulation Engine , Frontiers in Bioengineering and Biotechnology, 4: (55)
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Pai, V. P., Martyniuk, C. J., Echeverri, K., Sundelacruz, S., Kaplan, D. L., and Levin, M. (2016), Genome-wide analysis reveals conserved transcriptional responses downstream of resting potential change in Xenopus embryos, axolotl regeneration, and human mesenchymal cell differentiation, Regeneration, 3(1): 3-25
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Sullivan, K. G., Emmons-Bell, M., and Levin, M. (2016), Physiological Inputs Regulate Species-Specific Anatomy During Embryogenesis And Regeneration, Communicative & Integrative Biology, 9(4): e1192733
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Durant, F., Lobo, D., Hammelman, J., and Levin, M. (2016), Physiological controls of large-scale patterning in planarian regeneration: a molecular and computational perspective on growth and form, Regeneration, 3(2): 78-102
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Chernet, B. T., Adams, D. S., Lobikin, M., and Levin, M. (2016), Use of genetically encoded, light-gated ion translocators to control tumorigenesis, Oncotarget, 7(15): 19575-19588
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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), Bioelectric signaling via potassium channels: a mechanism for craniofacial dysmorphogenesis in KCNJ2-associated Andersen-Tawil Syndrome, Journal of Physiology, 594(12): 3245-3270
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Li, C., Levin, M., and Kaplan, D. (2016), Bioelectric modulation of macrophage polarization, Scientific Reports, 6: 21044
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Sundelacruz, S., Levin M., and Kaplan, D. L. (2015), Comparison of the depolarization response of human mesenchymal stem cells from different donors, Scientific Reports, 5: 18279
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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), Gap junctional blockade stochastically induces different species-specific head anatomies in genetically wild-type Girardia dorotocephala flatworms, International Journal of Molecular Sciences, 16(11): 27865-27896
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Pezzulo, G., and Levin, M. (2015), Re-membering the body: applications of computational neuroscience to the top-down control of regeneration of limbs and other complex organs, Integrative Biology, 7(12): 1487-1517
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Lobikin, M., Lobo, D., Blackiston, D. J., Martyniuk, C. J., Tkachenko, E., and Levin, M. (2015), Serotonergic regulation of melanocyte conversion: A bioelectrically regulated network for stochastic all-or-none hyperpigmentation, Science Signaling, 8(397): ra99
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Law, R., and Levin, M. (2015), Bioelectric memory: modeling resting potential bistability in amphibian embryos and mammalian cells, Theoretical Biology and Medical Modelling, 12(1): 22
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Schatzberg, D., Lawton, M., Hadyniak, S. E., Ross E. J., Carney, T., Beane, W. S., Levin, M., and Bradham, C. A. (2015), H+/K+-ATPase activity is required for biomineralization in sea urchin embryos, Developmental Biology, 406(2): 259-270
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Pai, V. P., Lemire, J. M., Chen, Y., Lin, G., and Levin, M. (2015), Local and long-range endogenous resting potential gradients antagonistically regulate apoptosis and proliferation in the embryonic CNS, International Journal of Developmental Biology, 59(7-9): 327-340
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Lobikin, M., Paré, J-F., Kaplan, D. L., and Levin, M. (2015), Selective Depolarization Of Transmembrane Potential Alters Muscle Patterning And Muscle Cell Localization In Embryonic Xenopus Laevis, International Journal of Developmental Biology, 59(7-9): 303-311
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Pai, V., Lemire, J. M., Paré, J-F., Lin, G., Chen, Y., and Levin, M. (2015), Endogenous gradients of resting potential instructively pattern embryonic neural tissue via Notch signaling and regulation of proliferation, Journal of Neuroscience, 35(10): 4366-4385
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Blackiston, D., Anderson, G. M., Rahman, N., Bieck, C., and Levin, M. (2015), A novel method for inducing nerve growth via modulation of host resting potential, Neurotherapeutics, 12(1): 170-184
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Chernet, B. T., Fields, C., and Levin, M. (2015), Long-range gap junctional signaling controls oncogene-mediated tumorigenesis in Xenopus laevis embryos, Frontiers in Physiology, 5: 519
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Özkucur, N., Quinn, K. P., Pang, J. C., Du, C., Georgakoudi, I., Miller, E., Levin, M., and Kaplan, D. L. (2015), Membrane potential depolarization causes alterations in neuron arrangement and connectivity in cocultures, Brain and Behavior, 5(1): 24-38
