{"id":527681,"date":"2026-07-01T14:01:38","date_gmt":"2026-07-01T14:01:38","guid":{"rendered":"https:\/\/www.newsbeep.com\/ie\/527681\/"},"modified":"2026-07-01T14:01:38","modified_gmt":"2026-07-01T14:01:38","slug":"the-chemistry-of-habitable-oceans-in-the-solar-system","status":"publish","type":"post","link":"https:\/\/www.newsbeep.com\/ie\/527681\/","title":{"rendered":"The chemistry of habitable oceans in the Solar system"},"content":{"rendered":"<p>Persistent liquid water is generally considered a prerequisite for planetary habitability<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 1\" title=\"Cockell, C. S. et al. Habitability: a review. Astrobiology 16, 89&#x2013;117 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR1\" id=\"ref-link-section-d198960051e798\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 2\" title=\"National Academies of Sciences and Medicine, E. Origins, Worlds, and Life: A Decadal Strategy for Planetary Science and Astrobiology 2023&#x2013;2032. (The National Academies Press, Washington, 2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR2\" id=\"ref-link-section-d198960051e801\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>. For all known life, which is to say life on Earth, water is the primary component of cells and serves as an indispensable matrix for biochemical processes, facilitating molecular transport, solvation, and reactivity<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 3\" title=\"Ball, P. Water as an active constituent in cell biology. Chem. Rev. 108, 74&#x2013;108 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR3\" id=\"ref-link-section-d198960051e805\" rel=\"nofollow noopener\" target=\"_blank\">3<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 4\" title=\"Ball, P. Water is an active matrix of life for cell and molecular biology. Proc. Natl. Acad. Sci. USA 114, 13327&#x2013;13335 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR4\" id=\"ref-link-section-d198960051e808\" rel=\"nofollow noopener\" target=\"_blank\">4<\/a>. On a macroscopic scale, the planetary water cycle helps maintain temperate climates, distributes substantial heat from equator to poles<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 5\" title=\"Pierrehumbert, R. T. Principles of Planetary Climate. (Cambridge Univ. Press, Cambridge, 2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR5\" id=\"ref-link-section-d198960051e812\" rel=\"nofollow noopener\" target=\"_blank\">5<\/a>, and drives the global biogeochemical cycling of bio-essential elements<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 6\" title=\"Berner, E. K. &amp; Berner, R. A. Global Environment: Water, Air, and Geochemical Cycles. (Princeton Univ. Pcess, Princeton, 2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR6\" id=\"ref-link-section-d198960051e816\" rel=\"nofollow noopener\" target=\"_blank\">6<\/a>. Water\u2019s possibly unique suitability for life stems from its exceptional physicochemical properties, including its potent polarity, high dielectric constant, and capacity for extensive hydrogen bonding<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 4\" title=\"Ball, P. Water is an active matrix of life for cell and molecular biology. Proc. Natl. Acad. Sci. USA 114, 13327&#x2013;13335 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR4\" id=\"ref-link-section-d198960051e820\" rel=\"nofollow noopener\" target=\"_blank\">4<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 7\" title=\"Chaplin, M. et al. Do we underestimate the importance of water in cell biology? Nat. Rev. Mol. Cell Biol. 7, 861&#x2013;866 (2006).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR7\" id=\"ref-link-section-d198960051e823\" rel=\"nofollow noopener\" target=\"_blank\">7<\/a>. While it has been suggested that under certain conditions of temperature and pressure, liquid ammonia and certain organic solvents might share water\u2019s bio-friendly attributes<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 8\" title=\"Benner, S. A., Ricardo, A. &amp; Carrigan, M. A. Is there a common chemical model for life in the universe? Curr. Opin. Chem. Biol. 8, 672&#x2013;689 (2004).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR8\" id=\"ref-link-section-d198960051e828\" rel=\"nofollow noopener\" target=\"_blank\">8<\/a>, such conditions are doubtful on the rocky planets and moons deemed as possible abodes for life in our solar system. Consequently, the search for extraterrestrial life remains focused on worlds that harbor liquid water (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>).<\/p>\n<p>Fig. 1: Evolution and distribution of potential ocean-bearing planetary bodies in the solar system.<img decoding=\"async\" aria-describedby=\"figure-1-desc\" src=\"https:\/\/www.newsbeep.com\/ie\/wp-content\/uploads\/2026\/07\/41467_2026_74955_Fig1_HTML.png\" alt=\"Fig. 1: Evolution and distribution of potential ocean-bearing planetary bodies in the solar system.\" loading=\"lazy\" width=\"685\" height=\"827\"\/><\/p>\n<p>The conventional habitable zone (HZ) is defined as the circumstellar region where Earth-like planets with CO2-H2O-dominated greenhouse atmospheres can sustain liquid water on their surfaces. In more optimistic formulations, the Solar System\u2019s HZ may extend from early Mars to recent Venus. Discoveries of subsurface liquid-water reservoirs and oceans in dwarf planets and icy moons of the outer Solar System have substantially broadened the concept of planetary habitability beyond the classical HZ. This extended view of habitability incorporates endogenous energy sources that can maintain liquid water in planetary interiors or subsurface reservoirs. Planetary bodies are not shown to scale.<\/p>\n<p>This emphasis on liquid water has strongly shaped the astronomical concept of the habitable zone (HZ), which is conventionally defined as the circumstellar region within which an Earth-like planet with the right atmospheric composition can sustain liquid water on the surface<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 9\" title=\"Huang, S. et al. Occurrence of Life in the Universe. Am. Sci. 47, 397&#x2013;402 (1959).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR9\" id=\"ref-link-section-d198960051e862\" rel=\"nofollow noopener\" target=\"_blank\">9<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 10\" title=\"Kasting, J. F., Kopparapu, R., Ramirez, R. M. &amp; Harman, C. E. Remote life-detection criteria, habitable zone boundaries, and the frequency of Earth-like planets around M and late K stars. Proc. Natl. Acad. Sci. USA 111, 12641&#x2013;12646 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR10\" id=\"ref-link-section-d198960051e865\" rel=\"nofollow noopener\" target=\"_blank\">10<\/a>. Commonly, the HZ boundaries are estimated as a function of stellar irradiance and orbital distance, assuming radiative balance between absorbed stellar flux and emitted thermal radiation, and a CO2-H2O greenhouse effect<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 11\" title=\"Kopparapu, R. K. et al. Habitable zones around main-sequence stars: new estimates. Astrophys. J. 765, 131 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR11\" id=\"ref-link-section-d198960051e873\" rel=\"nofollow noopener\" target=\"_blank\">11<\/a>. Under some optimistic formulations, the solar system\u2019s HZ may span from early Mars, which preserves abundant evidence for ancient surface water activities<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 12\" title=\"Wordsworth, R. D. The climate of early Mars. Annu. Rev. Earth Planet. Sci. 44, 381&#x2013;408 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR12\" id=\"ref-link-section-d198960051e877\" rel=\"nofollow noopener\" target=\"_blank\">12<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 13\" title=\"Haberle, R. M., Catling, D. C., Carr, M. H. &amp; Zahnle, K. J. The Early Mars Climate System. in The Atmosphere and Climate of Mars 526&#x2013;568. (Cambridge Univ. Press, Cambridge, 2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR13\" id=\"ref-link-section-d198960051e880\" rel=\"nofollow noopener\" target=\"_blank\">13<\/a>, to recent Venus, which assumes that Venus did not undergo a runaway transition to a hot climatic state until ~1 Ga (i.e., billion years) ago<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 14\" title=\"Yang, J., Bou&#xE9;, G., Fabrycky, D. C. &amp; Abbot, D. S. Strong dependence of the inner edge of the habitable zone on planetary rotation rate. Astrophys. J. 787, L2 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR14\" id=\"ref-link-section-d198960051e885\" rel=\"nofollow noopener\" target=\"_blank\">14<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 15\" title=\"Way, M. J. et al. Was Venus the first habitable world of our solar system? Geophys. Res. Lett. 43, 8376&#x2013;8383 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR15\" id=\"ref-link-section-d198960051e888\" rel=\"nofollow noopener\" target=\"_blank\">15<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>).<\/p>\n<p>However, a star-centric HZ definition overlooks the influence of planetary properties, including atmospheric and interior evolution, and some climate feedbacks. For example, recent models suggest that Venus may never have had a liquid-water ocean on its surface because of early climatic evolution and cloud feedbacks<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 16\" title=\"Turbet, M. et al. Day&#x2013;night cloud asymmetry prevents early oceans on Venus but not on Earth. Nature 598, 276&#x2013;280 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR16\" id=\"ref-link-section-d198960051e898\" rel=\"nofollow noopener\" target=\"_blank\">16<\/a>. Conversely, although modern Mars lies within the outer boundary of the conventional HZ, its low mass allowed for substantial atmospheric escape, leading to a tenuous atmosphere that cannot sustain persistent surface liquid water<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 13\" title=\"Haberle, R. M., Catling, D. C., Carr, M. H. &amp; Zahnle, K. J. The Early Mars Climate System. in The Atmosphere and Climate of Mars 526&#x2013;568. (Cambridge Univ. Press, Cambridge, 2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR13\" id=\"ref-link-section-d198960051e902\" rel=\"nofollow noopener\" target=\"_blank\">13<\/a>. Meanwhile, evidence for extant subsurface Martian aquifers remains elusive<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 17\" title=\"Luzzi, E. et al. Geomorphological evidence of near-surface ice at candidate landing sites in northern Amazonis Planitia, Mars. J. Geophys. Res. Planets 130, e2024JE008724 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR17\" id=\"ref-link-section-d198960051e906\" rel=\"nofollow noopener\" target=\"_blank\">17<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 18\" title=\"Morgan, G. A. et al. High frequency radar perspective of putative subglacial liquid water on Mars. Geophys. Res. Lett. 52, e2025GL118537 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR18\" id=\"ref-link-section-d198960051e909\" rel=\"nofollow noopener\" target=\"_blank\">18<\/a>. Furthermore, the conventional HZ concept neglects endogenous energy sources, such as tidal dissipation and radiogenic heating, which can maintain liquid water independently of stellar insolation. This limitation is underscored by growing evidence of subsurface liquid water bodies or oceans in icy bodies in the outer solar system, including dwarf planets like Ceres and Pluto and moons such as Europa, Ganymede, Callisto, Enceladus, and Titan<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 19\" title=\"Nimmo, F. &amp; Pappalardo, R. T. Ocean worlds in the outer solar system. J. Geophys. Res. Planets 121, 1378&#x2013;1399 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR19\" id=\"ref-link-section-d198960051e913\" rel=\"nofollow noopener\" target=\"_blank\">19<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Together, these discoveries have broadened the concept of planetary habitability beyond the classical HZ, in both space and time (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). This expanded perspective also extends to exoplanetary systems, where an increasing number of candidates may host surface oceans<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 20\" title=\"Hill, M. L. et al. A catalog of habitable zone exoplanets. Astron. J. 165, 34 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR20\" id=\"ref-link-section-d198960051e924\" rel=\"nofollow noopener\" target=\"_blank\">20<\/a>, potentially accompanied by ocean-bearing exomoons<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 21\" title=\"Kaltenegger, L. Characterizing habitable exomoons. Astrophys. J. Lett. 712, L125 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR21\" id=\"ref-link-section-d198960051e928\" rel=\"nofollow noopener\" target=\"_blank\">21<\/a>.<\/p>\n<p>However, life is not constituted by water alone; it requires a specific suite of bioessential elements dissolved within it. Beyond hydrogen and oxygen, a cell\u2019s core chemical formulae include carbon, nitrogen, phosphorus, and sulfur, alongside a range of minor and trace elements<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 22\" title=\"Rickaby, R. E. M. Goldilocks and the three inorganic equilibria: how Earth&#x2019;s chemistry and life coevolve to be nearly in tune. Philos. Trans. R. Soc. A 373, 20140188 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR22\" id=\"ref-link-section-d198960051e935\" rel=\"nofollow noopener\" target=\"_blank\">22<\/a>. The molar ratio of C:N:P in marine biomass on Earth, the Redfield ratio, is broadly consistent across diverse contemporary life forms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 23\" title=\"Redfield, A. C. The biological control of chemical factors in the environment. Am. Sci. 46, 230A&#x2013;221 (1958).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR23\" id=\"ref-link-section-d198960051e939\" rel=\"nofollow noopener\" target=\"_blank\">23<\/a>. Intriguingly, this ratio correlates with the elements\u2019 relative bioavailability in the modern ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 24\" title=\"Moore, C. M. et al. Processes and patterns of oceanic nutrient limitation. Nat. Geosci. 6, 701&#x2013;710 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR24\" id=\"ref-link-section-d198960051e943\" rel=\"nofollow noopener\" target=\"_blank\">24<\/a>, underscoring the critical roles of aquatic chemistry in supplying life\u2019s building blocks and life\u2019s ability to exert a strong control on elemental ratios within said aquatic chemistry. Consequently, the chemical composition of a planetary water reservoir is not only a critical factor in assessing its habitability<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 2\" title=\"National Academies of Sciences and Medicine, E. Origins, Worlds, and Life: A Decadal Strategy for Planetary Science and Astrobiology 2023&#x2013;2032. (The National Academies Press, Washington, 2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR2\" id=\"ref-link-section-d198960051e947\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a> but may also directly reflect the life within it. However, not all water is habitable; water activity (aw), controlled by the interactions between ions and dissolved organic molecules in aqueous solution, can inhibit cellular physiology when solutes reach high concentrations<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Tosca, N. J., Knoll, A. H. &amp; McLennan, S. M. Water activity and the challenge for life on early Mars. Science 320, 1204&#x2013;1207 (2008).\" href=\"#ref-CR25\" id=\"ref-link-section-d198960051e958\">25<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Stevens, A. H. &amp; Cockell, C. S. The water activity of mars-relevant multicomponent brines: The changing influence of perchlorate on habitability over time. Planet. Sci. J. 4, 6 (2023).\" href=\"#ref-CR26\" id=\"ref-link-section-d198960051e958_1\">26<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 27\" title=\"Stevenson, A. et al. Is there a common water-activity limit for the three domains of life? ISME J. 9, 1333&#x2013;1351 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR27\" id=\"ref-link-section-d198960051e961\" rel=\"nofollow noopener\" target=\"_blank\">27<\/a>.<\/p>\n<p>At steady state, planetary water chemistry is predominantly governed by water-rock interactions, processes that are highly sensitive to environmental conditions including temperature, pressure, pH, redox state, and rock composition. Throughout Earth\u2019s geological history, evolving surface conditions have driven profound changes in ocean chemistry, which both shaped and were shaped by the biosphere<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 28\" title=\"Anbar, A. D. Elements and evolution. Science 322, 1481&#x2013;1483 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR28\" id=\"ref-link-section-d198960051e969\" rel=\"nofollow noopener\" target=\"_blank\">28<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 29\" title=\"Moore, E. K., Jelen, B. I., Giovannelli, D., Raanan, H. &amp; Falkowski, P. G. Metal availability and the expanding network of microbial metabolisms in the Archaean eon. Nat. Geosci. 10, 629&#x2013;636 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR29\" id=\"ref-link-section-d198960051e972\" rel=\"nofollow noopener\" target=\"_blank\">29<\/a>. Other ocean worlds in our solar system, however, have followed vastly different evolutionary paths dictated by distinct accretion and orbital histories, tectonic regimes, impact fluxes, and atmospheric evolution<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 19\" title=\"Nimmo, F. &amp; Pappalardo, R. T. Ocean worlds in the outer solar system. J. Geophys. Res. Planets 121, 1378&#x2013;1399 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR19\" id=\"ref-link-section-d198960051e976\" rel=\"nofollow noopener\" target=\"_blank\">19<\/a>. Consequently, the chemical features and evolutionary histories of these extraterrestrial oceans likely differ significantly from Earth\u2019s (e.g., Xu et al<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 30\" title=\"Xu, W. et al. Enough sulfur and iron for potential life make Enceladus&#x2019;s ocean fully habitable. Astrophys. J. Lett. 980, L10 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR30\" id=\"ref-link-section-d198960051e980\" rel=\"nofollow noopener\" target=\"_blank\">30<\/a>.), with profound implications for their capacity to support life as we know it.<\/p>\n<p>In this review, we first synthesize the chemical features of water reservoirs and their biological implications on the modern Earth (Section 1). Based on this foundation, we then outline constraints on the evolutionary history of Earth\u2019s ocean chemistry and discuss its interplay with the biosphere (Section 2). Next, we evaluate geological evidence for aqueous chemistry on early Mars and discuss its consequences for habitability and biosignature preservation (Section 3). We then examine the inferred chemical environments of four representative ocean-bearing icy bodies, namely dwarf planet Ceres, Jupiter\u2019s moon Europa, and Saturn\u2019s moons Enceladus and Titan, and highlight current knowledge and key uncertainties (Section 4). Finally, we conclude by outlining open questions in planetary ocean chemistry that are paramount to address through future research to fully evaluate habitability across the solar system.