Classic understanding of membrane potential as due to electrogenic ion pumping is untenable. In the new Wet-and-Murburn- Membrane-Model (WMMM) the sub-threshold R-state is the high-energy low-entropy excited coherent state and all processes (hyperpolarization, rest, generator potentials) are equilibrium-based. A phase transition occurs upon surpassing threshold, whereby the energy stored in the R-state is transformed into an all-or-none propagating action potential, a soliton. Only during the refractive period metabolic energy is introduced into the system by murburn redox processes directly and indirectly through adsorption of ATP produced by murburn phosphorylation. All processes are primarily due to electron displacements in redoxes and Lewis-induction. Ion currents are secondary, local, and do not propagate longitudinally along the axon. Permeability changes of the membrane upon surpassing threshold are due to changes in the structure of water. The process of changing low-energetic non-polarized water into highly excited polarized multi-layer water with a fourth phase 3D-structure depends on the combination of ATP adsorption energy and murburn redox activity. The fourth phase structure can extract energy from the quantum vacuum (aether). After Thomson and Bourassa corrected a mistake in the mainstream concept of charge, they could derive a physical description of the aether as a rotating magnetic field. Hereby aether units can house elementary particles in regions where there is matter. When electrons are highly excited and highly ordered (as in fourth phase water) energy and information from the aether field can be tapped. The Aether Protoplasm Interaction Proposal (APIP) suggests that the transformation of fourth phase water into non-polarized/de-structured water (rest-to-action transition) goes along with lowered interaction with the aether. This is what is observed in MRI. Biomed Rev 2025; 36: 85-107
Jaeken L. Wet-and-Murburn-Membrane-Model. Part I: Solute distribution, transport, volume control and redox metabolism. Biomed. Rev. 2025; 36: this issue
Ling GN (2001) Life at the Cell and Below-Cell Level. The Hidden History of a Fundamental Revolution in Biology. Pacific Press, New York.
Ling GN (2007) Nano-protoplasm: the ultimate unit of life. Physiol. Chem. Phys. & Med. NMR 39: 111-234.
Manoj KM (2018) Aerobic respiration: criticism of the proton-centric explanation involving rotary adenosine triphosphate synthesis, chemiosmosis principle, proton pumps and electron transport chain. Biochem. Insights 11: 1178626418818442.
Manoj KM, Jaeken L, Bazhin NM, Tamagawa H, Kavdia M, Manekkathodi A (2023) Murburn concept in cellular function and bioenergetics, Part 1: Understanding mur- zymes at the cellular level. AIP Advances 13: 120702.
Manoj KM, Jaeken L, Bazhin NM, Tamagawa H, Gideon DA, Kavdia M (2023) Murburn concept in cel- lular function and bioenergetics, Part 2: Understanding integrations-translations from molecular to macroscopic levels. AIP Advances 13: 120701.
Manoj KM, Jaeken L, Gideon DA, Tamagawa H, Kavdia
M. (2026) Structure and mechanism of cellular cation- transporters: Affinity-binding and murburn models. Int. J. Biol. Macromol. 2026. https://doi.org/10.1016/ ju.ijbiomac.2026.1506.150614.
Jaeken L, Manoj KM (2025) Murburn bioenergetics and “origins-sustenance-termination-evolution of life”: Emergence of intelligence from a network of molecules, unbound ions, radicals and radiations. Int. J. Mol. Sci. 26(15): 7542.
Matveev VV (2019) Cell theory, intrinsically disordered proteins, and the physics of the origin of life. Progr. Bio- phys. Mol. Biol. 149: 114-130.
Ling GN (1978) Maintenance of low sodium and high potassium levels in resting muscle cells. J. Physiol. 280(1): 105-123.
Ho MW (2008) The Rainbow and the Worm. The Physics of Organisms. (3rd ed.) World Scientific, Singapore.
Ho MW (2012) Super-conducting liquid crystalline water aligned with collagen fibres in the fascia as acupuncture meridians of traditional Chinese medicine. Forum Im- mun. Dis. & Therap. 3(3-4): 221-230.