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Levin, M. (2014), Molecular bioelectricity: how endogenous voltage potentials control cell behavior and instruct pattern regulation in vivo, Molecular Biology of the Cell, 25(24): 3835-3850
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Adams, D. S., Lemire, J. M., Kramer, R. H., and Levin, M. (2014), Optogenetics in developmental biology: using light to control ion flux-dependent signals in Xenopus embryos, International Journal of Developmental Biology, 58(10-12): 851-861
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Lan, J. Y., Williams, C., Levin, M., and Black, L. D. III (2014), Depolarization of Cellular Resting Membrane Potential Promotes Neonatal Cardiomyocyte Proliferation In Vitro, Cellular and Molecular Bioengineering, 7(3): 432-445
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Mustard, J., and Levin, M. (2014), Bioelectrical mechanisms for programming growth and form: taming physiological networks for soft body robotics, Soft Robotics, 1(3): 169-191
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Chernet, B., and Levin, M. (2014), Transmembrane voltage potential of somatic cells controls oncogene-mediated tumorigenesis at long-range, Oncotarget, 5(10): 3287-3306
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Hernandez-Diaz, S., and Levin, M. (2014), Alteration of bioelectrically-controlled processes in the embryo: a teratogenic mechanism for anticonvulsants, Reproductive Toxicology, 47: 111-114
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Levin, M. (2014), Endogenous bioelectrical networks store non-genetic patterning information during development and regeneration, Journal of Physiology, 592(11): 2295-2305
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Pai, V., and Levin, M. (2014), Bioelectric controls of stem cell function, Chapter 5 in F. Calegari and C. Waskow (Eds.), Stem Cells: From Basic Research to Therapy, Volume 1, CRC Press: Boca Raton, FL, pp. 106-148
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Lobikin, M., and Levin, M. (2014), Endogenous bioelectric cues as morphogenetic signals in vivo, Chapter 15 in D. Fels, M. Cifra, and F. Scholkmann (Eds), Fields of the Cell, Research Signpost: Kerala, India, pp. 283-302
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Lange, C., Prenninger, S., Knuckles, P., Taylor, V., Levin, M., and Calegari, F. (2014), V-ATPase maintains neural stem cells in the developing mouse cortex, in S. Nakamuri (Ed.), Handbook of H+ ATPases, Pan Stanford Pte. Ltd: Singapore, pp. 413-430
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Sundelacruz, S., Li, C., Choi, Y. J., Levin, M., and Kaplan, D. L. (2013), Bioelectric modulation of wound healing in a 3D in vitro model of tissue-engineered bone, Biomaterials, 34(28): 6695-705
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Sundelacruz, S., Levin, M., and Kaplan, D. L. (2013), Depolarization Alters Phenotype, Maintains Plasticity of Predifferentiated Mesenchymal Stem Cells,Tissue Engineering Part A, 19(17-18): 1889-1908
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Chernet, B., and Levin, M. (2013), Endogenous voltage potentials and the microenvironment: bioelectric signals that reveal, induce, and normalize cancer, Journal of Clinical and Experimental Oncology, S1: doi:10.4172/2324-9110.S1-002
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Levin, M. (2013), Reprogramming cells and tissue patterning via bioelectrical pathways: molecular mechanisms and biomedical opportunities, Wiley Interdisciplinary Reviews: Systems Biology and Medicine, 5(6): 657-676
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Levin, M. (2013), Bioelectrical signaling has rich history, Physics Today, 66(9): 11
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Adams, D. S., and Levin, M. (2013), Endogenous Voltage Gradients as Mediators of Cell-Cell Communication: Strategies for Investigating Bioelectrical Signals During Pattern Formation, Cell and Tissue Research, 352(1):95-122
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Chernet, B. T., and Levin, M. (2013), Transmembrane voltage potential is an essential cellular parameter for the detection and control of tumor development in a Xenopus model, Disease Models & Mechanisms, 6(3): 595-607
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Beane, W. S., Morokuma, J., Lemire, J. M., and Levin, M. (2013), Bioelectric signaling regulates head and organ size during planarian regeneration, Development, 140(2): 313-322
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Tseng, A-S., and Levin, M. (2013), Cracking the bioelectric code: probing endogenous ionic controls of pattern formation, Communicative & Integrative Biology, 6(1): e22595
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Adams, D. S., Tseng, A-S. and Levin, M. (2013), Light-activation of the Archaerhodopsin H+ pump reverses age-dependent loss of vertebrate regeneration: sparking system-level controls in vivo, Biology Open, 2(3): 306-313