<\/p>\n<p>Section 1. Water bodies on the modern earth: a framework for planetary habitability<\/p>\n<p>Earth is predominantly covered by liquid water (~71% of its surface), and ~96.5% of its total surface water inventory resides in the ocean, while only a minor proportion is distributed among saline lakes and freshwater reservoirs<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 6\" title=\"Berner, E. K. &amp; Berner, R. A. Global Environment: Water, Air, and Geochemical Cycles. (Princeton Univ. Pcess, Princeton, 2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR6\" id=\"ref-link-section-d198960051e1006\" rel=\"nofollow noopener\" target=\"_blank\">6<\/a>. These surface water bodies are integral components of the global hydrological cycle yet exhibit striking chemical diversity (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>), primarily governed by the extent of water-rock interaction and local hydrodynamic conditions. The hydrological cycle begins with rainfall. Rainwater is dilute, with only minor dissolved solids (typically ~10\u2009mg\/L), oxidizing, and weakly acidic (pH ~ 4\u20136), due to equilibration with atmospheric CO2 and O2<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 6\" title=\"Berner, E. K. &amp; Berner, R. A. Global Environment: Water, Air, and Geochemical Cycles. (Princeton Univ. Pcess, Princeton, 2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR6\" id=\"ref-link-section-d198960051e1016\" rel=\"nofollow noopener\" target=\"_blank\">6<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 31\" title=\"Stumm, W. &amp; Morgan, J. J. Aquatic Chemistry: Chemical Equilibria and Rates in Natural Waters. (John Wiley &amp; Sons, New York, 2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR31\" id=\"ref-link-section-d198960051e1019\" rel=\"nofollow noopener\" target=\"_blank\">31<\/a>. In surface or near-surface drainage, rain and river waters cause chemical weathering of terrestrial rocks<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 6\" title=\"Berner, E. K. &amp; Berner, R. A. Global Environment: Water, Air, and Geochemical Cycles. (Princeton Univ. Pcess, Princeton, 2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR6\" id=\"ref-link-section-d198960051e1023\" rel=\"nofollow noopener\" target=\"_blank\">6<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 31\" title=\"Stumm, W. &amp; Morgan, J. J. Aquatic Chemistry: Chemical Equilibria and Rates in Natural Waters. (John Wiley &amp; Sons, New York, 2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR31\" id=\"ref-link-section-d198960051e1026\" rel=\"nofollow noopener\" target=\"_blank\">31<\/a>, releasing alkalinity and dissolved salts, including oxidants, which are ultimately transported to the ocean via rivers and subterranean groundwater discharge. In the marine environment, solutes accumulate until they are removed through the deposition and burial of authigenic minerals in local environments<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 32\" title=\"Burdige, D. J. Geochemistry of Marine Sediments. (Princeton Univ. Press, Princeton, 2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR32\" id=\"ref-link-section-d198960051e1031\" rel=\"nofollow noopener\" target=\"_blank\">32<\/a>, entering the large-scale, long-term deep element cycle via sediment recycling.<\/p>\n<p>Table 1 Examples of aquatic habitats and the characteristic metabolisms on the modern Earth<\/p>\n<p>Beyond these major surface water reservoirs, a diverse suite of smaller-volume water bodies exists with a very broad array of physicochemical profiles (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). These environments, which include hypersaline lakes, brine systems in sea ice or in sedimentary pore waters, subglacial lakes, and hydrothermal vent systems, can exhibit extreme ranges in temperature, pressure, pH, salinity, and redox conditions<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 33\" title=\"Merino, N. et al. Living at the extremes: extremophiles and the limits of life in a planetary context. Front. Microbiol. 10, 780 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR33\" id=\"ref-link-section-d198960051e1382\" rel=\"nofollow noopener\" target=\"_blank\">33<\/a>. Their chemical differences commonly reflect isolation from major surface water reservoirs within diverse lithologies, allowing them to sustain steep physicochemical gradients against the backdrop of Earth\u2019s generally mild and oxidizing surface conditions. Although these extreme environments are rare on the modern Earth, they are highly relevant as analogs for environments proposed for the origin of life on the early Earth and potentially in water bodies on other planetary surfaces and interiors. For instance, alkaline soda lakes and low-temperature serpentinization-driven hydrothermal systems have been proposed as key settings for the origin of life on Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Martin, W., Baross, J., Kelley, D. &amp; Russell, M. J. Hydrothermal vents and the origin of life. Nat. Rev. Microbiol. 6, 805&#x2013;814 (2008).\" href=\"#ref-CR34\" id=\"ref-link-section-d198960051e1386\">34<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Baross, J. A. &amp; Hoffman, S. E. Submarine hydrothermal vents and associated gradient environments as sites for the origin and evolution of life. Orig. Life Evol. Biosph. 15, 327&#x2013;345 (1985).\" href=\"#ref-CR35\" id=\"ref-link-section-d198960051e1386_1\">35<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 36\" title=\"Cohen, Z. R. et al. Natural soda lakes provide compatible conditions for RNA and membrane function that could have enabled the origin of life. PNAS Nexus 3, pgae084 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR36\" id=\"ref-link-section-d198960051e1389\" rel=\"nofollow noopener\" target=\"_blank\">36<\/a> as well as in the subsurface ocean of Enceladus<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 37\" title=\"Glein, C. R., Postberg, F., Vance, S. D. The Geochemistry of Enceladus: Composition and Controls. In Enceladus and the icy moons of Saturn. 39&#x2013;56Arizona Univ. Press: Tucson, 2018.\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR37\" id=\"ref-link-section-d198960051e1393\" rel=\"nofollow noopener\" target=\"_blank\">37<\/a>, due to steep chemical gradients and high availability of nutrients and reductants. Similarly, terrestrial hot spring systems have been proposed as analogs for ancient hydrothermal environments on early Mars<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 38\" title=\"Des Marais, D. J. &amp; Walter, M. R. Terrestrial hot spring systems: introduction. Astrobiology 19, 1419&#x2013;1432 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR38\" id=\"ref-link-section-d198960051e1397\" rel=\"nofollow noopener\" target=\"_blank\">38<\/a>. Consequently, the study of these extreme aquatic systems on Earth provides a critical foundation for assessing the habitability and potential biosignatures of extraterrestrial environments<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 33\" title=\"Merino, N. et al. Living at the extremes: extremophiles and the limits of life in a planetary context. Front. Microbiol. 10, 780 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR33\" id=\"ref-link-section-d198960051e1402\" rel=\"nofollow noopener\" target=\"_blank\">33<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 39\" title=\"Rothschild, L. J. &amp; Mancinelli, R. L. Life in extreme environments. Nature 409, 1092&#x2013;1101 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR39\" id=\"ref-link-section-d198960051e1405\" rel=\"nofollow noopener\" target=\"_blank\">39<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 40\" title=\"Pikuta, E. V., Hoover, R. B. &amp; Tang, J. Microbial extremophiles at the limits of life. Crit. Rev. Microbiol. 33, 183&#x2013;209 (2007).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR40\" id=\"ref-link-section-d198960051e1408\" rel=\"nofollow noopener\" target=\"_blank\">40<\/a>.<\/p>\n<p>Microbial life is ubiquitous in modern aquatic environments, with the active assemblage (encompassing a range of functionality and activity level) and the residence time of characteristic microbes largely determined by the physicochemical parameters of water. In sunlit water, oxygenic photoautotrophs\u2014including prokaryotic cyanobacteria (e.g., Prochlorococcus, Synechococcus) and eukaryotic phytoplankton\u2014 capture solar energy to drive primary production<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 41\" title=\"Dadheech, P. K. Cyanobacteria&#x2014;the Pioneering Photoautotrophs. in Cyanobacteria. Elsevier: Amsterdam, 2024). 1&#x2013;18.\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR41\" id=\"ref-link-section-d198960051e1421\" rel=\"nofollow noopener\" target=\"_blank\">41<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Dissolved organic matter derived from these photoautotrophs sustains heterotrophs, which remain numerically dominant across most aquatic systems<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 42\" title=\"Giovannoni, S. J. SAR11 bacteria: the most abundant plankton in the oceans. Ann. Rev. Mar. Sci. 9, 231&#x2013;255 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR42\" id=\"ref-link-section-d198960051e1428\" rel=\"nofollow noopener\" target=\"_blank\">42<\/a>. Going deeper down the water column, light becomes limiting, and thus, photosynthesis declines and then ceases. Organic matter derived from descending photosynthetic biomass fuels heterotrophic respiration as it settles into dark environments<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 43\" title=\"Herndl, G., Bayer, B., Baltar, F. &amp; Reinthaler, T. Prokaryotic life in the deep ocean&#x2019;s water column. Ann. Rev. Mar. Sci. 15, 461&#x2013;483 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR43\" id=\"ref-link-section-d198960051e1433\" rel=\"nofollow noopener\" target=\"_blank\">43<\/a>. In oxygen-minimum zones and anoxic sediments, aerobic respiration is replaced by anaerobic respiration, where the oxidation of organic carbon is coupled to the reduction of nitrate, sulfate, or other available oxidants<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 32\" title=\"Burdige, D. J. Geochemistry of Marine Sediments. (Princeton Univ. Press, Princeton, 2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR32\" id=\"ref-link-section-d198960051e1437\" rel=\"nofollow noopener\" target=\"_blank\">32<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 44\" title=\"J&#xF8;rgensen, B. B. Bacteria and Marine Biogeochemistry. in Marine Geochemistry 173&#x2013;207. (Springer, Berlin, Heidelberg, 2000).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR44\" id=\"ref-link-section-d198960051e1440\" rel=\"nofollow noopener\" target=\"_blank\">44<\/a>. Anoxic habitats also host microbial consortia capable of conducting anaerobic oxidation of methane coupled to sulfate reduction, a process largely mediated by consortia of anaerobic methanotrophic archaea (ANME) and sulfate-reducing bacteria (SRB)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 45\" title=\"Hinrichs, K.-U., Hayes, J. M., Sylva, S. P., Brewer, P. G. &amp; DeLong, E. F. Methane-consuming archaebacteria in marine sediments. Nature 398, 802&#x2013;805 (1999).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR45\" id=\"ref-link-section-d198960051e1444\" rel=\"nofollow noopener\" target=\"_blank\">45<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>).<\/p>\n<p>Elevated concentrations of metabolic substrates can be supplied by water-rock interactions (e.g., serpentinization) and associated abiotic synthesis of organic carbon (e.g., Fischer-Tropsch reaction), as well as during radiolysis. In some interface environments, the mixing of reduced (e.g., Fe2+, H2, CH4, H2S, and organics) and oxidized substrates (e.g., O2, Fe(III), and SO42-) can build redox disequilibrium that provides the energetic driving force for chemolithoautotrophic metabolisms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Lin, L.-H. et al. Long-term sustainability of a high-energy, low-diversity crustal biome. Science 314, 479&#x2013;482 (2006).\" href=\"#ref-CR46\" id=\"ref-link-section-d198960051e1468\">46<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Chivian, D. et al. Environmental genomics reveals a single-species ecosystem deep within Earth. Science 322, 275&#x2013;278 (2008).\" href=\"#ref-CR47\" id=\"ref-link-section-d198960051e1468_1\">47<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 48\" title=\"Kelley, D. S. et al. A serpentinite-hosted ecosystem: the Lost City hydrothermal field. Science 307, 1428&#x2013;1434 (2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR48\" id=\"ref-link-section-d198960051e1471\" rel=\"nofollow noopener\" target=\"_blank\">48<\/a>. These microorganisms dominate the base trophic level on the modern Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 49\" title=\"Lau, M. C. Y. et al. An oligotrophic deep-subsurface community dependent on syntrophy is dominated by sulfur-driven autotrophic denitrifiers. Proc. Natl. Acad. Sci. USA 113, E7927&#x2013;E7936 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR49\" id=\"ref-link-section-d198960051e1475\" rel=\"nofollow noopener\" target=\"_blank\">49<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 50\" title=\"Dick, G. J. The microbiomes of deep-sea hydrothermal vents: distributed globally, shaped locally. Nat. Rev. Microbiol. 17, 271&#x2013;283 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR50\" id=\"ref-link-section-d198960051e1478\" rel=\"nofollow noopener\" target=\"_blank\">50<\/a>, including anaerobes such as methanogens, ANME, acetogens and SRB, as well as aerobes like sulfur-oxidizing and hydrogen-oxidizing bacteria (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Even at extreme temperature, pH, pressure and salinity conditions, diverse microorganisms have evolved to thrive in hostile environments such as hydrothermal vents and subglacial lakes<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 33\" title=\"Merino, N. et al. Living at the extremes: extremophiles and the limits of life in a planetary context. Front. Microbiol. 10, 780 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR33\" id=\"ref-link-section-d198960051e1485\" rel=\"nofollow noopener\" target=\"_blank\">33<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 51\" title=\"Siegert, M. J. et al. Physical, chemical and biological processes in Lake Vostok and other Antarctic subglacial lakes. Nature 414, 603&#x2013;609 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR51\" id=\"ref-link-section-d198960051e1488\" rel=\"nofollow noopener\" target=\"_blank\">51<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). These discoveries have continuously expanded the boundary conditions of life<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 39\" title=\"Rothschild, L. J. &amp; Mancinelli, R. L. Life in extreme environments. Nature 409, 1092&#x2013;1101 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR39\" id=\"ref-link-section-d198960051e1496\" rel=\"nofollow noopener\" target=\"_blank\">39<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 40\" title=\"Pikuta, E. V., Hoover, R. B. &amp; Tang, J. Microbial extremophiles at the limits of life. Crit. Rev. Microbiol. 33, 183&#x2013;209 (2007).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR40\" id=\"ref-link-section-d198960051e1499\" rel=\"nofollow noopener\" target=\"_blank\">40<\/a>, and thus increase the plausibility of life surviving in other planetary waters<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 33\" title=\"Merino, N. et al. Living at the extremes: extremophiles and the limits of life in a planetary context. Front. Microbiol. 10, 780 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR33\" id=\"ref-link-section-d198960051e1503\" rel=\"nofollow noopener\" target=\"_blank\">33<\/a>.<\/p>\n<p>Importantly, there is often a large diversity of low-abundance taxa<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 52\" title=\"Pascoal, F., Costa, R. &amp; Magalh&#xE3;es, C. The microbial rare biosphere: current concepts, methods and ecological principles. FEMS Microbiol. Ecol. 97, fiaa227 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR52\" id=\"ref-link-section-d198960051e1511\" rel=\"nofollow noopener\" target=\"_blank\">52<\/a> and spores<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 53\" title=\"Lomstein, B. A., Langerhuus, A. T., D&#x2019;Hondt, S., J&#xF8;rgensen, B. B. &amp; Spivack, A. J. Endospore abundance, microbial growth and necromass turnover in deep sub-seafloor sediment. Nature 484, 101&#x2013;104 (2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR53\" id=\"ref-link-section-d198960051e1515\" rel=\"nofollow noopener\" target=\"_blank\">53<\/a> in addition to the dormant forms mentioned above, and together these constitute an integrated microbial community. At steady state, the overall biomass of each ecosystem is dependent upon the availability of energy or nutrients, which is strongly shaped by local or global hydrological processes, and can vary significantly across modern environments. For example, cell concentrations can reach &gt; 108 cells mL\u22121 in nutrient-rich eutrophic coastal waters (e.g., Li et al<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 54\" title=\"Li, H. et al. Eutrophication reshapes microbial communities and life-history strategies in the riverine ecosystems. Environ. Microbiol. Rep. 17, e70234 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR54\" id=\"ref-link-section-d198960051e1523\" rel=\"nofollow noopener\" target=\"_blank\">54<\/a>.; Yuan et al<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 55\" title=\"Yuan, X., He, L., Yin, K., Pan, G. &amp; Harrison, P. J. Bacterial distribution and nutrient limitation in relation to different water masses in the coastal and northwestern South China Sea in late summer. Cont. Shelf Res. 31, 1214&#x2013;1223 (2011).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR55\" id=\"ref-link-section-d198960051e1528\" rel=\"nofollow noopener\" target=\"_blank\">55<\/a>.), largely sustained by riverine transport and ocean upwelling<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 56\" title=\"Falkowski, P. G., Barber, R. T. &amp; Smetacek, V. Biogeochemical controls and feedbacks on ocean primary production. Science 281, 200&#x2013;206 (1998).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR56\" id=\"ref-link-section-d198960051e1532\" rel=\"nofollow noopener\" target=\"_blank\">56<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 57\" title=\"Mann, K. H. &amp; Lazier, J. R. N. Dynamics of Marine Ecosystems: Biological-Physical Interactions in the Oceans. (John Wiley &amp; Sons, Hoboken, 2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR57\" id=\"ref-link-section-d198960051e1535\" rel=\"nofollow noopener\" target=\"_blank\">57<\/a>, but decline to &lt;103 cells mL\u22121 in nutrient-poor subsurface waters<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 58\" title=\"Magnabosco, C. et al. The biomass and biodiversity of the continental subsurface. Nat. Geosci. 11, 707&#x2013;717 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR58\" id=\"ref-link-section-d198960051e1543\" rel=\"nofollow noopener\" target=\"_blank\">58<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 59\" title=\"Hofmann, R. &amp; Griebler, C. DOM and bacterial growth efficiency in oligotrophic groundwater: absence of priming and co-limitation by organic carbon and phosphorus. Aquat. Microb. Ecol. 81, 55&#x2013;71 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR59\" id=\"ref-link-section-d198960051e1546\" rel=\"nofollow noopener\" target=\"_blank\">59<\/a>. While saline lakes and freshwater lakes and rivers together have previously been estimated to host ~2\u2009\u00d7\u20091026 prokaryotic cells (i.e., archaea and bacteria)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 60\" title=\"Whitman, W. B., Coleman, D. C. &amp; Wiebe, W. J. Prokaryotes: the unseen majority. Proc. Natl. Acad. Sci. USA 95, 6578&#x2013;6583 (1998).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR60\" id=\"ref-link-section-d198960051e1553\" rel=\"nofollow noopener\" target=\"_blank\">60<\/a>, the majority of prokaryotic cells reside in the marine pelagic zones (~1\u2009\u00d7\u20091029 cells at 0\u20134000\u2009m water depth)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 61\" title=\"Bar-On, Y. M., Phillips, R. &amp; Milo, R. The biomass distribution on Earth. Proc. Natl. Acad. Sci. USA 115, 6506&#x2013;6511 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR61\" id=\"ref-link-section-d198960051e1559\" rel=\"nofollow noopener\" target=\"_blank\">61<\/a>. In addition, despite having extremely slow cellular turnover times<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 62\" title=\"Onstott, T. C. et al. Does aspartic acid racemization constrain the depth limit of the subsurface biosphere? Geobiology 12, 1&#x2013;19 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR62\" id=\"ref-link-section-d198960051e1563\" rel=\"nofollow noopener\" target=\"_blank\">62<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 63\" title=\"Braun, S. et al. Microbial turnover times in the deep seabed studied by amino acid racemization modelling. Sci. Rep. 7, 5680 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR63\" id=\"ref-link-section-d198960051e1566\" rel=\"nofollow noopener\" target=\"_blank\">63<\/a>, marine sediments and terrestrial groundwater can also host significant cell populations (&gt; 1029 cells)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 61\" title=\"Bar-On, Y. M., Phillips, R. &amp; Milo, R. The biomass distribution on Earth. Proc. Natl. Acad. Sci. USA 115, 6506&#x2013;6511 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR61\" id=\"ref-link-section-d198960051e1572\" rel=\"nofollow noopener\" target=\"_blank\">61<\/a>. Comparatively, waters on the modern Earth, characterized primarily by a moderate range of physio-chemical parameters, have built a robust and complex ecosystem with much higher primary productivity than the reconstructed levels for the ancient Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 64\" title=\"Crockford, P. W., Bar On, Y. M., Ward, L. M., Milo, R. &amp; Halevy, I. The geologic history of primary productivity. Curr. Biol. 33, 4741&#x2013;4750.e5 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR64\" id=\"ref-link-section-d198960051e1577\" rel=\"nofollow noopener\" target=\"_blank\">64<\/a> (see also Section 2) and theoretical estimates for extraterrestrial water bodies<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Chyba, C. F. &amp; Hand, K. P. Life without photosynthesis. Science 292, 2026&#x2013;2027 (2001).