Bioelectricity

Electroceuticals: emerging applications beyond the nervous system and excitable tissues
Balasubramanian, S., Weston, D., Levin, M., Davidian, D. C. C. (2024)
Cell Press, 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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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    

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

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

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

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

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

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

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

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

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

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

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

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

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: An Ion Channel Database with Small Molecule Modulators
Churchill, C. D. M., Winter, P., Tuszynski, J. A., and Levin, M. (2019)
iScience, 11: 42-56

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

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

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

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

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

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

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

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

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

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

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

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, p. 66-76

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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