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Pai, V. P., Vandenberg, L. N., Blackiston, D. J., and Levin, M. (2012), Neurally derived tissues in Xenopus laevis embryos exhibit a consistent bioelectrical left-right asymmetry, Stem Cells International, 2012: 353491
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Lobikin, M., Chernet, B., Lobo, D., and Levin, M. (2012), Resting potential, oncogene-induced tumorigenesis, and metastasis: the bioelectric basis of cancer in vivo, Physical Biology, 9(6): 065002
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Tseng, A.-S., and Levin, M. (2012), Transducing bioelectrical signals into epigenetic pathways during tadpole tail regeneration, Anatomical Record, 295(10): 1541-1551
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Levin, M., and Stevenson, C. (2012), Regulation of Cell Behavior and Tissue Patterning by Bioelectrical Signals: challenges and opportunities for biomedical engineering, Annual Reviews in Biomedical Engineering, 14: 295-323
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Adams, D. S., and Levin, M. (2012), General principles for measuring resting membrane potential and ion concentration using fluorescent bioelectricity reporters, Cold Spring Harbor Protocols, 2012(4): 385-397
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Adams, D. S., and M. Levin (2012), Measuring resting membrane potential using the fluorescent voltage reporters DiBAC4(3) and CC2-DMPE, CSHL Protocols, 2012(4): 459-464
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Levin, M. (2012), Molecular bioelectricity in developmental biology: new tools and recent discoveries, BioEssays, 34(3): 205-217
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Pai, V., Aw, S., Shomrat, T., Lemire, J. M., and Levin, M. (2012), Transmembrane voltage potential controls embryonic eye patterning in Xenopus laevis, Development, 139(2): 313-323
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Blackiston, D., Adams, D. S., Lemire, J. M., Lobikin, M., and Levin, M. (2011), Transmembrane potential of GlyCl-expressing instructor cells induces a neoplastic-like conversion of melanocytes via a serotonergic pathway, Disease Models and Mechanisms, 4(1): 67-85
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Beane, W. S., Morokuma, J., Adams, D. S., and Levin, M. (2011), A chemical genetics approach reveals H,K-ATPase-mediated membrane voltage is required for planarian head regeneration, Chemistry & Biology, 18(1): 77-89
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Lange, C., Prenninger, S., Knuckles, P., Taylor, V., Levin, M., and Calegari, F. (2011), The H(+) vacuolar ATPase maintains neural stem cells in the developing mouse cortex, Stem Cells and Development, 20(5): 843-850
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Levin, M. (2011), Endogenous Bioelectric Signals as Morphogenetic Controls of Development, Regeneration, and Neoplasm, in C. Pullar (Ed.), The Physiology of Bioelectricity in Development, Tissue Regeneration, and Cancer, CRC Press: Boca Raton, FL, pp. 39-89
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Levin, M. (2011), The wisdom of the body: future techniques and approaches to morphogenetic fields in regenerative medicine, developmental biology, and cancer, Regenerative Medicine, 6(6): 667-673
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Aw, S., Koster, J., Pearson, W., Nicols, C., Shi, N. Q., Carneiro, K., and Levin, M. (2010), The ATP-sensitive K+-channel (KATP) controls early left-right patterning in Xenopus and chick embryos, Developmental Biology, 346(1): 39-53
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Tseng, A-S., Beane, W. S., Lemire, J. M., Masi, A., and Levin, M. (2010), Induction of vertebrate regeneration by a transient sodium current, Journal of Neuroscience, 30(39): 13192-13200
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Zhang, Y., and Levin, M. (2009), Particle tracking model of electrophoretic morphogen movement reveals stochastic dynamics of embryonic gradient, Developmental Dynamics, 238(8): 1923-1935
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Levin, M. (2009), Bioelectric mechanisms in regeneration: Unique aspects and future perspectives, Seminars in Cell and Developmental Biology, 20(5): 543-556
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Sundelacruz, S., Levin M., and Kaplan, D. L. (2009), Role of membrane potential in the regulation of cell proliferation and differentiation, Stem Cell Reviews, 5(3): 231-246
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Blackiston, D. J., McLaughlin, K., and Levin, M. (2009), Bioelectric controls of cell proliferation: ion channels, membrane voltage, and the cell cycle, Cell Cycle, 8(21): 3527-3536
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Morokuma, J., Blackiston, D., and Levin, M. (2008), KCNQ1 and KCNE1 K+ channel components are involved in early left-right patterning in Xenopus embryos, Cellular Physiology and Biochemistry, 21(5-6): 357-372