\" href=\"#ref-CR65\" id=\"ref-link-section-d198960051e1584\">65<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Taubner, R.-S. et al. Biological methane production under putative Enceladus-like conditions. Nat. Commun. 9, 748 (2018).\" href=\"#ref-CR66\" id=\"ref-link-section-d198960051e1584_1\">66<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 67\" title=\"Hand, K. P., Carlson, R. W. &amp; Chyba, C. F. Energy, chemical disequilibrium, and geological constraints on Europa. Astrobiology 7, 1006&#x2013;1022 (2007).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR67\" id=\"ref-link-section-d198960051e1587\" rel=\"nofollow noopener\" target=\"_blank\">67<\/a> (Section 3 &amp; 4).<\/p>\n<p>Section 2. Co-evolution of the hydrosphere and biosphere on the Earth<\/p>\n<p>Earth\u2019s surface has hosted a liquid hydrosphere since the early Hadean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 68\" title=\"Sleep, N. H. The Hadean-Archaean environment. Cold Spring Harb. Perspect. Biol. 2, a002527 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR68\" id=\"ref-link-section-d198960051e1605\" rel=\"nofollow noopener\" target=\"_blank\">68<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 69\" title=\"Wilde, S. A., Valley, J. W., Peck, W. H. &amp; Graham, C. M. Evidence from detrital zircons for the existence of continental crust and oceans on the Earth 4.4 Gyr ago. Nature 409, 175&#x2013;178 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR69\" id=\"ref-link-section-d198960051e1608\" rel=\"nofollow noopener\" target=\"_blank\">69<\/a>. The planet\u2019s water is thought to have been delivered by H-bearing planetesimals and meteorites and\/or formed during the interaction of Earth\u2019s early magma ocean with nebular gas, though the relative proportions remain debated<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 70\" title=\"Horn, H. W., Vazan, A., Chariton, S., Prakapenka, V. B. &amp; Shim, S.-H. Building wet planets through high-pressure magma&#x2013;hydrogen reactions. Nature 646, 1069&#x2013;1074 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR70\" id=\"ref-link-section-d198960051e1612\" rel=\"nofollow noopener\" target=\"_blank\">70<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 71\" title=\"Piani, L. et al. Earth&#x2019;s water may have been inherited from material similar to enstatite chondrite meteorites. Science 369, 1110&#x2013;1113 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR71\" id=\"ref-link-section-d198960051e1615\" rel=\"nofollow noopener\" target=\"_blank\">71<\/a>. Nevertheless, high oxygen isotope ratios in detrital zircons suggest the presence of liquid water near Earth\u2019s surface as early as 4.4 Ga<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 69\" title=\"Wilde, S. A., Valley, J. W., Peck, W. H. &amp; Graham, C. M. Evidence from detrital zircons for the existence of continental crust and oceans on the Earth 4.4 Gyr ago. Nature 409, 175&#x2013;178 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR69\" id=\"ref-link-section-d198960051e1619\" rel=\"nofollow noopener\" target=\"_blank\">69<\/a>. Despite significantly lower solar luminosity at this time, a \u201cFaint Young Sun\u201d was likely compensated for by high atmospheric concentrations of greenhouse gases<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 72\" title=\"Feulner, G. The faint young Sun problem. Rev. Geophys. 50, (2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR72\" id=\"ref-link-section-d198960051e1623\" rel=\"nofollow noopener\" target=\"_blank\">72<\/a>, which maintained temperate conditions for sustaining a persistent liquid surface ocean. Direct geological constraints on the Hadean ocean\u2019s chemistry, including its pH, salinity, and trace element inventory, are sparse and contentious<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Trail, D. &amp; McCollom, T. M. Relatively oxidized fluids fed Earth&#x2019;s earliest hydrothermal systems. Science 379, 582&#x2013;586 (2023).\" href=\"#ref-CR73\" id=\"ref-link-section-d198960051e1627\">73<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Pinti, D. L. The Origin and Evolution of the Oceans. in Lectures in Astrobiology: Volume I 83&#x2013;112 (Springer, Berlin, Heidelberg, 2005).\" href=\"#ref-CR74\" id=\"ref-link-section-d198960051e1627_1\">74<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 75\" title=\"Holland, H. D. The Chemical Evolution of the Atmosphere and Oceans. (Princeton Univ. Press, 1984).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR75\" id=\"ref-link-section-d198960051e1630\" rel=\"nofollow noopener\" target=\"_blank\">75<\/a>. Furthermore, the habitability of these primordial waters was likely frequently perturbed by intense impact bombardment<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 76\" title=\"Zahnle, K. J., Lupu, R., Catling, D. C. &amp; Wogan, N. Creation and evolution of impact-generated reduced atmospheres of early Earth. Planet. Sci. J. 1, 11 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR76\" id=\"ref-link-section-d198960051e1635\" rel=\"nofollow noopener\" target=\"_blank\">76<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 77\" title=\"Kadoya, S., Krissansen-Totton, J. &amp; Catling, D. C. Probable cold and alkaline surface environment of the Hadean Earth caused by impact ejecta weathering. Geochem. Geophys. Geosyst. 21, e2019GC008734 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR77\" id=\"ref-link-section-d198960051e1638\" rel=\"nofollow noopener\" target=\"_blank\">77<\/a>.<\/p>\n<p>Despite the above considerations, principles of water-rock interaction allow for general inferences about early aquatic environments (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). High volcanic outgassing fluxes of CO2 and other acidic volatiles likely rendered most surface waters weakly acidic to circumneutral<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 78\" title=\"Krissansen-Totton, J., Arney, G. N. &amp; Catling, D. C. Constraining the climate and ocean pH of the early Earth with a geological carbon cycle model. Proc. Natl. Acad. Sci. USA 115, 4105&#x2013;4110 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR78\" id=\"ref-link-section-d198960051e1650\" rel=\"nofollow noopener\" target=\"_blank\">78<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 79\" title=\"Halevy, I. &amp; Bachan, A. The geologic history of seawater pH. Science 355, 1069&#x2013;1071 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR79\" id=\"ref-link-section-d198960051e1653\" rel=\"nofollow noopener\" target=\"_blank\">79<\/a>. However, progressive silicate weathering would have gradually increased ocean alkalinity and pH over time<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 78\" title=\"Krissansen-Totton, J., Arney, G. N. &amp; Catling, D. C. Constraining the climate and ocean pH of the early Earth with a geological carbon cycle model. Proc. Natl. Acad. Sci. USA 115, 4105&#x2013;4110 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR78\" id=\"ref-link-section-d198960051e1657\" rel=\"nofollow noopener\" target=\"_blank\">78<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 79\" title=\"Halevy, I. &amp; Bachan, A. The geologic history of seawater pH. Science 355, 1069&#x2013;1071 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR79\" id=\"ref-link-section-d198960051e1660\" rel=\"nofollow noopener\" target=\"_blank\">79<\/a>, perhaps buffering at a value around 6.5-7 in the Archean (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2a<\/a>). Before the emergence of large continental landmasses, alkalinity and critical greenhouse gases (such as CH4<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 80\" title=\"Etiope, G. &amp; Sherwood Lollar, B. Abiotic methane on earth. Rev. Geophys. 51, 276&#x2013;299 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR80\" id=\"ref-link-section-d198960051e1669\" rel=\"nofollow noopener\" target=\"_blank\">80<\/a> and potentially H2<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 81\" title=\"Leong, J. A. M., Ely, T. &amp; Shock, E. L. Decreasing extents of Archean serpentinization contributed to the rise of an oxidized atmosphere. Nat. Commun. 12, 7341 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR81\" id=\"ref-link-section-d198960051e1674\" rel=\"nofollow noopener\" target=\"_blank\">81<\/a>) released from seafloor basalt weathering and impact ejecta probably served as a principal feedback stabilizing Earth\u2019s early climate<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 72\" title=\"Feulner, G. The faint young Sun problem. Rev. Geophys. 50, (2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR72\" id=\"ref-link-section-d198960051e1678\" rel=\"nofollow noopener\" target=\"_blank\">72<\/a>, although these reactions remain poorly quantified<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 77\" title=\"Kadoya, S., Krissansen-Totton, J. &amp; Catling, D. C. Probable cold and alkaline surface environment of the Hadean Earth caused by impact ejecta weathering. Geochem. Geophys. Geosyst. 21, e2019GC008734 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR77\" id=\"ref-link-section-d198960051e1682\" rel=\"nofollow noopener\" target=\"_blank\">77<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 78\" title=\"Krissansen-Totton, J., Arney, G. N. &amp; Catling, D. C. Constraining the climate and ocean pH of the early Earth with a geological carbon cycle model. Proc. Natl. Acad. Sci. USA 115, 4105&#x2013;4110 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR78\" id=\"ref-link-section-d198960051e1685\" rel=\"nofollow noopener\" target=\"_blank\">78<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 82\" title=\"Coogan, L. A. &amp; Gillis, K. M. Low-Temperature Alteration of the Seafloor: Impacts on Ocean Chemistry. Annu. Rev. Earth Planet. Sci. 46, 21&#x2013;45 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR82\" id=\"ref-link-section-d198960051e1688\" rel=\"nofollow noopener\" target=\"_blank\">82<\/a>. These extensive water-rock interactions might also have helped maintain reducing surface waters and an oxygen-poor atmosphere in the early Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 81\" title=\"Leong, J. A. M., Ely, T. &amp; Shock, E. L. Decreasing extents of Archean serpentinization contributed to the rise of an oxidized atmosphere. Nat. Commun. 12, 7341 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR81\" id=\"ref-link-section-d198960051e1692\" rel=\"nofollow noopener\" target=\"_blank\">81<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 83\" title=\"Kasting, J. F. What caused the rise of atmospheric O2? Chem. Geol. 362, 13&#x2013;25 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR83\" id=\"ref-link-section-d198960051e1695\" rel=\"nofollow noopener\" target=\"_blank\">83<\/a>. Ocean salinity in the Hadean and Archean is poorly constrained<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 84\" title=\"Marty, B., Avice, G., Bekaert, D. V. &amp; Broadley, M. W. Salinity of the Archaean oceans from analysis of fluid inclusions in quartz. C. R. Geosci. 350, 154&#x2013;163 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR84\" id=\"ref-link-section-d198960051e1700\" rel=\"nofollow noopener\" target=\"_blank\">84<\/a> but might have reached levels close to or higher than the modern, given greater volcanic volatile input<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 74\" title=\"Pinti, D. L. The Origin and Evolution of the Oceans. in Lectures in Astrobiology: Volume I 83&#x2013;112 (Springer, Berlin, Heidelberg, 2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR74\" id=\"ref-link-section-d198960051e1704\" rel=\"nofollow noopener\" target=\"_blank\">74<\/a> and a reduced evaporite sink in the absence of extensive subaerial continents<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 85\" title=\"Knauth, L. P. Salinity History of the Earth&#x2019;s Ocean. in Encyclopedia of Geobiology 769&#x2013;772. (Springer, Dordrecht, 2011).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR85\" id=\"ref-link-section-d198960051e1708\" rel=\"nofollow noopener\" target=\"_blank\">85<\/a>. Intense hydrothermal alteration, coupled with limited riverine input, may have led to seawater enriched in Ca2+ but relatively depleted in Mg2+ compared to today<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 78\" title=\"Krissansen-Totton, J., Arney, G. N. &amp; Catling, D. C. Constraining the climate and ocean pH of the early Earth with a geological carbon cycle model. Proc. Natl. Acad. Sci. USA 115, 4105&#x2013;4110 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR78\" id=\"ref-link-section-d198960051e1716\" rel=\"nofollow noopener\" target=\"_blank\">78<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 86\" title=\"Jones, C., Nomosatryo, S., Crowe, S. A., Bjerrum, C. J. &amp; Canfield, D. E. Iron oxides, divalent cations, silica, and the early Earth phosphorus crisis. Geology 43, 135&#x2013;138 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR86\" id=\"ref-link-section-d198960051e1719\" rel=\"nofollow noopener\" target=\"_blank\">86<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2d<\/a>). Moreover, surface waters would accumulate reductants like Fe2+ and Mn2+ under a weakly reducing atmosphere devoid of biogenic oxygen <a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 75\" title=\"Holland, H. D. The Chemical Evolution of the Atmosphere and Oceans. (Princeton Univ. Press, 1984).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR75\" id=\"ref-link-section-d198960051e1731\" rel=\"nofollow noopener\" target=\"_blank\">75<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 87\" title=\"Hao, J., Sverjensky, D. A. &amp; Hazen, R. M. A model for late Archean chemical weathering and world average river water. Earth Planet. Sci. Lett. 457, 191&#x2013;203 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR87\" id=\"ref-link-section-d198960051e1734\" rel=\"nofollow noopener\" target=\"_blank\">87<\/a>, in contrast to the modern oxic surface waters (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2b &amp; 2c<\/a>). In such anoxic and ferruginous seawater, rapid reduction and reaction with abundant hydrothermal Ba2+ and Fe2+ would have kept dissolved sulfur levels in the micromolar range<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 88\" title=\"Crowe, S. A. et al. Sulfate was a trace constituent of Archean seawater. Science 346, 735&#x2013;739 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR88\" id=\"ref-link-section-d198960051e1746\" rel=\"nofollow noopener\" target=\"_blank\">88<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 89\" title=\"Li, X. et al. Isotopic evidence for oceanic barium cycling in the initial stage of the mesoproterozoic. Earth Planet. Sci. Lett. 658, 119314 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR89\" id=\"ref-link-section-d198960051e1749\" rel=\"nofollow noopener\" target=\"_blank\">89<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2d<\/a>). Ammonium (NH4+) would have been the thermodynamically stable form of dissolved nitrogen<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 90\" title=\"Hao, J., Sverjensky, D. A. &amp; Hazen, R. M. Mobility of nutrients and trace metals during weathering in the late Archean. Earth Planet. Sci. Lett. 471, 148&#x2013;159 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR90\" id=\"ref-link-section-d198960051e1759\" rel=\"nofollow noopener\" target=\"_blank\">90<\/a>, and small amounts of NH4+ were probably produced in hydrothermal settings<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 91\" title=\"Sun, L., Li, K., Sun, Z., Zhang, Y. &amp; Li, L. Abiotic N2 reduction in submarine hydrothermal systems could quickly fertilize prebiotic oceans. Nat. Commun. 16, 10608 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR91\" id=\"ref-link-section-d198960051e1767\" rel=\"nofollow noopener\" target=\"_blank\">91<\/a> and via the reduction of lightning-generated nitrogen oxides<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 92\" title=\"Laneuville, M., Kameya, M. &amp; Cleaves, H. Earth without life: a systems model of a global abiotic nitrogen cycle. Astrobiology 18, 897&#x2013;914 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR92\" id=\"ref-link-section-d198960051e1771\" rel=\"nofollow noopener\" target=\"_blank\">92<\/a>. In the absence of biosilicification, the primitive ocean would also have been rich in dissolved silica (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2d<\/a>), derived from hydrothermal leaching of the seafloor<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 93\" title=\"Hofmann, A. &amp; Harris, C. Silica alteration zones in the Barberton greenstone belt: A window into subseafloor processes 3.5&#x2013;3.3 Ga ago. Chem. Geol. 257, 221&#x2013;239 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR93\" id=\"ref-link-section-d198960051e1778\" rel=\"nofollow noopener\" target=\"_blank\">93<\/a>, leading to widespread silica precipitation<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 94\" title=\"Manning-Berg, A. R. &amp; Kah, L. C. Proterozoic microbial mats and their constraints on environments of silicification. Geobiology 15, 469&#x2013;483 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR94\" id=\"ref-link-section-d198960051e1783\" rel=\"nofollow noopener\" target=\"_blank\">94<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 95\" title=\"Gauvey, K. &amp; Kah, L. C. Unravelling neomorphism: recrystallization pathways in Proterozoic microfossiliferous chert. Front. Earth Sci. 13, 1598200 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR95\" id=\"ref-link-section-d198960051e1786\" rel=\"nofollow noopener\" target=\"_blank\">95<\/a> and silicification of early sediments<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 96\" title=\"Siever, R. The silica cycle in the Precambrian. Geochim. Cosmochim. Acta 56, 3265&#x2013;3272 (1992).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR96\" id=\"ref-link-section-d198960051e1790\" rel=\"nofollow noopener\" target=\"_blank\">96<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 97\" title=\"Conley, D. J. et al. Biosilicification drives a decline of dissolved Si in the oceans through geologic time. Front. Mar. Sci. 4, 397 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR97\" id=\"ref-link-section-d198960051e1793\" rel=\"nofollow noopener\" target=\"_blank\">97<\/a>. High dissolved silica is also proposed to favor reverse weathering in the early ocean, which might help stabilize the long-term evolution of climate and marine pH<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 98\" title=\"Isson, T. T. &amp; Planavsky, N. J. Reverse weathering as a long-term stabilizer of marine pH and planetary climate. Nature 560, 471&#x2013;475 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR98\" id=\"ref-link-section-d198960051e1797\" rel=\"nofollow noopener\" target=\"_blank\">98<\/a>.<\/p>\n<p>Fig. 2: Co-evolution of life and Earth\u2019s surface environments.<img decoding=\"async\" aria-describedby=\"figure-2-desc\" src=\"https:\/\/www.newsbeep.com\/ie\/wp-content\/uploads\/2026\/07\/41467_2026_74955_Fig2_HTML.png\" alt=\"Fig. 2: Co-evolution of life and Earth&#x2019;s surface environments.\" loading=\"lazy\" width=\"685\" height=\"362\"\/><\/p>\n<p>a Reconstructed evolution of ocean pH. The solid blue lines show median pH estimates from Krissansen-Totton et al<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 78\" title=\"Krissansen-Totton, J., Arney, G. N. &amp; Catling, D. C. Constraining the climate and ocean pH of the early Earth with a geological carbon cycle model. Proc. Natl. Acad. Sci. USA 115, 4105&#x2013;4110 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR78\" id=\"ref-link-section-d198960051e1815\" rel=\"nofollow noopener\" target=\"_blank\">78<\/a>., with the blue shading denoting the 95% confidence interval. Yellow dashed lines indicate the 95% confidence interval from Halevy and Bachan<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 79\" title=\"Halevy, I. &amp; Bachan, A. The geologic history of seawater pH. Science 355, 1069&#x2013;1071 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR79\" id=\"ref-link-section-d198960051e1819\" rel=\"nofollow noopener\" target=\"_blank\">79<\/a>. b Evolution of ocean redox stratification structure<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 239\" title=\"Amor, M., Mathon, F. P., Monteil, C. L., Busigny, V. &amp; Lefevre, C. T. Iron-biomineralizing organelle in magnetotactic bacteria: function, synthesis and preservation in ancient rock samples. Environ. Microbiol. 22, 3611&#x2013;3632 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR239\" id=\"ref-link-section-d198960051e1826\" rel=\"nofollow noopener\" target=\"_blank\">239<\/a>. c Reconstructed evolution of atmospheric O2 and CO2 levels through time<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 121\" title=\"Lyons, T. W. et al. Co-evolution of early Earth environments and microbial life. Nat. Rev. Microbiol. 