Hodgkin AL, Huxley AF, Katz B (1952) Measurement of current-voltage relations in the membrane of the giant axon of Loligo. J. Physiol. (London) 116: 424-434
Hodgkin AL, Huxley AF (1952) A quantitative description of membrane current and its application to conduction and excitation in nerve. J. Physiol. London 117: 500-544.
Hodgkin AL, Katz B (1949) The effect of sodium ions on the electrical activity of the giant axon of the squid. J. Physiol. 108: 37-37.
Goldman DE (1943) Potential, impedance and rectifica- tion in membranes. J. Gen. Physiol. 27: 37-60
Delalande B, Tamagawa H, Matveev V (2020) From Nernst to Bernstein and beyond. Open Access Library
J. 7: e7047. (update, not peer reviewed: Delalande B, Tamagawa H, Matveev V (2025) The Goldman Hodgkin and Katz Errors).
Mosgaard LD, Zecchi KA, Heimburg T (2015) Mechano- capacitive properties of polarized membranes. Soft Mat- ter, 11: 7899.
Ling GN (1982) The cellular resting and action potential: interpretation based on the AI hypothesis. Physiol. Chem. Phys. 14: 47-96.
Pollack GH (2013) The Fourth Phase of Water: Beyond Solid, Liquid and Vapor. Ebner & Sons, Seattle.
Heimburg T (2022) The thermodynamic soliton theory of the nerve impulse and possible medical implications. Progr. Biophys. Mol. Biol. 173: 24-35.
Mitchell P (1961) Coupling of phosphorylation to elec- tron and hydrogen transfer by a chemiosmotic type of mechanism. Nature 191: 144-148.
Hodgkin AL, Horowicz P (1959) The influence of potas- sium and chloride on the membrane potential of single muscle fibers. J. Physiol. 148: 127-160.
Ritchie JM, Keynes RD (1985) The production and absorption of heat associated with electrical activity in nerve and electric organ. Q. Rev. Biophys. 18: 451-476.
Cope FW (1965) Kinetic criteria of solid-state mecha- nisms in biology. Bull. Math. Biophys. 27: 113- 118.
Cope FW (1968) Evidence for semiconduction in Aplysia nerve membrane. Proc. Natl. Acad. Sci. USA 61: 905- 908.
Cope FW (1977) Solid state physical theory of nerve im- pulse conduction: Elovich kinetics of axon repolarisation currents. Physiol. Chem. Phys. 9: 383-387.
Cope FW (1976) Solid state theory of competitive diffu- sion of associated Na+ and K+ in cells by free cation and vacancy (hole) mechanisms, with application to nerve. Physiol. Chem. Phys. 9: 389-398.
Cope FW (1977) Solid state physical replacement of Hodgkin-Huxley theory. Phase transformation kinetics of axonal potassium conductance? Physiol. Chem. Phys. 9: 155-160.
Cope FW (1977) Solid state physical theory of nerve im- pulse conduction: Elovich kinetics of axon repolarisation currents. Physiol. Chem. Phys. 9: 383-387.
Cope FW (1973) Electron-phonon (trapped photon) coupling and infrared coaxial transmission line theory of energy transport in mitochondria and nerve. Bull. Math. Biol. 35: 627-644.
Cope FW (1975) A Review of the Applications of Solid- State Physics Concepts to Biological Systems. J. Biol. Phys. 3(1): 1-41.
Davydov AS (1973) The theory of soliton contraction of proteins under their excitation. J. theor. Biol. 38: 559-569.
Manoj KM, Tamagawa H, Bazhin N, Jaeken L, Nirusim- han V, Faraci F, Gideon DA (2022) Murburn model of vision, precepts and proof of concept. J. Cell. Physiol. 237(8): 3338-3355.
Manoj KM, Gideon DA, Bazhin N, Tamagawa H, Nirusimhan, V, Kavdia M, Jaeken L (2023). Na,K- ATPase: A murzyme facilitating thermodynamic equilib- riums at the membrane interface. J. Cell. Physiol. 2023, 238: 109-136.