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Morokuma, J., Blackiston, D., Adams, D. S., Seebohm, G., Trimmer, B., and Levin, M. (2008), Modulation of potassium channel confers a hyperproliferative invasive phenotype on embryonic stem cells, Proceedings of the National Academy of Sciences of the United States, 105(43): 16608-16613
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Sundelacruz, S., Levin, M., and Kaplan, D. L. (2008), Membrane potential controls adipogenic and osteogenic differentiation of mesenchymal stem cells, PLoS One, 3(11): e3737
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Oviedo, N. J., Nicolas, C. L., Adams, D. S., and Levin, M. (2008), Live imaging of planarian membrane potential using DiBAC4(3), Cold Spring Harbor Protocols, 3(10): pdb.prot5055
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Adams, D. S., Masi, A., and Levin, M. (2007), H+ Pump-dependent changes in membrane voltage are an early mechanism necessary and sufficient to induce Xenopus tail regeneration, Development, 134(7): 1323-1335
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Aw, S., Adams, D. S., Qiu, D., and Levin, M. (2007), H,K-ATPase protein localization and Kir4.1 function reveal concordance of three axes during early determination of left-right asymmetry, Mechanisms of Development, 125(3-4): 353-372
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Levin, M. (2007), Large-Scale Biophysics: Ion Flows and Regeneration, Trends in Cell Biology, 17(6): 261-270
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Hibino, T., Ishii, Y., Levin, M., and Nishino, A. (2006), Ion flow regulates left-right asymmetry in sea urchin development, Development Genes and Evolution, 216(5): 265-76
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Shimeld, S. M., and Levin, M. (2006), Evidence for the regulation of left-right asymmetry in Ciona intestinalis by ion flux, Developmental Dynamics, 235(6): 1543-1553
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Adams, D. S., Robinson, K. R., Fukumoto, T., Yuan, S., Albertson, R. C., Yelick, P., Kuo, L., McSweeney, M., and Levin, M. (2006), Early, H+-V-ATPase-dependent proton flux is necessary for consistent left-right patterning of non-mammalian vertebrates, Development, 133(9): 1657-1671
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Esser, A. T., Smith, K. C., Weaver, J. C., and Levin, M. (2006), Mathematical Model of Morphogen Electrophoresis through Gap Junctions, Developmental Dynamics, 235(8): 2144-2159
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Adams, D. S., and Levin, M. (2006), Inverse Drug Screens: a rapid and inexpensive method for implicating molecular targets, Genesis, 44(11): 530-540
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Adams, D., and Levin, M. (2006), Strategies and techniques for investigation of biophysical signals in patterning, in Analysis of Growth Factor signaling in Embryos, M. Whitman and A. K. Sater eds., pp. 177-264, Methods in Signal Transduction Series, CRC Press

Levin, M. (2003), Bioelectromagnetics in morphogenesis, Bioelectromagnetics, 24(5): 295-315
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Levin, M. (2003), Motor protein control of ion flux is an early step in embryonic Left-Right asymmetry, BioEssays, 25(10): 1002-1010
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Levin, M., Thorlin, T., Robinson, K., Nogi, T., and Mercola, M. (2002), Asymmetries in H+/K+-ATPase and cell membrane potentials comprise a very early step in left-right patterning, Cell, 111(1): 77-89
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Rutenberg, J., Cheng, S. M., and Levin, M. (2002), Early embryonic expression of ion channels and pumps in chick and Xenopus development, Developmental Dynamics, 225(4): 469-484
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Cheng, S. M., Chen, I., and Levin, M. (2002), Katp channel activity is required for hatching in Xenopus embryos, Developmental Dynamics, 225(4): 588-591
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Levin, M., (1999), Endogenous electromagnetic fields and radiations in regeneration, development, and neoplasm, Proceedings of the First World Congress on the Effects of Electricity and Magnetism in the Natural World, Madeira, Portugal

Levin, M., and Ernst, S. G. (1997), Applied DC magnetic fields cause alterations in the time of cell divisions and developmental abnormalities in early sea urchin embryos, Bioelectromagnetics, 18(3): 255-263
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Levin, M., and Ernst, S. G. (1995), Applied AC and DC Magnetic Fields Cause Alterations in the Mitotic Cycle of Early Sea Urchin Embryos, Bioelectromagnetics, 16(4): 231-240
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Levin, M. (1993), Current and potential applications of bioelectromagnetics in medicine, Subtle Energies & Energy Medicine Journal, 4(1): 77-87
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ORCID ID: 0000-0001-7292-8084