22, 572&#x2013;586 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR121\" id=\"ref-link-section-d198960051e1838\" rel=\"nofollow noopener\" target=\"_blank\">121<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 240\" title=\"Catling, D. C. &amp; Zahnle, K. J. The Archean atmosphere. Sci. Adv. 6, eaax1420 (2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR240\" id=\"ref-link-section-d198960051e1841\" rel=\"nofollow noopener\" target=\"_blank\">240<\/a>. d Evolution of dissolved bio-essential elements in subsurface seawater and their dominant chemical forms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 28\" title=\"Anbar, A. D. Elements and evolution. Science 322, 1481&#x2013;1483 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR28\" id=\"ref-link-section-d198960051e1848\" rel=\"nofollow noopener\" target=\"_blank\">28<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 86\" title=\"Jones, C., Nomosatryo, S., Crowe, S. A., Bjerrum, C. J. &amp; Canfield, D. E. Iron oxides, divalent cations, silica, and the early Earth phosphorus crisis. Geology 43, 135&#x2013;138 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR86\" id=\"ref-link-section-d198960051e1851\" rel=\"nofollow noopener\" target=\"_blank\">86<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 96\" title=\"Siever, R. The silica cycle in the Precambrian. Geochim. Cosmochim. Acta 56, 3265&#x2013;3272 (1992).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR96\" id=\"ref-link-section-d198960051e1854\" rel=\"nofollow noopener\" target=\"_blank\">96<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 97\" title=\"Conley, D. J. et al. Biosilicification drives a decline of dissolved Si in the oceans through geologic time. Front. Mar. Sci. 4, 397 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR97\" id=\"ref-link-section-d198960051e1857\" rel=\"nofollow noopener\" target=\"_blank\">97<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Poulton, S. W. Early phosphorus redigested. Nat. Geosci. 10, 75&#x2013;76 (2017).\" href=\"#ref-CR241\" id=\"ref-link-section-d198960051e1860\">241<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Large, R. R. et al. Ocean and atmosphere geochemical proxies derived from trace elements in marine pyrite: Implications for ore genesis in sedimentary basins. Econ. Geol. 112, 423&#x2013;450 (2017).\" href=\"#ref-CR242\" id=\"ref-link-section-d198960051e1860_1\">242<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 243\" title=\"Johnson, J. E., Present, T. M. &amp; Valentine, J. S. Iron: Life&#x2019;s primeval transition metal. Proc. Natl. Acad. Sci. USA 121, e2318692121 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR243\" id=\"ref-link-section-d198960051e1863\" rel=\"nofollow noopener\" target=\"_blank\">243<\/a>. The [P] (% modern levels) axis represents the percentage of P concentrations in the ocean relative to modern levels. e Major evolutionary milestones of life, shown by red symbols and lines (upper), together with the evolution of total primary productivity, shown by the blue dashed field and lines (lower)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 64\" title=\"Crockford, P. W., Bar On, Y. M., Ward, L. M., Milo, R. &amp; Halevy, I. The geologic history of primary productivity. Curr. Biol. 33, 4741&#x2013;4750.e5 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR64\" id=\"ref-link-section-d198960051e1871\" rel=\"nofollow noopener\" target=\"_blank\">64<\/a>. f Geological constraints on the emergence of major microbial metabolisms during the Archean Eon<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 29\" title=\"Moore, E. K., Jelen, B. I., Giovannelli, D., Raanan, H. &amp; Falkowski, P. G. Metal availability and the expanding network of microbial metabolisms in the Archaean eon. Nat. Geosci. 10, 629&#x2013;636 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR29\" id=\"ref-link-section-d198960051e1878\" rel=\"nofollow noopener\" target=\"_blank\">29<\/a>. Dark age bars indicate relatively well-constrained age estimates, whereas lighter age bars indicate more uncertain ranges.<\/p>\n<p>This geochemical backdrop, or at least portions of it, provided the physical setting for life\u2019s emergence. Reducing conditions favored the abiotic synthesis and stability of organic molecules and their precursors<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 99\" title=\"Shock, E. &amp; Canovas, P. The potential for abiotic organic synthesis and biosynthesis at seafloor hydrothermal systems. Geofluids 10, 161&#x2013;192 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR99\" id=\"ref-link-section-d198960051e1893\" rel=\"nofollow noopener\" target=\"_blank\">99<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 100\" title=\"Patel, B. H., Percivalle, C., Ritson, D. J., Duffy, C. D. &amp; Sutherland, J. D. Common origins of RNA, protein and lipid precursors in a cyanosulfidic protometabolism. Nat. Chem. 7, 301&#x2013;307 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR100\" id=\"ref-link-section-d198960051e1896\" rel=\"nofollow noopener\" target=\"_blank\">100<\/a>. Key nutrients like nitrogen and phosphorus may have been more abundant or in more bioavailable forms than today<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 90\" title=\"Hao, J., Sverjensky, D. A. &amp; Hazen, R. M. Mobility of nutrients and trace metals during weathering in the late Archean. Earth Planet. Sci. Lett. 471, 148&#x2013;159 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR90\" id=\"ref-link-section-d198960051e1900\" rel=\"nofollow noopener\" target=\"_blank\">90<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 101\" title=\"Galloway, T., Baidya, A. S., Cousins, C. &amp; St&#xFC;eken, E. E. Planetary sources of bio-essential nutrients on a prebiotic world. Philos. Trans. R. Soc. B 380, 20240288 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR101\" id=\"ref-link-section-d198960051e1903\" rel=\"nofollow noopener\" target=\"_blank\">101<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 102\" title=\"Hao, J., Li, X., Ju, P. &amp; Pasek, M. Active redox cycling of phosphorus on the early Earth. Nat. Commun. 16, 4596 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR102\" id=\"ref-link-section-d198960051e1906\" rel=\"nofollow noopener\" target=\"_blank\">102<\/a>. Furthermore, extensive water-rock interactions produced catalytic mineral surfaces (e.g., phyllosilicates, native metals, metal oxides, and sulfides)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 34\" title=\"Martin, W., Baross, J., Kelley, D. &amp; Russell, M. J. Hydrothermal vents and the origin of life. Nat. Rev. Microbiol. 6, 805&#x2013;814 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR34\" id=\"ref-link-section-d198960051e1910\" rel=\"nofollow noopener\" target=\"_blank\">34<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 103\" title=\"Morrison, S. M., Prabhu, A. &amp; Hazen, R. M. An evolutionary system of mineralogy, Part VI: Earth&#x2019;s earliest Hadean crust (&gt; 4370&#x2009;Ma). Am. Mineral. 108, 42&#x2013;58 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR103\" id=\"ref-link-section-d198960051e1913\" rel=\"nofollow noopener\" target=\"_blank\">103<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 104\" title=\"Hazen, R. M. &amp; Sverjensky, D. A. Mineral surfaces, geochemical complexities, and the origins of life. Cold Spring Harb. Perspect. Biol. 2, a002162 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR104\" id=\"ref-link-section-d198960051e1916\" rel=\"nofollow noopener\" target=\"_blank\">104<\/a> that could have facilitated prebiotic chemistry. Meanwhile, large thermal and chemical gradients, sustained by surface radiation, impacts, and geothermal heating, provided the thermodynamic disequilibrium necessary to drive early metabolisms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 34\" title=\"Martin, W., Baross, J., Kelley, D. &amp; Russell, M. J. Hydrothermal vents and the origin of life. Nat. Rev. Microbiol. 6, 805&#x2013;814 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR34\" id=\"ref-link-section-d198960051e1920\" rel=\"nofollow noopener\" target=\"_blank\">34<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 104\" title=\"Hazen, R. M. &amp; Sverjensky, D. A. Mineral surfaces, geochemical complexities, and the origins of life. Cold Spring Harb. Perspect. Biol. 2, a002162 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR104\" id=\"ref-link-section-d198960051e1923\" rel=\"nofollow noopener\" target=\"_blank\">104<\/a>. Phylogenetic reconstructions indicate that the Last Universal Common Ancestor (LUCA) was likely a chemoautotroph, exploiting these abundant chemical energy sources<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 105\" title=\"Moody, E. R. R. et al. The nature of the last universal common ancestor and its impact on the early Earth system. Nat. Ecol. Evol. 8, 1654&#x2013;1666 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR105\" id=\"ref-link-section-d198960051e1927\" rel=\"nofollow noopener\" target=\"_blank\">105<\/a>.<\/p>\n<p>Life probably emerged in the late Hadean or early Archean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 105\" title=\"Moody, E. R. R. et al. The nature of the last universal common ancestor and its impact on the early Earth system. Nat. Ecol. Evol. 8, 1654&#x2013;1666 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR105\" id=\"ref-link-section-d198960051e1934\" rel=\"nofollow noopener\" target=\"_blank\">105<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2e<\/a>), coinciding with a decline in impact frequency<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 106\" title=\"Koeberl, C. The record of impact processes on the early Earth: A review of the first 2.5 billion years. Geol. Soc. Am. Spec. Pap. 405, 1&#x2013;22 (2006).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR106\" id=\"ref-link-section-d198960051e1941\" rel=\"nofollow noopener\" target=\"_blank\">106<\/a>. At the same time, continuous dissipation of Earth\u2019s internal heat would have resulted in a secular decline in geothermal heat flow<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 107\" title=\"Herzberg, C., Condie, K. &amp; Korenaga, J. Thermal history of the Earth and its petrological expression. Earth Planet. Sci. Lett. 292, 79&#x2013;88 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR107\" id=\"ref-link-section-d198960051e1945\" rel=\"nofollow noopener\" target=\"_blank\">107<\/a>, leading to reduced volcanic activity<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 108\" title=\"Viehmann, S. et al. Europium traces the impact of high temperature hydrothermal systems on the early oceans. Geochem. Perspect. Lett. 34, 57&#x2013;61 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR108\" id=\"ref-link-section-d198960051e1949\" rel=\"nofollow noopener\" target=\"_blank\">108<\/a>. In the meantime, continental landmass grew substantially in areal extent<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 109\" title=\"Taylor, S. R. &amp; McLennan, S. M. The geochemical evolution of the continental crust. Rev. Geophys. 33, 241&#x2013;265 (1995).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR109\" id=\"ref-link-section-d198960051e1954\" rel=\"nofollow noopener\" target=\"_blank\">109<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 110\" title=\"Cawood, P. A. et al. Secular evolution of continents and the Earth system. Rev. Geophys. 60, e2022RG000789 (2022).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR110\" id=\"ref-link-section-d198960051e1957\" rel=\"nofollow noopener\" target=\"_blank\">110<\/a>; however, much of it may have remained largely submerged until the middle or late Archean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 111\" title=\"Chowdhury, P., Cawood, P. A. &amp; Mulder, J. A. Subaerial emergence of continents on Archean Earth. Annu. Rev. Earth Planet. Sci. 53, 443&#x2013;478 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR111\" id=\"ref-link-section-d198960051e1961\" rel=\"nofollow noopener\" target=\"_blank\">111<\/a>. These shifts drove a decrease in atmospheric pCO2 and a rise in ocean pH and alkalinity<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 78\" title=\"Krissansen-Totton, J., Arney, G. N. &amp; Catling, D. C. Constraining the climate and ocean pH of the early Earth with a geological carbon cycle model. Proc. Natl. Acad. Sci. USA 115, 4105&#x2013;4110 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR78\" id=\"ref-link-section-d198960051e1970\" rel=\"nofollow noopener\" target=\"_blank\">78<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2a<\/a>). The emergence of continental landmasses also established modern-style lacustrine and riverine systems, promoting large-scale continental weathering and marine upwelling supplies of nutrients that could support rising biological productivity<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 64\" title=\"Crockford, P. W., Bar On, Y. M., Ward, L. M., Milo, R. &amp; Halevy, I. The geologic history of primary productivity. Curr. Biol. 33, 4741&#x2013;4750.e5 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR64\" id=\"ref-link-section-d198960051e1978\" rel=\"nofollow noopener\" target=\"_blank\">64<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 90\" title=\"Hao, J., Sverjensky, D. A. &amp; Hazen, R. M. Mobility of nutrients and trace metals during weathering in the late Archean. Earth Planet. Sci. Lett. 471, 148&#x2013;159 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR90\" id=\"ref-link-section-d198960051e1981\" rel=\"nofollow noopener\" target=\"_blank\">90<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 112\" title=\"Hao, J., Knoll, A. H., Huang, F., Hazen, R. M. &amp; Daniel, I. Cycling phosphorus on the Archean Earth: Part I. Continental weathering and riverine transport of phosphorus. Geochim. Cosmochim. Acta 273, 70&#x2013;84 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR112\" id=\"ref-link-section-d198960051e1984\" rel=\"nofollow noopener\" target=\"_blank\">112<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 113\" title=\"Kienert, H., Feulner, G. &amp; Petoukhov, V. Albedo and heat transport in 3-D model simulations of the early Archean climate. Climate 9, 1841&#x2013;1862 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR113\" id=\"ref-link-section-d198960051e1987\" rel=\"nofollow noopener\" target=\"_blank\">113<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2e<\/a>). Moreover, the establishment of these environmental settings might have spurred a diversification of microbial metabolisms (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2f<\/a>) as more oxidants and nutrients became available<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 29\" title=\"Moore, E. K., Jelen, B. I., Giovannelli, D., Raanan, H. &amp; Falkowski, P. G. Metal availability and the expanding network of microbial metabolisms in the Archaean eon. Nat. Geosci. 10, 629&#x2013;636 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR29\" id=\"ref-link-section-d198960051e1997\" rel=\"nofollow noopener\" target=\"_blank\">29<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 114\" title=\"Bianchini, G., Hagemann, M. &amp; S&#xE1;nchez-Baracaldo, P. Stochastic character mapping, Bayesian model selection, and biosynthetic pathways shed new light on the evolution of habitat preference in cyanobacteria. Syst. Biol. 73, 644&#x2013;665 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR114\" id=\"ref-link-section-d198960051e2000\" rel=\"nofollow noopener\" target=\"_blank\">114<\/a>. Among them, one critical innovation is the emergence of cyanobacteria probably in the middle to late Archean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 115\" title=\"Fischer, W. W., Hemp, J. &amp; Johnson, J. E. Evolution of oxygenic photosynthesis. Annu. Rev. Earth Planet. Sci. 44, 647&#x2013;683 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR115\" id=\"ref-link-section-d198960051e2004\" rel=\"nofollow noopener\" target=\"_blank\">115<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 116\" title=\"S&#xE1;nchez-Baracaldo, P. &amp; Cardona, T. On the origin of oxygenic photosynthesis and Cyanobacteria. N. 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Lett. 471, 148&#x2013;159 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR90\" id=\"ref-link-section-d198960051e2044\" rel=\"nofollow noopener\" target=\"_blank\">90<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 118\" title=\"Crockford, P. W. et al. Revisiting the greatness of Earth&#x2019;s great oxidation. Commun. Earth Environ. 7, 348 (2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR118\" id=\"ref-link-section-d198960051e2047\" rel=\"nofollow noopener\" target=\"_blank\">118<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 120\" title=\"Konhauser, K. O. et al. Aerobic bacterial pyrite oxidation and acid rock drainage during the Great Oxidation Event. Nature 478, 369&#x2013;373 (2011).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR120\" id=\"ref-link-section-d198960051e2050\" rel=\"nofollow noopener\" target=\"_blank\">120<\/a>. However, atmospheric O2 levels likely remained low (&lt;10% of present levels) during the Proterozoic<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 118\" title=\"Crockford, P. W. et al. Revisiting the greatness of Earth&#x2019;s great oxidation. Commun. Earth Environ. 7, 348 (2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR118\" id=\"ref-link-section-d198960051e2056\" rel=\"nofollow noopener\" target=\"_blank\">118<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 121\" title=\"Lyons, T. W. et al. Co-evolution of early Earth environments and microbial life. Nat. Rev. Microbiol. 22, 572&#x2013;586 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR121\" id=\"ref-link-section-d198960051e2059\" rel=\"nofollow noopener\" target=\"_blank\">121<\/a>, resulting in a stratified ocean: oxygenated at the surface but anoxic and largely ferruginous, with localized euxinic margins (H2S-rich) at depth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 122\" title=\"Poulton, S. W. &amp; Canfield, D. E. Ferruginous conditions: a dominant feature of the ocean through Earth&#x2019;s history. Elements 7, 107&#x2013;112 (2011).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR122\" id=\"ref-link-section-d198960051e2065\" rel=\"nofollow noopener\" target=\"_blank\">122<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2b<\/a>). The reasons for this protracted period of intermediate oxygen levels despite cyanobacterial dominance remain unclear, but may be linked to persistently low primary productivity, as suggested by invariant carbon isotope records and significantly negative triple oxygen isotope values<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 123\" title=\"Buick, R., Des Marais, D. J. &amp; Knoll, A. H. 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N. et al. The supercontinent cycle. Nat. Rev. Earth Environ. 2, 358&#x2013;374 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR125\" id=\"ref-link-section-d198960051e2080\" rel=\"nofollow noopener\" target=\"_blank\">125<\/a>, coupled with overall low continental topographic relief<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 126\" title=\"Tang, M., Chu, X., Hao, J. &amp; Shen, B. Orogenic quiescence in Earth&#x2019;s middle age. Science 371, 728&#x2013;731 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR126\" id=\"ref-link-section-d198960051e2084\" rel=\"nofollow noopener\" target=\"_blank\">126<\/a>, might have limited nutrient supply via continental weathering and ocean upwelling<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 126\" title=\"Tang, M., Chu, X., Hao, J. &amp; Shen, B. Orogenic quiescence in Earth&#x2019;s middle age. Science 371, 728&#x2013;731 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR126\" id=\"ref-link-section-d198960051e2088\" rel=\"nofollow noopener\" target=\"_blank\">126<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 127\" title=\"Olson, S. L., Jansen, M. &amp; Abbot, D. S. Oceanographic considerations for exoplanet life detection. Astrophys. J. 895, 19 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR127\" id=\"ref-link-section-d198960051e2091\" rel=\"nofollow noopener\" target=\"_blank\">127<\/a>, constraining marine productivity<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 128\" title=\"Laakso, T. A. &amp; Schrag, D. P. A theory of atmospheric oxygen. Geobiology 15, 366&#x2013;384 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR128\" id=\"ref-link-section-d198960051e2095\" rel=\"nofollow noopener\" target=\"_blank\">128<\/a>. Despite this apparent environmental stability, the mid-Proterozoic witnessed the emergence of eukaryotes and multicellular organisms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 129\" title=\"Brocks, J. J. et al. Lost world of complex life and the late rise of the eukaryotic crown. Nature 618, 767&#x2013;773 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR129\" id=\"ref-link-section-d198960051e2100\" rel=\"nofollow noopener\" target=\"_blank\">129<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2e<\/a>), potentially driven by local habitat heterogeneity or environmental perturbations<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 130\" title=\"Zhang, S. et al. Subaerial volcanism broke mid-Proterozoic environmental stasis. Sci. Adv. 10, eadk5991 (2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR130\" id=\"ref-link-section-d198960051e2107\" rel=\"nofollow noopener\" target=\"_blank\">130<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 131\" title=\"Diamond, C. W., Ernst, R. E., Zhang, S.