Edelmann L (1973) The influence of rubidium and cesium ions on the resting potential of guinea pig heart muscle cells as predicted by the association-induction hypothesis. Ann. N. Y. Acad. Sci. 204(1): 534-537.
Ling GN (1959) On the mechanism of cell potential. Fed. Proc. 18: 371.
Thoke HS, Bagatolli LA, Olsen LF (2018) Effect of macromolecular crowding on the kinetics of glycolytic enzymes and the behaviour of glycolysis in yeast. Integr. Biol. (Camb) 10(10): 587-597.
Tamagawa H (2019) Mathematical expression of mem- brane potential based on Ling’s adsorption theory is approximately the same as the Goldman-Hodgkin-Katz equation. J. Biol. Phys. 45(1): 13-30.
Ling GN (1964) The role of inductive effect in coopera- tive phenomena of proteins.
Ling GN (1984) In Search of the Physical Basis of Life.
Plenum Press, New York.
Zheng JM, Pollack GH (2003) Long-range forces extend- ing from polymer-gel surfaces. Phys. Rev. E, 68: 031408.
Yoo H; Baker DR, Pirie CM, Hovakeemoan B, Pollack GH (2011) Characteristics of water adjacent to hydro- philic interfaces. In: Water: The Forgotten Biological Molecule. Le Bihan D, Fukuyama H., eds. Pan Stanford Publishing Pte. Ltd., pp. 123-136.
Bunkin NF, Kozlov VA, Ignatiev PS, Suyazov NV, Shki- rin AV, Alakhodzhaev IA (2012). Refractiv coefficient of water and aqueous solutions in the optical frequency range in the vicinity of Nafion. Biophysics 57: 733-749.
Zhao Q, Ovchinnikova K, Chai B, Yoo H, Magula J, Pollack GH (2009) Role of proton gradients in the mecha- nism of osmosis. J. Phys. Chem. B, 113: 10708-10714.
Ovchinnikova K, Pollack GH (2009) Can water store charge? Langmuir 25: 542-547.
Chai B, Yoo H, Pollack GH (2009) Effect of radiant energy on near-surface water. J. Phys. Chem. B, 113: 13953-13958.
Voeikov VL, Del Giudice E (2009) Water respiration – the basis of the living state. Water 1: 52-73. Doi: 10.14294/ WATER.2009.4.
Del Giudice E, Doglia S, Milani M and Vitiello G (1986) Collective properties of biological systems: solitons and coherent electric waves in a quantum field theoretical approach. In: Modern Bioelectrochemistry. Gutmann F and Keyzer H., eds., Plenum, New York, pp. 263-288.
Del Giudice E, Elia V, Tedeschi A (2009) Role of water in the living organisms. Neural netw. World 19(4): 355- 360.
Del Giudice E, Spinetti PR and Tedeschi A (2010) Water Dynamics at the Root of Metamorphosis in Living Organ- isms. Water 2: 566-586.
Del Giudice E, Voeikov V, Tedeshi A and Vitiello G (2015) The origin and the special role of coherent water in living systems. In: Fields of the Cell. Fels D, Cifra M and Scholkmann F, eds., Research Signpost 37/661(2), Fort P.O., Trivandrum, Kerala, India, pp. 95-111.
King MB (1989) Tapping the Zero-Point Energy. Para- clete Publishing. Provo, Utah.
Chatzidimitriou-Dreismann, C. A., Abdul Redah, T., Streffer, R. M. F. and Mayers, J. (1997) Anomalous deep inelastic neutron scattering from liquid H2O-D2O: evidence of nuclear quantum entanglement. Phys. Rev. Lett. 79(15): 2839-2842.
Schurr JM (2013) Phenomena associated with gel-water interfaces. Analyses and alternatives to the long-range water hypothesis. J. Phys. Chem. B 117(25): 7653-7674.
Elton, D. C., Spencer, P. D., Riches, J. D. Williams, E. D. (2020) Exclusion Zone Phenomena in Water—A Critical Review of Experimental Findings and Theories. Intern. J. Mol. Sci. 21: 5041.
Huang SG, Ki Hong J, Sharma A, Pollack GH, Bahrig GW (2018) Exclusion zone and heterogeneous water structure at ambient temperature. PLoS ONE 13(4): e0195057.