-H. &amp; Lyons, T. W. Breaking the Boring Billion. in Large Igneous Provinces 487&#x2013;501. (Wiley, Hoboken, 2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR131\" id=\"ref-link-section-d198960051e2110\" rel=\"nofollow noopener\" target=\"_blank\">131<\/a>. The subsequent breakup of Rodinia and increased orogenic activity in the late Proterozoic enhanced continental weathering<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 132\" title=\"Donnadieu, Y., Godd&#xE9;ris, Y., Ramstein, G., N&#xE9;d&#xE9;lec, A. &amp; Meert, J. A &#x2018;snowball Earth&#x2019; climate triggered by continental break-up through changes in runoff. Nature 428, 303&#x2013;306 (2004).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR132\" id=\"ref-link-section-d198960051e2114\" rel=\"nofollow noopener\" target=\"_blank\">132<\/a>, nutrient supply, and oxygen production, culminating in the second oxygenation of the atmosphere and ocean (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2b, c<\/a>) and the radiation of complex eukaryotic life in the Ediacaran and early Phanerozoic<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 133\" title=\"Zhang, X. et al. Triggers for the Cambrian explosion: hypotheses and problems. Gondwana Res. 25, 896&#x2013;909 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR133\" id=\"ref-link-section-d198960051e2121\" rel=\"nofollow noopener\" target=\"_blank\">133<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2e<\/a>).<\/p>\n<p>The rise of complex Phanerozoic ecosystems further shaped biogeochemical cycles. The colonization of land by plants in the Ordovician to early Devonian significantly amplified the continental supply of weathering products<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 134\" title=\"Dahl, T. W. &amp; Arens, S. K. M. The impacts of land plant evolution on Earth&#x2019;s climate and oxygenation state &#x2013; An interdisciplinary review. Chem. Geol. 547, 119665 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR134\" id=\"ref-link-section-d198960051e2132\" rel=\"nofollow noopener\" target=\"_blank\">134<\/a>, including bio-limiting nutrients (e.g., phosphorus), to the oceans, promoting organic carbon burial and further oxygenation of the atmosphere and deep ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 135\" title=\"Tostevin, R. &amp; Mills, B. J. W. Reconciling proxy records and models of Earth&#x2019;s oxygenation during the Neoproterozoic and Palaeozoic. Interface Focus 10, 20190137 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR135\" id=\"ref-link-section-d198960051e2136\" rel=\"nofollow noopener\" target=\"_blank\">135<\/a>. This established or helped sustain a fully oxygenated ocean with high levels of oxidants (including oxygen, sulfate, and nitrate) but low dissolved iron, similar to the modern (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2b, d<\/a>). The end-Permian mass extinction created ecological niches for modern eukaryotic phytoplankton to radiate in the Mesozoic, notably dinoflagellates, coccolithophores, and diatoms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 136\" title=\"Falkowski, P. G. et al. The evolution of modern eukaryotic phytoplankton. Science 305, 354&#x2013;360 (2004).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR136\" id=\"ref-link-section-d198960051e2143\" rel=\"nofollow noopener\" target=\"_blank\">136<\/a>. These groups might greatly enhance the efficiency of the biological pump and carbonate factory, increasing carbon export to deep sediments<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 136\" title=\"Falkowski, P. G. et al. The evolution of modern eukaryotic phytoplankton. Science 305, 354&#x2013;360 (2004).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR136\" id=\"ref-link-section-d198960051e2147\" rel=\"nofollow noopener\" target=\"_blank\">136<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 137\" title=\"Ridgwell, A. Evolution of the ocean&#x2019;s &#x201C;biological pump&#x201D;. Proc. Natl. Acad. Sci. USA 108, 16485&#x2013;16486 (2011).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR137\" id=\"ref-link-section-d198960051e2150\" rel=\"nofollow noopener\" target=\"_blank\">137<\/a>. Moreover, diatoms began to efficiently strip dissolved silica from seawater, resulting in low silica oceans<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 97\" title=\"Conley, D. J. et al. Biosilicification drives a decline of dissolved Si in the oceans through geologic time. Front. Mar. Sci. 4, 397 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR97\" id=\"ref-link-section-d198960051e2155\" rel=\"nofollow noopener\" target=\"_blank\">97<\/a>. Stepwise, these biological innovations drove the ocean toward its modern chemical state (as detailed in Section 1).<\/p>\n<p>In summary, the chemical evolution of Earth\u2019s hydrosphere and its habitability have been jointly dictated by the co-evolution of geosphere and biosphere (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). The secular cooling of the Earth reduced volcanic and hydrothermal activities, while the emergence of land increased continental weathering and marine upwelling supplies of nutrients. These tandem processes drew down atmospheric CO2, raised ocean pH, and altered nutrient inventories (e.g., sulfur, phosphorus, nitrogen, and trace metals), which in turn controlled biological primary productivity and metabolic evolution. The subsequent rise of complex life amplified organic carbon burial, leading to a fully oxygenated surface environment that ultimately ushered in the robust and complex ecosystems of today.<\/p>\n<p>While this Section focuses on the generic evolutionary history of Earth\u2019s oceans, it is crucial to acknowledge the persistently important roles of diverse aquatic environments, such as lakes, rivers, hot springs, and hydrothermal systems. Such environments would have been established as soon as the surface hydrosphere appeared, although their relative abundance may have fluctuated considerably over geological time. Furthermore, evidence suggests that the colonization of terrestrial aquatic habitats may have begun as early as the Archean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 138\" title=\"Zaharescu, D. G. et al. Biological Weathering in the Terrestrial System. in Biogeochemical Cycles: Ecological Drivers and Environmental Impact 1&#x2013;32. (Wiley, Hoboken, 2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR138\" id=\"ref-link-section-d198960051e2176\" rel=\"nofollow noopener\" target=\"_blank\">138<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 139\" title=\"Beraldi-Campesi, H. Early life on land and the first terrestrial ecosystems. Ecol. Process. 2, 1 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR139\" id=\"ref-link-section-d198960051e2179\" rel=\"nofollow noopener\" target=\"_blank\">139<\/a>, with these environments becoming increasingly important following the emergence of complex life (e.g., plants<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 140\" title=\"Donoghue, P. C. J., Harrison, C. J., Paps, J. &amp; Schneider, H. The evolutionary emergence of land plants. Curr. Biol. 31, R1281&#x2013;R1298 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR140\" id=\"ref-link-section-d198960051e2183\" rel=\"nofollow noopener\" target=\"_blank\">140<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 141\" title=\"Boyce, C. K. &amp; Nelsen, M. P. Terrestrialization: toward a shared framework for ecosystem evolution. Paleobiology 51, 174&#x2013;194 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR141\" id=\"ref-link-section-d198960051e2186\" rel=\"nofollow noopener\" target=\"_blank\">141<\/a>; Fig.<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2e<\/a>). The biological activity within these environments, in turn, significantly shaped the habitability of the terrestrial realm<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 138\" title=\"Zaharescu, D. G. et al. Biological Weathering in the Terrestrial System. in Biogeochemical Cycles: Ecological Drivers and Environmental Impact 1&#x2013;32. (Wiley, Hoboken, 2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR138\" id=\"ref-link-section-d198960051e2193\" rel=\"nofollow noopener\" target=\"_blank\">138<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 141\" title=\"Boyce, C. K. &amp; Nelsen, M. P. Terrestrialization: toward a shared framework for ecosystem evolution. Paleobiology 51, 174&#x2013;194 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR141\" id=\"ref-link-section-d198960051e2196\" rel=\"nofollow noopener\" target=\"_blank\">141<\/a>. However, compared to the marine record, the long-term chemical evolution of these continental and niche water bodies is poorly constrained, due to a general scarcity of continuous sedimentary archives. This challenge is compounded by their intrinsic chemical heterogeneity, which is highly sensitive to local environmental and hydrological conditions.<\/p>\n<p>Section 3. Evolution of hydrosphere and habitability on Mars<\/p>\n<p>The present-day surface of Mars is forbiddingly cold and dry, making it an unlikely habitat for life. Orbital and rover exploration over the past half century, however, demonstrates that Mars was not always like this<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 142\" title=\"Jakosky, B. M. et al. The history of Martian water during the Hesperian and Amazonian epochs. Icarus 443, 116782 (2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR142\" id=\"ref-link-section-d198960051e2208\" rel=\"nofollow noopener\" target=\"_blank\">142<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 143\" title=\"Carr, M. H. &amp; Head, J. W. Geologic history of Mars. Earth Planet. Sci. Lett. 294, 185&#x2013;203 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR143\" id=\"ref-link-section-d198960051e2211\" rel=\"nofollow noopener\" target=\"_blank\">143<\/a>. Early in its history, our planetary neighbor boasted rivers and lakes, perhaps even an ocean (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3<\/a>). Early orbiters, in particular those of NASA\u2019s Mariner and Viking programs, clearly imaged drainage channels much like those incised by flowing water on Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 144\" title=\"Baker, V. R. et al. The Channels of Mars. (University of Texas Press, Austin, 1982).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR144\" id=\"ref-link-section-d198960051e2218\" rel=\"nofollow noopener\" target=\"_blank\">144<\/a>. Ensuing spectroscopic mapping by NASA and ESA orbiters documented the widespread distribution of clays, hematite and other minerals generally formed in association with liquid water<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 145\" title=\"Ehlmann, B. L. &amp; Edwards, C. S. Mineralogy of the Martian surface. Annu. Rev. Earth Planet. Sci. 42, 291&#x2013;315 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR145\" id=\"ref-link-section-d198960051e2222\" rel=\"nofollow noopener\" target=\"_blank\">145<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 146\" title=\"Bibring, J.-P. et al. Global mineralogical and aqueous Mars history derived from OMEGA\/Mars Express data. Science 312, 400&#x2013;404 (2006).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR146\" id=\"ref-link-section-d198960051e2225\" rel=\"nofollow noopener\" target=\"_blank\">146<\/a>. Moreover, observations by the Mars Odyssey orbiter documented high concentrations of hydrogen at or just beneath the Martian surface, especially at higher latitudes<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 147\" title=\"Boynton, W. V. et al. Distribution of hydrogen in the near surface of Mars: Evidence for subsurface ice deposits. Science 297, 81&#x2013;85 (2002).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR147\" id=\"ref-link-section-d198960051e2229\" rel=\"nofollow noopener\" target=\"_blank\">147<\/a>. The interpretation of these observations as a signature for water ice was corroborated by orbital imaging of surface features similar to those associated with near-surface ground ice on Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 17\" title=\"Luzzi, E. et al. Geomorphological evidence of near-surface ice at candidate landing sites in northern Amazonis Planitia, Mars. J. Geophys. Res. Planets 130, e2024JE008724 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR17\" id=\"ref-link-section-d198960051e2234\" rel=\"nofollow noopener\" target=\"_blank\">17<\/a>.<\/p>\n<p>Fig. 3: Temporal evolution of key surface and interior processes on Mars.<img decoding=\"async\" aria-describedby=\"figure-3-desc\" src=\"https:\/\/www.newsbeep.com\/ie\/wp-content\/uploads\/2026\/07\/41467_2026_74955_Fig3_HTML.png\" alt=\"Fig. 3: Temporal evolution of key surface and interior processes on Mars.\" loading=\"lazy\" width=\"685\" height=\"677\"\/><\/p>\n<p>a Schematic evolution of Martian atmospheric density, which remains poorly constrained. Geological and geochemical evidence suggests that early Mars may have sustained a denser atmosphere than today, at least episodically. b Conceptual evolution history of liquid-water reservoirs on Mars. Early Mars may have hosted relatively stable surface-water bodies, including lakes and channel-forming fluvial systems. As atmospheric lost progressively, stable surface liquid water became increasingly difficult to sustain, and any remaining liquid water was likely restricted mainly to the subsurface. c Timing of major impact events on Mars. Question marks indicate uncertainties in the duration and intensity of impact episodes. d Evolution of major volcanic episodes on Mars. e Representative secondary minerals formed through water-rock interactions on Mars. Based on data from Sasselov et al<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 218\" title=\"Sasselov, D. D., Grotzinger, J. P. &amp; Sutherland, J. D. The origin of life as a planetary phenomenon. Sci. Adv. 6, eaax3419 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR218\" id=\"ref-link-section-d198960051e2264\" rel=\"nofollow noopener\" target=\"_blank\">218<\/a>., Wordsworth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 12\" title=\"Wordsworth, R. D. The climate of early Mars. Annu. Rev. Earth Planet. Sci. 44, 381&#x2013;408 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR12\" id=\"ref-link-section-d198960051e2269\" rel=\"nofollow noopener\" target=\"_blank\">12<\/a>, Ehlmann et al<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 244\" title=\"Ehlmann, B. L. et al. Subsurface water and clay mineral formation during the early history of Mars. Nature 479, 53&#x2013;60 (2011).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR244\" id=\"ref-link-section-d198960051e2273\" rel=\"nofollow noopener\" target=\"_blank\">244<\/a>., Ramirez and Craddock<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 245\" title=\"Ramirez, R. M. &amp; Craddock, R. A. The geological and climatological case for a warmer and wetter early Mars. Nat. Geosci. 11, 230&#x2013;237 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR245\" id=\"ref-link-section-d198960051e2277\" rel=\"nofollow noopener\" target=\"_blank\">245<\/a>.<\/p>\n<p>With the successful landing and deployment of the NASA rovers Spirit and Opportunity in 2004, it became possible for the first time to study Mars and its history in much the same way that field geologists do on Earth, via the physical and geochemical investigation of ancient Martian sedimentary rocks exposed in outcrop, largely associated with meteorite impacts (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3c<\/a>). Opportunity\u2019s exploration of sedimentary rocks in Meridiani planum found evidence of ripple marks known to be shaped by flowing water on Earth; Opportunity also discovered abundant evaporite minerals that record rock-water interactions (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3e<\/a>). Early diagenetic hematite concretions additionally record iron oxidation associated with ground water percolation. Taken together, evidence from Meridiani planum indicates that Mars hosted a regionally extensive arid, acidic and oxidizing environment more than three billion years ago<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 148\" title=\"Squyres, S. W. &amp; Knoll, A. H. Sedimentary rocks at Meridiani Planum: Origin, diagenesis, and implications for life on Mars. Earth Planet. Sci. Lett. 240, 1&#x2013;10 (2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR148\" id=\"ref-link-section-d198960051e2298\" rel=\"nofollow noopener\" target=\"_blank\">148<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 149\" title=\"Jolliff, B. L. et al. Mars Exploration Rover Opportunity: Water and Other Volatiles on Ancient Mars. in Volatiles in the Martian Crust. 285&#x2013;328Elsevier: Amsterdam, 2019.\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR149\" id=\"ref-link-section-d198960051e2301\" rel=\"nofollow noopener\" target=\"_blank\">149<\/a>.<\/p>\n<p>Opportunity\u2019s twin rover Spirit, operating on the other side of the planet, found further evidence of rock-water interactions but within older strata<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 150\" title=\"Arvidson, R. E. et al. Spirit Mars rover mission to the Columbia Hills, Gusev Crater: Mission overview and selected results from the Cumberland Ridge to Home Plate. J. Geophys. Res. 113, E12S33 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR150\" id=\"ref-link-section-d198960051e2308\" rel=\"nofollow noopener\" target=\"_blank\">150<\/a>. Notably, a failed wheel, dragged through surficial regolith in Gusev Crater, fortuitously discovered a white material beneath surficial dust \u2013 amorphous silica, likely formed by the alteration of basalt by hydrothermal fluids<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 151\" title=\"Squyres, S. W. et al. Detection of silica-rich deposits on Mars. Science 320, 1063&#x2013;1067 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR151\" id=\"ref-link-section-d198960051e2312\" rel=\"nofollow noopener\" target=\"_blank\">151<\/a>. Rovers Opportunity and Spirit, thus, corroborated and enriched orbital evidence that liquid water once existed at the Martian surface. But they did more than that, providing insights into the properties and persistence of early water bodies. Water is key to life as we know it, but not all water bodies are habitable. In particular, aw sets an environmental limit for essentially all species found on Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 27\" title=\"Stevenson, A. et al. Is there a common water-activity limit for the three domains of life? ISME J. 9, 1333&#x2013;1351 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR27\" id=\"ref-link-section-d198960051e2322\" rel=\"nofollow noopener\" target=\"_blank\">27<\/a>. Ions and dissolved molecules bond weakly with H2O, limiting the amount of water available for physical and biological activities. Every species has a lower limit of tolerance for aw, and calculation of aw in some ancient Martian waters suggests that evaporation could drive it to low levels that may become uninhabitable for most known life on Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 25\" title=\"Tosca, N. J., Knoll, A. H. &amp; McLennan, S. M. Water activity and the challenge for life on early Mars. Science 320, 1204&#x2013;1207 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR25\" id=\"ref-link-section-d198960051e2341\" rel=\"nofollow noopener\" target=\"_blank\">25<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 26\" title=\"Stevens, A. H. &amp; Cockell, C. S. The water activity of mars-relevant multicomponent brines: The changing influence of perchlorate on habitability over time. Planet. Sci. J. 4, 6 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR26\" id=\"ref-link-section-d198960051e2344\" rel=\"nofollow noopener\" target=\"_blank\">26<\/a>.<\/p>\n<p>The persistence of surficial water is another important consideration for ancient habitability within the Martian environment. Geochemical observations of jarosite, amorphous silica and smectite clays in ancient Martian sedimentary rocks provide compelling evidence for liquid water at their time of formation (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3e<\/a>). However, the fact that continued interactions with water generally transform such minerals into one other, more stable species on Earth suggests that on Mars these minerals have not seen much water since they formed billions of years ago<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 152\" title=\"Tosca, N. J. &amp; Knoll, A. H. Juvenile chemical sediments and the long term persistence of water at the surface of Mars. Earth Planet. Sci. Lett. 286, 379&#x2013;386 (2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR152\" id=\"ref-link-section-d198960051e2354\" rel=\"nofollow noopener\" target=\"_blank\">152<\/a>, thus suggesting Mars has been persistently dry for a long time.<\/p>\n<p>In the wake of Opportunity\u2019s and Spirit\u2019s pioneering exploration, two more recent rovers have greatly improved our understanding of Martian surficial environments in both space and time. Since 2012, Curiosity has explored a thick succession of sedimentary rocks exposed in Gale crater, and, since 2020, Perseverance has been investigating sedimentary deposits in and around an ancient delta preserved in Jezero crater. Mars chronology is currently in a state of flux, as confidence in the Late Heavy Bombardment hypothesis has waned (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3c<\/a>), but in-situ geochronological analysis indicates that sedimentary rocks within Gale crater are 4.1 &#8211; 3.6 billion years old<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 153\" title=\"Farley, K. A. et al. In situ radiometric and exposure age dating of the martian surface. Science 343, 1247166 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR153\" id=\"ref-link-section-d198960051e2365\" rel=\"nofollow noopener\" target=\"_blank\">153<\/a>, older, on stratigraphic grounds than the evaporite-rich rocks in Meridiani planum (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3e<\/a>). Deltaic and delta-front deposits in Jezero crater may be comparably old\u2013again, both older and environmentally distinct from those discovered in Meridiani planum. At both newer localities, the rovers discovered lacustrine sedimentary rocks that not only preserve sedimentological and chemical evidence of water, but also have organic molecules, phosphorus, nitrogen, redox gradients, and a more bio-friendly aw, collectively opening the possibility that these environments were, in principle, habitable<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 154\" title=\"Grotzinger, J. P. et al. A habitable fluvio-lacustrine environment at Yellowknife Bay, Gale Crater, Mars. Science 343, 1242777 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR154\" id=\"ref-link-section-d198960051e2378\" rel=\"nofollow noopener\" target=\"_blank\">154<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 155\" title=\"Eigenbrode, J. L. et al. Organic matter preserved in 3-billion-year-old mudstones at Gale crater, Mars. Science 360, 1096&#x2013;1101 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR155\" id=\"ref-link-section-d198960051e2381\" rel=\"nofollow noopener\" target=\"_blank\">155<\/a>. Indeed, in 2025, a silty sedimentary rock bed in Jezero crater was found to contain abundant small, irregular patches containing vivianite, an iron phosphate mineral, and greigite, an iron sulfide, interpreted as a potential biosignature<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 156\" title=\"Hurowitz, J. A. et al. Redox-driven mineral and organic associations in Jezero Crater, Mars. Nature 645, 332&#x2013;340 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR156\" id=\"ref-link-section-d198960051e2386\" rel=\"nofollow noopener\" target=\"_blank\">156<\/a>. The pattern is indeed suggestive, although other explanations are possible. Rock samples were cached for eventual return to Earth, and if and when that happens, scientists may be able to discriminate among various hypotheses for the rock\u2019s origins.<\/p>\n<p>With questions of biology in mind, it is useful to return to questions of persistence. In comparison to the Earth, the window of habitability on early Mars may have been cut short. For decades, Martian environmental history was characterized in terms of a warm wet interval that persisted for at least hundreds of millions of years in Mars\u2019 youth. However, today, there is strong reason to interpret the history of Martian surface water as one of episodic warmth and surface water, with clement environments persisting for perhaps tens of thousands of years every so often during the planet\u2019s early history. Several hypotheses have been proposed to explain the episodicity of surface water, including transient warmth and wetness engendered by large impacts and volcanic eruptions (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3c,<\/a>d), as well as episodic shifts in obliquity<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 157\" title=\"Wordsworth, R. et al. A coupled model of episodic warming, oxidation and geochemical transitions on early Mars. Nat. Geosci. 14, 127&#x2013;132 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR157\" id=\"ref-link-section-d198960051e2398\" rel=\"nofollow noopener\" target=\"_blank\">157<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 158\" title=\"Kite, E. S. &amp; Conway, S. Geological evidence for multiple climate transitions on Early Mars. Nat. Geosci. 17, 10&#x2013;19 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR158\" id=\"ref-link-section-d198960051e2401\" rel=\"nofollow noopener\" target=\"_blank\">158<\/a>. In any event, drivers of transient warmth and wetness declined in frequency and size, eventually resulting in the Mars we see today.<\/p>\n<p>So, what happened to Mars\u2019 surface water? Isotopic ratios of various gases support the hypothesis that large impacts and hydrodynamic escape removed the primordial atmosphere, and once Mars\u2019 short-lived magnetic field dissipated, solar wind interactions stripped away remaining or secondary atmospheric gases through time<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 159\" title=\"Jakosky, B. M. &amp; Hallis, L. J. Fate of an earth-like water inventory on Mars. J. Geophys. Res. Planets 129, e2023JE008159 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR159\" id=\"ref-link-section-d198960051e2408\" rel=\"nofollow noopener\" target=\"_blank\">159<\/a>. These data also indicate that much of the Martian atmosphere-surface H2O reservoir was lost rapidly, perhaps as early as ~3 billion years ago<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 160\" title=\"Scheller, E. L., Ehlmann, B. L., Hu, R., Adams, D. J. &amp; Yung, Y. L. Long-term drying of Mars by sequestration of ocean-scale volumes of water in the crust. Science 372, 56&#x2013;62 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR160\" id=\"ref-link-section-d198960051e2414\" rel=\"nofollow noopener\" target=\"_blank\">160<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig3\" rel=\"nofollow noopener\" target=\"_blank\">3a<\/a>). In contrast to Earth\u2019s subduction-driven mode of volatile recycling between its interior and surface, the lack of a similar mechanism on Mars may have set it on an irreversible path toward desiccation as volatiles were permanently lost to its ancient crust through alteration processes<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 160\" title=\"Scheller, E. L., Ehlmann, B. L., Hu, R., Adams, D. J. &amp; Yung, Y. L. Long-term drying of Mars by sequestration of ocean-scale volumes of water in the crust. Science 372, 56&#x2013;62 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR160\" id=\"ref-link-section-d198960051e2421\" rel=\"nofollow noopener\" target=\"_blank\">160<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 161\" title=\"Mustard, J. F. et al. Sequestration of Volatiles in the Martian Crust Through Hydrated Minerals: A Significant Planetary Reservoir of Water. in Volatiles in the Martian Crust 247&#x2013;263. (Elsevier, Amsterdam, 2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR161\" id=\"ref-link-section-d198960051e2424\" rel=\"nofollow noopener\" target=\"_blank\">161<\/a>. Some water vapor would also have been lost by photodissociation of water vapor or ice in the upper atmosphere<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 162\" title=\"Jakosky, B. M. et al. Loss of the Martian atmosphere to space: Present-day loss rates determined from MAVEN observations and integrated loss through time. Icarus 315, 146&#x2013;157 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR162\" id=\"ref-link-section-d198960051e2429\" rel=\"nofollow noopener\" target=\"_blank\">162<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 163\" title=\"Chaffin, M. S., Deighan, J., Schneider, N. M. &amp; Stewart, A. I. F. Elevated atmospheric escape of atomic hydrogen from Mars induced by high-altitude water. Nat. Geosci. 10, 174&#x2013;178 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR163\" id=\"ref-link-section-d198960051e2432\" rel=\"nofollow noopener\" target=\"_blank\">163<\/a>, resulting in hydrogen escape to space while leaving oxygen gas to be consumed by the oxidation of reduced iron at the surface. Of course, as noted above, a considerable amount of H2O remains as ice within surface rocks and regolith, as well as water ice in polar ice caps. Recent geophysical data that are interpreted as requiring a large reservoir of water at depths of 11 to 20 kilometers below the Martian surface<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 164\" title=\"Wright, V., Morzfeld, M. &amp; Manga, M. Liquid water in the Martian mid-crust. Proc. Natl. Acad. Sci. USA 121, e2409983121 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR164\" id=\"ref-link-section-d198960051e2438\" rel=\"nofollow noopener\" target=\"_blank\">164<\/a> can be also explained without liquid water<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 165\" title=\"Xiao, W., Pan, L., Wang, Y. &amp; Li, J. Liquid water might not be the only answer: Evaluating pore-filling materials in the Martian crust. Proc. Natl. Acad. Sci. USA 122, e2503071122 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR165\" id=\"ref-link-section-d198960051e2442\" rel=\"nofollow noopener\" target=\"_blank\">165<\/a> and so remain controversial. In summary, while present day Mars is sufficiently cold that liquid water is not stable at its surface, during its early history it supported episodic rivers and lakes, some of them at least transiently habitable.<\/p>\n<p>Section 4. Subsurface oceans in icy bodies of the outer solar system<\/p>\n<p>Beyond the terrestrial planets, our solar system hosts a diverse population of ocean worlds among dwarf planets and icy satellites, including Ceres, Europa, Ganymede, Callisto, Enceladus, Titan, Triton, and Pluto (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). In stark contrast to Earth\u2019s surface oceans, these extraterrestrial bodies harbor their liquid water reservoirs beneath thick icy crusts, often sandwiched between layer(s) of ice and rock (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig4\" rel=\"nofollow noopener\" target=\"_blank\">4<\/a>). Some subsurface oceans are believed to have persisted for at least 107 years<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Courville, S. W., Castillo-Rogez, J. C., Daswani, M. M., Robare, J. &amp; O&#x2019;Rourke, J. G. Core metamorphism controls the dynamic habitability of mid-sized ocean worlds&#x2014;The case of Ceres. Sci. Adv. 11, eadt3283 (2025).\" href=\"#ref-CR166\" id=\"ref-link-section-d198960051e2462\">166<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Trinh, K. T., Bierson, C. J., O&#x2019;Rourke, J. G. &amp; al., et. Slow evolution of Europa&#x2019;s interior: Metamorphic ocean origin, delayed metallic core formation, and limited seafloor volcanism. Sci. Adv. 9, eadf3955 (2023).\" href=\"#ref-CR167\" id=\"ref-link-section-d198960051e2462_1\">167<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Tobie, G., Grasset, O., Lunine, J. I., Mocquet, A. &amp; Sotin, C. Titan&#x2019;s internal structure inferred from a coupled thermal-orbital model. Icarus 175, 496&#x2013;502 (2005).\" href=\"#ref-CR168\" id=\"ref-link-section-d198960051e2462_2\">168<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 169\" title=\"Choblet, G. et al. Powering prolonged hydrothermal activity inside Enceladus. Nat. Astron. 1, 841&#x2013;847 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR169\" id=\"ref-link-section-d198960051e2465\" rel=\"nofollow noopener\" target=\"_blank\">169<\/a>, sustained by heat from radioactive decay and\/or tidal dissipation<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 19\" title=\"Nimmo, F. &amp; Pappalardo, R. T. Ocean worlds in the outer solar system. J. Geophys. Res. Planets 121, 1378&#x2013;1399 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR19\" id=\"ref-link-section-d198960051e2469\" rel=\"nofollow noopener\" target=\"_blank\">19<\/a>. There are also strong signs of past or ongoing hydrothermal activities in some planetary bodies, particularly Enceladus<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 170\" title=\"Hsu, H.-W. et al. Ongoing hydrothermal activities within Enceladus. Nature 519, 207&#x2013;210 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR170\" id=\"ref-link-section-d198960051e2474\" rel=\"nofollow noopener\" target=\"_blank\">170<\/a>. Moreover, remote geochemical analyses of their surface and plume materials suggest availability of bio-essential nutrients and organic species in the oceans (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). These lines of evidence jointly indicate that these subsurface oceans are likely habitable and potentially conducive to the origin and survival of life. This section synthesizes the up to date understanding of the ocean chemistry and habitability of four key representatives: Ceres, Europa, Enceladus, and Titan.<\/p>\n<p>Fig. 4: Interior structures and possible material exchange pathways in ocean-bearing icy moons.<img decoding=\"async\" aria-describedby=\"figure-4-desc\" src=\"https:\/\/www.newsbeep.com\/ie\/wp-content\/uploads\/2026\/07\/41467_2026_74955_Fig4_HTML.png\" alt=\"Fig. 4: Interior structures and possible material exchange pathways in ocean-bearing icy moons.\" loading=\"lazy\" width=\"685\" height=\"399\"\/><\/p>\n<p>This schematic illustration compares two major types of ocean-bearing icy moons: a smaller icy moons, such as Enceladus and Europa, in which a subsurface ocean may directly overlie a poorly differentiated silicate interior; and (b) larger, more strongly differentiated bodies, such as Titan and Ganymede, in which high-pressure ice layers separate the ocean from the deep rocky interior. Chemical species produced at the surface or within the interior may be exchanged through ice-shell recycling or subduction, ocean circulation, plume activity, as well as convection and melting within high-pressure ice layers. These possible transport processes regulate ocean chemistry and, consequently, the habitability of subsurface oceans. The structure is not shown to scale.<\/p>\n<p>Table 2 Four representative icy worlds with putative subsurface liquid water bodies in the solar systemCeres: a relic ocean world<\/p>\n<p>Ceres, a dwarf planet in the asteroid belt, is a highly hydrated body. Our understanding of Ceres was revolutionized by the Dawn spacecraft (2015-2018). Thermal evolution models suggest that Ceres once hosted a global subsurface ocean, formed early in its history by the metamorphic dehydration of hydrated materials<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 166\" title=\"Courville, S. W., Castillo-Rogez, J. C., Daswani, M. M., Robare, J. &amp; O&#x2019;Rourke, J. G. Core metamorphism controls the dynamic habitability of mid-sized ocean worlds&#x2014;The case of Ceres. Sci. Adv. 11, eadt3283 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR166\" id=\"ref-link-section-d198960051e2803\" rel=\"nofollow noopener\" target=\"_blank\">166<\/a>. While this global ocean likely froze due to limited radiogenic or tidal heating, evidence points to the existence of regional brines in the subsurface<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 171\" title=\"Castillo-Rogez, J. et al. Concepts for the future exploration of dwarf planet Ceres&#x2019; habitability. Planet. Sci. J. 3, 41 (2022).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR171\" id=\"ref-link-section-d198960051e2807\" rel=\"nofollow noopener\" target=\"_blank\">171<\/a>. These brines are implicated in forming recent geological features and precipitating salts and secondary minerals on the surface. The brine is inferred to be reducing and alkaline, as indicated by observations of Fe(II)-bearing minerals and brucite<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 172\" title=\"Milliken, R. E. &amp; Rivkin, A. S. Brucite and carbonate assemblages from altered olivine-rich materials on Ceres. Nat. Geosci. 2, 258&#x2013;261 (2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR172\" id=\"ref-link-section-d198960051e2811\" rel=\"nofollow noopener\" target=\"_blank\">172<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). The presence of ammonium-bearing phyllosilicates and carbonate salts (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>) further suggests considerable levels of dissolved NH3 and carbonate ions<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 173\" title=\"Ammannito, E. et al. Distribution of phyllosilicates on the surface of Ceres. Science 353, aaf4279 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR173\" id=\"ref-link-section-d198960051e2824\" rel=\"nofollow noopener\" target=\"_blank\">173<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 174\" title=\"Castillo-Rogez, J. et al. Insights into Ceres&#x2019;s evolution from surface composition. Meteorit. Planet. Sci. 53, 1820&#x2013;1843 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR174\" id=\"ref-link-section-d198960051e2827\" rel=\"nofollow noopener\" target=\"_blank\">174<\/a>. Moreover, surface analyses indicate a high abundance of organic compounds (potentially up to 30\u2009wt%)<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 175\" title=\"De Sanctis, M. C. et al. Localized aliphatic organic material on the surface of Ceres. Science 355, 719&#x2013;722 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR175\" id=\"ref-link-section-d198960051e2831\" rel=\"nofollow noopener\" target=\"_blank\">175<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>), likely from a large subsurface reservoir. In summary, Ceres may host relict brines whose chemistry has been largely shaped by serpentinization reactions<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 176\" title=\"De Sanctis, M. C. et al. Relict ocean worlds: Ceres. Space Sci. Rev. 216, 60 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR176\" id=\"ref-link-section-d198960051e2838\" rel=\"nofollow noopener\" target=\"_blank\">176<\/a>, analogous to terrestrial serpentinizing systems.<\/p>\n<p>Europa: a tidally-locked ocean world<\/p>\n<p>Unlike Ceres, Europa is robustly inferred to host a global, long-lived ocean sandwiched between its icy shell and partially differentiated core<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 177\" title=\"Kivelson, M. G. et al. Galileo magnetometer measurements: a stronger case for a subsurface ocean at Europa. Science 289, 1340&#x2013;1343 (2000).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR177\" id=\"ref-link-section-d198960051e2850\" rel=\"nofollow noopener\" target=\"_blank\">177<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). This ocean likely formed early via dehydration of hydrated minerals<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 167\" title=\"Trinh, K. T., Bierson, C. J., O&#x2019;Rourke, J. G. &amp; al., et. Slow evolution of Europa&#x2019;s interior: Metamorphic ocean origin, delayed metallic core formation, and limited seafloor volcanism. Sci. Adv. 9, eadf3955 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR167\" id=\"ref-link-section-d198960051e2857\" rel=\"nofollow noopener\" target=\"_blank\">167<\/a> and is sustained today by intense tidal heating<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 178\" title=\"Bierson, C. J. &amp; Steinbr&#xFC;gge, G. Tidal heating did not dry out Io and Europa. Planet. Sci. J. 2, 89 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR178\" id=\"ref-link-section-d198960051e2861\" rel=\"nofollow noopener\" target=\"_blank\">178<\/a>. While transient water plumes have been observed<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 179\" title=\"Roth, L. et al. Transient water vapor at Europa&#x2019;s south pole. Science 343, 171&#x2013;174 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR179\" id=\"ref-link-section-d198960051e2865\" rel=\"nofollow noopener\" target=\"_blank\">179<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 180\" title=\"Jia, X., Kivelson, M. G., Khurana, K. K. &amp; Kurth, W. S. Evidence of a plume on Europa from Galileo magnetic and plasma wave signatures. Nat. Astron. 