Jaeken L (2026) Wonderful oxygen, wonderful (cell) wa- ter, and their wonderful equilibrium: a new bioenergetic foundation for cell physiology. In: Recent Advances in Bioelectromagnetism and subtle energies. Implications for health and Disease. Martel J, ed., Acad. Press, Lon- don, Cambridge MA, pp. 55-83.
Chaplin M (2006). Do we underestimate the importance of water in cell biology? Nat. Rev. Mol. Cell Biol. 7(11):861-866.
Pauling L (1935) The structure and entropy of ice and of other crystals with some randomness of atomic arrange- ment. J. Am. Chem. Soc. 57: 2680-2684.
Kanarev PM (2003, and updates online) The Foundations of Physchemistry of Microworld. 10th ed. (in English) http://guns.connect.fi/innoplaza/energy/story/Kanarev
Jaeken L (2021) Worries of a bio-scientist: Do we need a new atomic model? In: New Insights into Physical Science. Vol. 5, Tüzemen S, ed., Book Publisher Inter- national, India, UK, pp. 97-113.
Trevors JT, Pollack GH (2012) Origin of microbial life hypothesis: a gel cytoplasm lacking a bilayer membrane, with infrared radiation producing exclusion zone (EZ) water, hydrogen as an energy source and thermosynthesis for bioenergetics. Biochimie 94(1): 258-262.
Kell DB (1979) On the functional proton current pathway of electron transport phosphorylation: an electrodic view. Biochim. Biophys. Acta 549: 55-99.
Nanninga LB and Mommaerts WFHM (1957) On the binding of adenosine triphosphate by actomyosin. Proc. Natl. Acad. Sci. USA 43(6): 540-542.
Asakura S (1961) The interaction of G-actin and ATP. Arch. Biochem. Biophys. 92(1): 140-149.
Kinnunen PKJ, Virtanen JA (1986) A qualitative molecu- lar model of the nerve impulse. Conductive properties of unsaturated lyotropic liquid crystals. In: Modern Bioel- ectrochemistry. Gutmann F, Keyzer H, eds., chapter 17, pp. 457-480. Plenum, New York.
Schmidt D, Jiang QX, MacKinnon R (2006) Phospho- lipids and the origin of cationic gating charges in voltage sensors. Nature 444/ 775-779.
Jiang QX (2019) Cholesterol dependent gating effects on ion channels. Adv. Exp. Med. Biol. 1115: 167-190.
Scott AC (1995) Solitary waves in biology. In: Nonlinear Excitations in Biomolecules. Peyrard M, ed. Springer, Berlin, pp. 249-268.
Damadian R (1971) Tumor detection by nuclear magnetic resonance. Science 171: 1151-1153.
Plankar M, Jerman I and Krasovec R (2011) On the origin of cancer: can we ignore coherence? Progr. Biophys. Mol. Biol. 106(2): 380-390.
Popp FA, Chang JJ (1998) The physical background and the informational character of biophoton emission. In: Biophotons. Chang JJ, Fisch J, Popp FA, eds. Kluwer Academic, Dordrecht, The Netherlands, pp.239-250.
Drukarch B, Micha MMW, Shrivastava S. (2021) The thermodynamic theory of action potential propagation: a sound basis for unification of the physics of nerve im- pulses. Rev. Neurosci. 33(3): 285-302.
Einstein A (1950) Out of my later years. Philosophical library, p. 54.
Fröhlich H (1984) General theory of coherent excitations in biological systems. In: Nonlinear Electrodynamics in Biological Systems. Adey WR and Lawrence AF; eds., Plenum, New York, pp. 491-496.
Fröhlich H (1986) Coherent excitations in active biologi- cal systems. In: Modern Bioelectrochemistry. Gutmann F and Keyzer H., eds., Plenum, New York, pp. 246-261.
Jaeken L (2021) The Coacervate-Coherence Nature of Life. Fundamentals of Cell Physiology. BioMedES, Inverurie, UK (distributed by Amazon).