2, 459&#x2013;464 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR180\" id=\"ref-link-section-d198960051e2868\" rel=\"nofollow noopener\" target=\"_blank\">180<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>), the lack of direct sampling means its composition must be inferred from remote sensing of the surface. Current models suggest a water mass dominated by Na+, Mg2+, and Cl-<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 181\" title=\"Johnson, P. V., Hodyss, R., Vu, T. H. &amp; Choukroun, M. Insights into Europa&#x2019;s ocean composition derived from its surface expression. Icarus 321, 857&#x2013;865 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR181\" id=\"ref-link-section-d198960051e2881\" rel=\"nofollow noopener\" target=\"_blank\">181<\/a> with K+ and Ca2+<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 182\" title=\"Zolotov, M. Y. &amp; Kargel, J. S. On the Chemical Composition of Europa&#x2019;s Icy Shell, Ocean, and Underlying Rocks. In Europa 431&#x2013;458. (University of Arizona Press, Tucson, 2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR182\" id=\"ref-link-section-d198960051e2889\" rel=\"nofollow noopener\" target=\"_blank\">182<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>), but a central controversy surrounds the origin of surface sulfates &#8211; whether they are sourced from the ocean or produced in situ by radiolysis of sulfides<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 183\" title=\"Zolotov, M. Y. &amp; Shock, E. L. Composition and stability of salts on the surface of Europa and their oceanic origin. J. Geophys. Res. Planets 106, 32815&#x2013;32827 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR183\" id=\"ref-link-section-d198960051e2896\" rel=\"nofollow noopener\" target=\"_blank\">183<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 184\" title=\"Brown, M. E. &amp; Hand, K. Salts and radiation products on the surface of Europa. Astron. J. 145, 110 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR184\" id=\"ref-link-section-d198960051e2899\" rel=\"nofollow noopener\" target=\"_blank\">184<\/a>. This ambiguity propagates into major uncertainties in the ocean\u2019s pH and redox state<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 182\" title=\"Zolotov, M. Y. &amp; Kargel, J. S. On the Chemical Composition of Europa&#x2019;s Icy Shell, Ocean, and Underlying Rocks. In Europa 431&#x2013;458. (University of Arizona Press, Tucson, 2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR182\" id=\"ref-link-section-d198960051e2903\" rel=\"nofollow noopener\" target=\"_blank\">182<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 185\" title=\"Pasek, M. A. &amp; Greenberg, R. Acidification of Europa&#x2019;s subsurface ocean as a consequence of oxidant delivery. Astrobiology 12, 151&#x2013;159 (2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR185\" id=\"ref-link-section-d198960051e2906\" rel=\"nofollow noopener\" target=\"_blank\">185<\/a>. Recent observations of surface CO2 (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>) have been interpreted as evidence for an acidic and oxidizing ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 186\" title=\"Trumbo, S. K. &amp; Brown, M. E. The distribution of CO2 on Europa indicates an internal source of carbon. Science 381, 931&#x2013;934 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR186\" id=\"ref-link-section-d198960051e2916\" rel=\"nofollow noopener\" target=\"_blank\">186<\/a>, despite of several alternative interpretations in lack of reliable constraints. However, if true, this condition would not favor the stability of organics from accretion or impact delivery<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 182\" title=\"Zolotov, M. Y. &amp; Kargel, J. S. On the Chemical Composition of Europa&#x2019;s Icy Shell, Ocean, and Underlying Rocks. In Europa 431&#x2013;458. (University of Arizona Press, Tucson, 2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR182\" id=\"ref-link-section-d198960051e2920\" rel=\"nofollow noopener\" target=\"_blank\">182<\/a>, which may partly explain the lack of detectable organic materials to date. Moreover, any organics emplaced on the surface would be subjected to intense ionizing radiation from Jupiter<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 187\" title=\"Meitzler, R. et al. Investigating Europa&#x2019;s radiation environment with the Europa Clipper radiation monitor. Space Sci. Rev. 219, 61 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR187\" id=\"ref-link-section-d198960051e2924\" rel=\"nofollow noopener\" target=\"_blank\">187<\/a>, undermining their survival in surface ice<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 188\" title=\"Nordheim, T. A., Hand, K. P. &amp; Paranicas, C. Preservation of potential biosignatures in the shallow subsurface of Europa. Nat. Astron. 2, 673&#x2013;679 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR188\" id=\"ref-link-section-d198960051e2928\" rel=\"nofollow noopener\" target=\"_blank\">188<\/a>. The surface radiation could also generate potent oxidizing species, such as O2 and H2O2 at Europa\u2019s surface<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 189\" title=\"Carlson, R. W. et al. Europa&#x2019;s Surface Composition. In Europa 283&#x2013;327. (University of Arizona Press, Tucson, 2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR189\" id=\"ref-link-section-d198960051e2939\" rel=\"nofollow noopener\" target=\"_blank\">189<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). Possible ice tectonics, driven by tidal flexing<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 190\" title=\"Kattenhorn, S. A. &amp; Prockter, L. M. Evidence for subduction in the ice shell of Europa. Nat. Geosci. 7, 762&#x2013;767 (2014).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR190\" id=\"ref-link-section-d198960051e2946\" rel=\"nofollow noopener\" target=\"_blank\">190<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig4\" rel=\"nofollow noopener\" target=\"_blank\">4a<\/a>), may help transport these oxidants to the subsurface ocean, providing oxidant substrates for possible chemosynthetic life<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 67\" title=\"Hand, K. P., Carlson, R. W. &amp; Chyba, C. F. Energy, chemical disequilibrium, and geological constraints on Europa. Astrobiology 7, 1006&#x2013;1022 (2007).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR67\" id=\"ref-link-section-d198960051e2954\" rel=\"nofollow noopener\" target=\"_blank\">67<\/a>. However, recent geophysical studies suggest that Europa\u2019s seafloor may not currently host active volcanism<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 191\" title=\"Green, A. P., Elder, C. M., Bland, M. T., Tackley, P. J. &amp; Byrne, P. K. No magmatic driving force for Europan sea-floor volcanism. Nat. Astron. 9, 640&#x2013;649 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR191\" id=\"ref-link-section-d198960051e2958\" rel=\"nofollow noopener\" target=\"_blank\">191<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 192\" title=\"Byrne, P. K. et al. Little to no active faulting likely at Europa&#x2019;s seafloor today. Nat. Commun. 17, 4 (2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR192\" id=\"ref-link-section-d198960051e2961\" rel=\"nofollow noopener\" target=\"_blank\">192<\/a> or significant ongoing water-rock interaction<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 167\" title=\"Trinh, K. T., Bierson, C. J., O&#x2019;Rourke, J. G. &amp; al., et. Slow evolution of Europa&#x2019;s interior: Metamorphic ocean origin, delayed metallic core formation, and limited seafloor volcanism. Sci. Adv. 9, eadf3955 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR167\" id=\"ref-link-section-d198960051e2965\" rel=\"nofollow noopener\" target=\"_blank\">167<\/a>, limiting the release of reductants and buildup of redox gradients necessary to sustain life on Europa<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 67\" title=\"Hand, K. P., Carlson, R. W. &amp; Chyba, C. F. Energy, chemical disequilibrium, and geological constraints on Europa. Astrobiology 7, 1006&#x2013;1022 (2007).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR67\" id=\"ref-link-section-d198960051e2969\" rel=\"nofollow noopener\" target=\"_blank\">67<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 193\" title=\"McCollom, T. M. Methanogenesis as a potential source of chemical energy for primary biomass production by autotrophic organisms in hydrothermal systems on Europa. J. Geophys. Res. Planets 104, 30729&#x2013;30742 (1999).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR193\" id=\"ref-link-section-d198960051e2972\" rel=\"nofollow noopener\" target=\"_blank\">193<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 194\" title=\"Zolotov, M. Y. &amp; Shock, E. L. A model for low-temperature biogeochemistry of sulfur, carbon, and iron on Europa. J. Geophys. Res. Planets 109, E06003 (2004).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR194\" id=\"ref-link-section-d198960051e2975\" rel=\"nofollow noopener\" target=\"_blank\">194<\/a>.<\/p>\n<p>Apart from tectonic activities, ocean circulation is another key control on the transport and mixing of surface-derived oxidants and interior-derived reductants, and thus on the buildup of chemical disequilibria for potential metabolisms. Unlike Earth\u2019s oceans, icy moons\u2019 subsurface oceans lack wind forcing and are thought to be driven primarily by buoyancy contrasts associated with basal heating and ice-shell melting and freezing<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 195\" title=\"Soderlund, K. M., Rovira-Navarro, M., Le Bars, M., Schmidt, B. E. &amp; Gerkema, T. The physical oceanography of ice-covered moons. Ann. Rev. Mar. Sci. 16, 25&#x2013;53 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR195\" id=\"ref-link-section-d198960051e2982\" rel=\"nofollow noopener\" target=\"_blank\">195<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 196\" title=\"Lobo, A. H., Thompson, A. F., Vance, S. D. &amp; Tharimena, S. A pole-to-equator ocean overturning circulation on Enceladus. Nat. Geosci. 14, 185&#x2013;189 (2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR196\" id=\"ref-link-section-d198960051e2985\" rel=\"nofollow noopener\" target=\"_blank\">196<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig4\" rel=\"nofollow noopener\" target=\"_blank\">4a<\/a>). Although this circulation may operate under a different and possibly less efficient mixing regime than that of Earth\u2019s ocean, it also governs the extent to which hydrothermal products and other potential biosignatures can be transported from the seafloor to the ice shell or surface<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 197\" title=\"Ames, F., Ferreira, D., Czaja, A. &amp; Masters, A. Ocean stratification impedes particulate transport to the plumes of Enceladus. Commun. Earth Environ. 6, 63 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR197\" id=\"ref-link-section-d198960051e2992\" rel=\"nofollow noopener\" target=\"_blank\">197<\/a>, thereby strongly influencing their detectability.<\/p>\n<p>Enceladus: an active ocean world<\/p>\n<p>Among icy bodies, Enceladus provides the most direct evidence of a subsurface ocean, i.e., the ongoing water plume observed by the Cassini spacecraft in its south polar region<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 198\" title=\"Porco, C. C. et al. Cassini observes the active south pole of Enceladus. Science 311, 1393&#x2013;1401 (2006).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR198\" id=\"ref-link-section-d198960051e3004\" rel=\"nofollow noopener\" target=\"_blank\">198<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). Geophysical models further indicate a global, long-lived liquid ocean (&gt; 108 years), sustained by tidal dissipation along with radiogenic heating<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 169\" title=\"Choblet, G. et al. Powering prolonged hydrothermal activity inside Enceladus. Nat. Astron. 1, 841&#x2013;847 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR169\" id=\"ref-link-section-d198960051e3013\" rel=\"nofollow noopener\" target=\"_blank\">169<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 199\" title=\"Nimmo, F., Neveu, M. &amp; Howett, C. Origin and evolution of Enceladus&#x2019;s tidal dissipation. Space Sci. Rev. 219, 57 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR199\" id=\"ref-link-section-d198960051e3016\" rel=\"nofollow noopener\" target=\"_blank\">199<\/a>. Detailed analyses of the emitted gases and ice grains by the Cassini\u2019s spectrometers reveal a reducing chemical environment rich in CO2, H2, CH4, and NH3, alongside a diverse suite of organic compounds<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Waite, J. H. Jr et al. Liquid water on Enceladus from observations of ammonia and 40Ar in the plume. Nature 460, 487&#x2013;490 (2009).\" href=\"#ref-CR200\" id=\"ref-link-section-d198960051e3029\">200<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Waite, J. H. et al. Cassini finds molecular hydrogen in the Enceladus plume: Evidence for hydrothermal processes. Science 356, 155&#x2013;159 (2017).\" href=\"#ref-CR201\" id=\"ref-link-section-d198960051e3029_1\">201<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 202\" title=\"Postberg, F. et al. Macromolecular organic compounds from the depths of Enceladus. Nature 558, 564&#x2013;568 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR202\" id=\"ref-link-section-d198960051e3032\" rel=\"nofollow noopener\" target=\"_blank\">202<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). The salt composition of the ocean, inferred from analyses of the emitted ice grains, is dominated by NaCl, KCl, and NaHCO3\/Na2CO3<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 203\" title=\"Postberg, F. et al. Sodium salts in E-ring ice grains from an ocean below the surface of Enceladus. Nature 459, 1098&#x2013;1101 (2009).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR203\" id=\"ref-link-section-d198960051e3045\" rel=\"nofollow noopener\" target=\"_blank\">203<\/a> (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>). Recently, phosphorus, a key limiting nutrient for life on Earth, has been revealed to accumulate to significant abundance (~mM) within Enceladus\u2019s ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 204\" title=\"Hao, J. et al. Abundant phosphorus expected for possible life in Enceladus&#x2019;s ocean. Proc. Natl. Acad. Sci. USA 119, e2201388119 (2022).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR204\" id=\"ref-link-section-d198960051e3052\" rel=\"nofollow noopener\" target=\"_blank\">204<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 205\" title=\"Postberg, F. et al. Detection of phosphates originating from Enceladus&#x2019;s ocean. Nature 618, 489&#x2013;493 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR205\" id=\"ref-link-section-d198960051e3055\" rel=\"nofollow noopener\" target=\"_blank\">205<\/a>, which is at least three orders of magnitude higher than modern levels in Earth\u2019s oceans (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Compared with other icy bodies, Enceladus has the most compelling evidence for ongoing high-temperature (&gt; 90 \u00b0C) hydrothermal activity with the detection of silica nanoparticles in the plume<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 170\" title=\"Hsu, H.-W. et al. Ongoing hydrothermal activities within Enceladus. Nature 519, 207&#x2013;210 (2015).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR170\" id=\"ref-link-section-d198960051e3063\" rel=\"nofollow noopener\" target=\"_blank\">170<\/a>, although recent geophysical model suggests the ocean stratification would impede particulate transport from the hydrothermal vents to the plumes of Enceladus<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 197\" title=\"Ames, F., Ferreira, D., Czaja, A. &amp; Masters, A. Ocean stratification impedes particulate transport to the plumes of Enceladus. Commun. Earth Environ. 6, 63 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR197\" id=\"ref-link-section-d198960051e3067\" rel=\"nofollow noopener\" target=\"_blank\">197<\/a>. Nevertheless, intensive water-rock interaction and potential hydrothermal activity would help build chemical and energetic gradients as well as establish mineral interfaces in the Enceladus ocean floor, which are essential for the origin and sustainability of life<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 104\" title=\"Hazen, R. M. &amp; Sverjensky, D. A. Mineral surfaces, geochemical complexities, and the origins of life. Cold Spring Harb. Perspect. Biol. 2, a002162 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR104\" id=\"ref-link-section-d198960051e3071\" rel=\"nofollow noopener\" target=\"_blank\">104<\/a>.<\/p>\n<p>Titan: an organic-rich ocean world<\/p>\n<p>Titan is unique among icy worlds because of its dense nitrogen-methane atmosphere (~1.5\u2009bar; Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>) and thick orange haze<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 206\" title=\"Tomasko, M. G. et al. Rain, winds and haze during the Huygens probe&#x2019;s descent to Titan&#x2019;s surface. Nature 438, 765&#x2013;778 (2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR206\" id=\"ref-link-section-d198960051e3086\" rel=\"nofollow noopener\" target=\"_blank\">206<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 207\" title=\"H&#xF6;rst, S. M. Titan&#x2019;s atmosphere and climate. J. Geophys. Res. Planets 122, 432&#x2013;482 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR207\" id=\"ref-link-section-d198960051e3089\" rel=\"nofollow noopener\" target=\"_blank\">207<\/a>. The successful Cassini-Huygens mission fundamentally advanced our understanding of Titan\u2019s complex organic chemistry (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>) and revealed a possibly global subsurface ocean beneath its icy shell<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 208\" title=\"Iess, L. et al. The tides of Titan. Science 337, 457&#x2013;459 (2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR208\" id=\"ref-link-section-d198960051e3096\" rel=\"nofollow noopener\" target=\"_blank\">208<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 209\" title=\"Sulaiman, A. H. et al. Enceladus and Titan: emerging worlds of the Solar System. Exp. Astron. 54, 849&#x2013;876 (2022).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR209\" id=\"ref-link-section-d198960051e3099\" rel=\"nofollow noopener\" target=\"_blank\">209<\/a>. Radar and landmark-tracking observations identified surface motions decoupled from the solid interior, indicating an outer ice shell floating above a liquid layer<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 210\" title=\"Lorenz, R. D. et al. Titan&#x2019;s rotation reveals an internal ocean and changing zonal winds. Science 319, 1649&#x2013;1651 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR210\" id=\"ref-link-section-d198960051e3103\" rel=\"nofollow noopener\" target=\"_blank\">210<\/a>. Gravity and rotational data further support a differentiated interior, consisting of a low-density outer shell overlying a denser rock\/ice core with an intervening global ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 208\" title=\"Iess, L. et al. The tides of Titan. Science 337, 457&#x2013;459 (2012).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR208\" id=\"ref-link-section-d198960051e3108\" rel=\"nofollow noopener\" target=\"_blank\">208<\/a>. However, latest analyses suggest that this liquid reservoir, if present today, may be spatially limited or replaced by a partially molten high-pressure ice layer rather than global ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 211\" title=\"Petricca, F. et al. Titan&#x2019;s strong tidal dissipation precludes a subsurface ocean. Nature 648, 556&#x2013;561 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR211\" id=\"ref-link-section-d198960051e3112\" rel=\"nofollow noopener\" target=\"_blank\">211<\/a>. The liquid layer, if it exists, is likely tens to hundreds of kilometers thick, situated between an ice Ih shell and high-pressure ice mantle<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 168\" title=\"Tobie, G., Grasset, O., Lunine, J. I., Mocquet, A. &amp; Sotin, C. Titan&#x2019;s internal structure inferred from a coupled thermal-orbital model. Icarus 175, 496&#x2013;502 (2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR168\" id=\"ref-link-section-d198960051e3116\" rel=\"nofollow noopener\" target=\"_blank\">168<\/a>. Its composition probably includes water, ammonia, dissolved salts (e.g., Na+, K+, Cl-), and possibly organics delivered from the surface and\/or subsurface<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 212\" title=\"Leitner, M. A. &amp; Lunine, J. I. Modeling early Titan&#x2019;s ocean composition. Icarus 333, 61&#x2013;70 (2019).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR212\" id=\"ref-link-section-d198960051e3127\" rel=\"nofollow noopener\" target=\"_blank\">212<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 213\" title=\"Lopes, R. M. C. et al. Cryovolcanism on titan: new results from cassini RADAR and VIMS. J. Geophys. Res. Planets 118, 416&#x2013;435 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR213\" id=\"ref-link-section-d198960051e3130\" rel=\"nofollow noopener\" target=\"_blank\">213<\/a>. Ammonia may act as an antifreeze, which helps maintain the liquid water layer(s) under Titan\u2019s frigid temperatures<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 214\" title=\"Choukroun, M., Grasset, O., Tobie, G. &amp; Sotin, C. Stability of methane clathrate hydrates under pressure: Influence on outgassing processes of methane on Titan. Icarus 205, 581&#x2013;593 (2010).