Luckey M (2014) Membrane Structural Biology. 2nd ed. Cambridge University Press, New York.
Heimburg T (2025) The mechanical properties of nerves, the size of action potential, and consequences for the brain. Chem. Physics. Lipids 267: 105461. Doi: 10.1016/j.chemphyslip.2024.105461.
Heimburg T (2009) Die Physik der Nerven. Physik Jour- nal 8(3): 33-39 (translation: The physics of nerves. arXiv: 1008.4279v. (25 Aug 2010).
Nasonov DN (1962) Local Reaction of Protoplasm and Gradual Excitation. (English translation by Halpern YS), National Science Foundation, available at Office of Technical Services, Department of Commerce, US: Washington DC.
Iwasa K, Tasaki I, Gibbons RC (1980) Swelling of nerve fibers associated with action potentials. Science 210: 338-339. Doi: 10.1126/science.7423196.
Tasaki I (1999) Evidence for phase transition in nerve fibers, cells and synapses. Ferroelectrics 220: 305-316.
Heimburg T and Jackson AD (2005) On soliton propaga- tion in biomembranes and nerves. Proc. Natl. Acad. Sci. USA 102(28): 9790-9795.
Shrivastava S, Kang K, Schneider MF (2015) Solitary shock waves and adiabatic phase transitions in lipid interfaces and nerves. Phys. Rev. E, 91: 012715.
Ishima Y, Przybylski AT, Fox SW (1981) Spontaneous membrane phenomena in spherules from proteoid and lecithin. Biosystems, 13(4): 243-251.
Heimburg T (2010) Lipid ion channels. Biophys. Chem. 150(1-3): 2-22. Doi: 10.1016/j.bpc.2010.02.018.
Blicher A, Heimburg T (2013) Voltage-gated lipid ion channels. PLOS 0ne !(6): e65707.
Heimburg T (2014) Nerves and anesthesia: a physics per- spective on medicine. arXiv: 1409.2430v1 (8 Sep 2014).
Cole KS (1941) Rectification and inductance in the squid giant axon. J. Gen. Physiol. 25: 29-51.
Delalande B, Tamagawa H, Matveev V. (2026) Be- yond Hodgkin-Huxley – The ionic-mechano-hydraulic (IMH) model of nerve conduction. Doi: 10.20944/pre- prints202603.0725.v1.
Bagatolli LA, Ipsen JH, Simonsen AC, Mouritsen OG (2010) An outlook on organization of lipids in mem- branes; Searching for a realistic connection with the organization of biological membranes. Prog. Lipid Res. 49(4): 378-389.
Kabir SR, Yokoyama K, Minasi K, Kodama T, Suzuki M (2003) Hyper-mobile water is induced around actin filaments. Biophys. J. 85(5): 3154-3164.
Wheatley DN, Robertson JH, Giddings mR (1979) Amino acid analogues: uptake, pool formation and incorpora- tion of phenylalanine and two halogenated derivatives in cultured mammalian cells. Cytobios 24(94): 75-98.
Ling GN (1973) What component of the living cell is responsible for its semipermeable properties: polarized water or lipids? Biophys. J. 13:807.
Wheatley DN, Redfern A and Johnson RPC (1990) Ef- fects of osmotic manipulation of intracellular hydration of HeLa S-3 cells on their proton NMR relaxation times. Magn. Reson. Imaging 8(3): 285-293.
Jerman I (2016) The origin of life from quantum vacuum, water and polar molecules. Am. J; Modern Phys. 5(4-1): 34-43.
Thomson DW, Bourassa JD (2004) Secrets of the Aether: Unified Force Theory, Dark Matter and Consciousness. The Quantum Aether Dynamics Institute. (Short update, 2012: A New Foundation for Physics).
Voeikov VL (2007) Fundamental role of water in bioener- getics. In: Biophotonics and Coherent Systems in Biology. Voeikov VL, Martynyuk VS, eds. Springer, New York, pp. 89-104.
Rao ML, Sedlmayer SR, Roy R, Kanzius J. Polarized micro- wave and RF radiation effects on the structure and stability of liquid water. Current Sci 2010; 98(11): 1500-1504.
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