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR214\" id=\"ref-link-section-d198960051e3134\" rel=\"nofollow noopener\" target=\"_blank\">214<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 215\" title=\"Chua, B. H. et al. Low-temperature specific heat capacity of water&#x2013;ammonia mixtures down to the eutectic. ACS Earth Space Chem. 7, 1971&#x2013;1979 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR215\" id=\"ref-link-section-d198960051e3137\" rel=\"nofollow noopener\" target=\"_blank\">215<\/a>. Potential energy sources that sustain the ocean include tidal heating and radiogenic decay, although the energy fluxes are modest compared to other active ocean worlds<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 168\" title=\"Tobie, G., Grasset, O., Lunine, J. I., Mocquet, A. &amp; Sotin, C. Titan&#x2019;s internal structure inferred from a coupled thermal-orbital model. Icarus 175, 496&#x2013;502 (2005).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR168\" id=\"ref-link-section-d198960051e3141\" rel=\"nofollow noopener\" target=\"_blank\">168<\/a>. Nevertheless, several dynamic processes, such as cryovolcanism, impact-driven melting, and diapiric transport, are proposed to facilitate the exchange of material and energy between Titan\u2019s surface organic reservoir and its interior ocean<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 207\" title=\"H&#xF6;rst, S. M. Titan&#x2019;s atmosphere and climate. J. Geophys. Res. Planets 122, 432&#x2013;482 (2017).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR207\" id=\"ref-link-section-d198960051e3145\" rel=\"nofollow noopener\" target=\"_blank\">207<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 213\" title=\"Lopes, R. M. C. et al. Cryovolcanism on titan: new results from cassini RADAR and VIMS. J. Geophys. Res. Planets 118, 416&#x2013;435 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR213\" id=\"ref-link-section-d198960051e3148\" rel=\"nofollow noopener\" target=\"_blank\">213<\/a>.<\/p>\n<p>Crucially, Titan\u2019s atmosphere provides an important analog for understanding prebiotic synthesis under reducing atmospheric conditions, both on Titan and on early Earth. Photochemical reactions involving CH4 and N2 produce nitriles and other complex organics, while the formation of an organic haze exerts significant climatic effects \u2013 phenomena often invoked in models of Earth\u2019s prebiotic environment following large impacts or other transient reducing episodes<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 76\" title=\"Zahnle, K. J., Lupu, R., Catling, D. C. &amp; Wogan, N. Creation and evolution of impact-generated reduced atmospheres of early Earth. Planet. Sci. J. 1, 11 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR76\" id=\"ref-link-section-d198960051e3159\" rel=\"nofollow noopener\" target=\"_blank\">76<\/a>. These atmospheric processes establish a continuous source of chemically reactive organics to Titan\u2019s surface and interior, potentially feeding aqueous environments with prebiotic precursors. Titan\u2019s liquid water layer(s) could therefore be potentially habitable, supporting aqueous geochemistry, redox reactions, mineral dissolution, and organic processing<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 216\" title=\"McKay, C. P. Titan as the abode of life. Life 6, 8 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR216\" id=\"ref-link-section-d198960051e3163\" rel=\"nofollow noopener\" target=\"_blank\">216<\/a>. However, major uncertainties remain regarding the ocean\u2019s redox balance, energy availability, and inventory of bio-essential elements. Despite these uncertainties, Titan stands out as a complex, organic-rich ocean world. Its subsurface water-ammonia liquid layer(s), coupled to an active surface and thick organic atmosphere, makes Titan a compelling target for exploring habitability beyond Earth<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 217\" title=\"Lorenz, R. D. et al. Dragonfly: A rotorcraft lander concept for scientific exploration at Titan. Johns. Hopkins APL Tech. Dig. 34, 14 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR217\" id=\"ref-link-section-d198960051e3167\" rel=\"nofollow noopener\" target=\"_blank\">217<\/a>.<\/p>\n<p>In summary, subsurface oceans on dwarf planets or icy moons fulfill several basic criteria of planetary habitability: active and persistent liquid water, available bio-essential elements and energy, suitable physicochemical conditions, and prolonged environmental stability<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 2\" title=\"National Academies of Sciences and Medicine, E. Origins, Worlds, and Life: A Decadal Strategy for Planetary Science and Astrobiology 2023&#x2013;2032. (The National Academies Press, Washington, 2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR2\" id=\"ref-link-section-d198960051e3174\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>. However, these environments are inherently more limited than the complex planetary systems of early Earth or Mars, although possibly conducive to some origin of life processes<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 218\" title=\"Sasselov, D. D., Grotzinger, J. P. &amp; Sutherland, J. D. The origin of life as a planetary phenomenon. Sci. Adv. 6, eaax3419 (2020).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR218\" id=\"ref-link-section-d198960051e3178\" rel=\"nofollow noopener\" target=\"_blank\">218<\/a>. While a diverse array of organic molecules has been detected, key biomolecules like amino acids and nucleotides remain elusive. Furthermore, prebiotic pathways that rely on environmental cycles\u2014such as wet-dry cycles for polymerization\u2014are difficult to envision in these perpetually submerged, ice-covered settings (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig4\" rel=\"nofollow noopener\" target=\"_blank\">4<\/a>). Nevertheless, if life could have originated in these environments, it is most likely chemotrophic due to the lack of sunlight<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 65\" title=\"Chyba, C. F. &amp; Hand, K. P. Life without photosynthesis. Science 292, 2026&#x2013;2027 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR65\" id=\"ref-link-section-d198960051e3185\" rel=\"nofollow noopener\" target=\"_blank\">65<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 66\" title=\"Taubner, R.-S. et al. Biological methane production under putative Enceladus-like conditions. Nat. Commun. 9, 748 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR66\" id=\"ref-link-section-d198960051e3188\" rel=\"nofollow noopener\" target=\"_blank\">66<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 219\" title=\"Weber, J. M. et al. A review on hypothesized metabolic pathways on Europa and Enceladus: space-flight detection considerations. Life 13, 1726 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR219\" id=\"ref-link-section-d198960051e3191\" rel=\"nofollow noopener\" target=\"_blank\">219<\/a>. Speculative theoretical estimates of potential biomass, though highly model-dependent, suggest plausible populations for ecosystems of methanogens on Enceladus<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 65\" title=\"Chyba, C. F. &amp; Hand, K. P. Life without photosynthesis. Science 292, 2026&#x2013;2027 (2001).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR65\" id=\"ref-link-section-d198960051e3195\" rel=\"nofollow noopener\" target=\"_blank\">65<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 66\" title=\"Taubner, R.-S. et al. Biological methane production under putative Enceladus-like conditions. Nat. Commun. 9, 748 (2018).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR66\" id=\"ref-link-section-d198960051e3198\" rel=\"nofollow noopener\" target=\"_blank\">66<\/a> or SRB on Europa<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 67\" title=\"Hand, K. P., Carlson, R. W. &amp; Chyba, C. F. Energy, chemical disequilibrium, and geological constraints on Europa. Astrobiology 7, 1006&#x2013;1022 (2007).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR67\" id=\"ref-link-section-d198960051e3203\" rel=\"nofollow noopener\" target=\"_blank\">67<\/a>. However, these estimated biomasses are many orders of magnitude lower than those in Earth\u2019s oceans, which may limit the evolutionary rate of potential ecosystems per se on such bodies<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 64\" title=\"Crockford, P. W., Bar On, Y. M., Ward, L. M., Milo, R. &amp; Halevy, I. The geologic history of primary productivity. Curr. Biol. 33, 4741&#x2013;4750.e5 (2023).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR64\" id=\"ref-link-section-d198960051e3207\" rel=\"nofollow noopener\" target=\"_blank\">64<\/a>. This profound difference, coupled with the formidable technical challenge of probing through kilometers of ice, defines the primary obstacle for future missions: the detection of a potentially simple and sparse biosphere in a vast, hidden ocean.<\/p>\n<p>Moving forward<\/p>\n<p>Across ocean-bearing worlds, several cross-cutting factors emerge as universal controls on habitability. First, fluid-rock interactions act as a geochemical engine, regulating pH, alkalinity, and redox state while mobilizing nutrients and generating chemical energy. Second, redox gradients and chemical disequilibria provide sustained energy sources for metabolism, whether through radiolytic oxidants on Europa or potential hydrothermally-derived reductants on Enceladus (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig4\" rel=\"nofollow noopener\" target=\"_blank\">4<\/a>). Third, the persistence and stability of liquid water set the temporal window for prebiotic chemistry and evolution; Mars likely experienced episodic clement intervals and short-lived lakes, whereas Europa and Enceladus may maintain long-lived subsurface oceans sustained by tidal and radiogenic heating; Titan\u2019s water-ammonia liquid layer(s) demonstrates persistence at cryogenic temperatures (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Fourth, the availability and speciation of bioessential elements, including carbon, nitrogen, phosphorus, and sulfur, govern feasible metabolisms and energy yields. Finally, environmental extremes impose physiological limits. On Mars, brines often drove aw below terrestrial thresholds for biological activities; on Europa, higher pressure shapes fluid properties and mineral equilibria; on Titan, ammonia lower the freezing point and modulates buffering capacity. Temperature is a universal constraint: on Earth, the upper thermal limit for life approaches ~120 \u00b0C in high-pressure environments where water remains liquid.<\/p>\n<p>Taken together, these controls suggest a multidimensional framework for habitability\u2013integrating the persistence of liquid water, energy from redox disequilibria, nutrient supply and speciation, and the envelope of physicochemical limits (including aw, pressure, and temperature). Factors specific to each planetary body then modulate this framework: Mars\u2019 episodicity and periodically low aw, Ceres\u2019 relict brines shaped by serpentinization, Europa\u2019s oxidant delivery through its ice shell, Enceladus\u2019 hydrothermal flux and phosphate abundance, and Titan\u2019s ammonia-water chemistry and atmospherically-supplied organics. Viewed through this lens, the following unknowns and aspirations naturally resolve into questions about (i) how redox gradients are generated, maintained, and transported; (ii) how nutrients are partitioned and speciated in time and space; (iii) how complex organics form, cycle, and persist; and (iv) how microenvironments at mineral interfaces, within ice, and in porous crust, link chemical conditions to physiological limits.<\/p>\n<p>While the habitability of the ancient Earth and extraterrestrial oceans is often assessed through the lens of modern terrestrial biogeochemistry<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 220\" title=\"Falkowski, P. G., Fenchel, T. &amp; Delong, E. F. The microbial engines that drive Earth&#x2019;s biogeochemical cycles. Science 320, 1034&#x2013;1039 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR220\" id=\"ref-link-section-d198960051e3249\" rel=\"nofollow noopener\" target=\"_blank\">220<\/a>, significant knowledge gaps persist. Key questions remain regarding the generation and maintenance of redox gradients, the speciation and bioavailability of essential nutrients and trace metals, and the production and cycling of complex organic matter. Furthermore, our understanding of chemical microenvironments\u2014within biofilms, at mineral interfaces, and in concealed habitats like deep aquifers and subglacial lakes\u2014lags far behind that of bulk solutions. A fundamental challenge is to quantitatively link chemical conditions to the energetic and physiological limits of life, determining how spatial and temporal variability dictates microbial survival and growth (Table\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"table anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Tab1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Finally, in an era of unprecedented anthropogenic change, the impact of human-driven perturbations on these fundamental aquatic processes and the microbial communities they support represents a critical and poorly constrained frontier of research.<\/p>\n<p>Within our solar system, only on Earth has life persisted for some four billion years and transformed the planet<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 221\" title=\"Knoll, A. H. et al. Earth and Life: A Four Billion Year Conversation. (Princeton Univ. Press, Princeton, 2026).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR221\" id=\"ref-link-section-d198960051e3259\" rel=\"nofollow noopener\" target=\"_blank\">221<\/a>. The modern Earth has been profoundly shaped by the long-term evolution of its geosphere and biosphere<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 220\" title=\"Falkowski, P. G., Fenchel, T. &amp; Delong, E. F. The microbial engines that drive Earth&#x2019;s biogeochemical cycles. Science 320, 1034&#x2013;1039 (2008).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR220\" id=\"ref-link-section-d198960051e3263\" rel=\"nofollow noopener\" target=\"_blank\">220<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 222\" title=\"Knoll, A. H. et al. A Brief History of Earth: Four Billion Years in Eight Chapters. (HarperCollins, New York, 2021).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR222\" id=\"ref-link-section-d198960051e3266\" rel=\"nofollow noopener\" target=\"_blank\">222<\/a> (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>); however, major gaps persist in our understanding of this intertwined history. Although a surface hydrosphere was established early, the evolutionary trajectories of bio-essential elements, including key biomolecules and potentially limiting nutrients like phosphorus and trace metals, remain poorly constrained. Very likely, the origin of life and the environmental settings essential for its emergence differed significantly from the habitable conditions of modern organisms<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 1\" title=\"Cockell, C. S. et al. Habitability: a review. Astrobiology 16, 89&#x2013;117 (2016).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR1\" id=\"ref-link-section-d198960051e3273\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>, representing another fundamental frontier. Furthermore, while processes such as photochemistry, impacts, volcanic activities, and glaciation events are known to have heavily perturbed surface conditions, their specific effects on aquatic chemistry are not well characterized. In spite of the seemingly dynamic interplay between the long-term evolution of the biosphere and geosphere (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig2\" rel=\"nofollow noopener\" target=\"_blank\">2<\/a>), how and to what extent they actually co-evolved also requires deeper investigation. Compounding these challenges, the geological record is heavily biased towards marine sediments, leaving the evolution and role of non-marine aquatic environments throughout Earth\u2019s history poorly constrained.<\/p>\n<p>Beyond the Earth, ocean-bearing worlds, from early Mars to dwarf planets and icy moons, represent prime targets in the search for habitable environments and extraterrestrial life. However, Mars and possibly Venus are informative examples of bodies that have likely long passed their habitable windows (Fig.\u00a0<a data-track=\"click\" data-track-label=\"link\" data-track-action=\"figure anchor\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#Fig1\" rel=\"nofollow noopener\" target=\"_blank\">1<\/a>). Yet, fundamental questions persist regarding the evolution and longevity of their aqueous environments throughout the solar system, as well as the availability of critical bio-elements. These uncertainties are even greater for exoplanetary oceans, hindering a full assessment of their potential for prebiotic chemistry and their possibility for life. Moreover, a paramount challenge remains the detection and robust interpretation of potential biosignatures.<\/p>\n<p>Fortunately, a new generation of missions is poised to transform our understanding. ESA\u2019s Jupiter Icy Moons Explorer (JUICE) mission, launched in 2023, will arrive at Jupiter in 2031 and is designed to conduct detailed observations of three large ocean-bearing moons: Europa, Ganymede, and Callisto<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 223\" title=\"Grasset, O. et al. JUpiter ICy moons Explorer (JUICE): An ESA mission to orbit Ganymede and to characterise the Jupiter system. Planet. Space Sci. 78, 1&#x2013;21 (2013).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR223\" id=\"ref-link-section-d198960051e3293\" rel=\"nofollow noopener\" target=\"_blank\">223<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 224\" title=\"Van Hoolst, T. et al. Geophysical characterization of the interiors of Ganymede, Callisto and Europa by ESA&#x2019;s JUpiter ICy moons explorer. Space Sci. Rev. 220, 54 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR224\" id=\"ref-link-section-d198960051e3296\" rel=\"nofollow noopener\" target=\"_blank\">224<\/a>. Meanwhile, NASA\u2019s Europa Clipper mission, which is also en route to Jupiter and is expected to arrive in 2030, aims to search for water plumes on Europa and, if encountered, characterize material derived from its subsurface ocean through plume fly-throughs<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Waite, J. H. et al. MASPEX-Europa: the Europa Clipper neutral gas mass spectrometer investigation. Space Sci. Rev. 220, 30 (2024).\" href=\"#ref-CR225\" id=\"ref-link-section-d198960051e3303\">225<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" title=\"Kempf, S. et al. SUDA: A SUrface Dust Analyser for compositional mapping of the Galilean moon Europa. Space Sci. Rev. 221, 10 (2025).\" href=\"#ref-CR226\" id=\"ref-link-section-d198960051e3303_1\">226<\/a>,<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 227\" title=\"Pappalardo, R. T. et al. Science overview of the Europa clipper mission. Space Sci. Rev. 220, 40 (2024).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR227\" id=\"ref-link-section-d198960051e3306\" rel=\"nofollow noopener\" target=\"_blank\">227<\/a>. Moreover, NASA and ESA initiated the Mars Sample Return (MSR) program; accordingly, the Perseverance rover has already collected surface samples for potential future return. In parallel, China is implementing the Tianwen-III mission, planned for launch in 2028, which is designed to return drilled Martian samples potentially by 2031<a data-track=\"click\" data-track-action=\"reference anchor\" data-track-label=\"link\" data-test=\"citation-ref\" aria-label=\"Reference 228\" title=\"Hou, Z. et al. In search of signs of life on Mars with China&#x2019;s sample return mission Tianwen-3. Nat. Astron. 9, 783&#x2013;792 (2025).\" href=\"http:\/\/www.nature.com\/articles\/s41467-026-74955-4#ref-CR228\" id=\"ref-link-section-d198960051e3314\" rel=\"nofollow noopener\" target=\"_blank\">228<\/a>. These Martian samples will provide unprecedented insight into the history of water and the potential for life on Mars. Together, these endeavors will not only constrain the habitability of other worlds but also offer a critical lens through which to re-examine the poorly preserved record of life\u2019s origins and early history on Earth.<\/p>\n","protected":false},"excerpt":{"rendered":"Persistent liquid water is generally considered a prerequisite for planetary habitability1,2. For all known life, which is to&hellip;\n","protected":false},"author":2,"featured_media":527682,"comment_status":"","ping_status":"","sticky":false,"template":"","format":"standard","meta":{"footnotes":""},"categories":[23],"tags":[13633,34598,7692,2026,61,60,4397,2027,40016,82,247],"class_list":["post-527681","post","type-post","status-publish","format-standard","has-post-thumbnail","category-space","tag-astrobiology","tag-element-cycles","tag-geochemistry","tag-humanities-and-social-sciences","tag-ie","tag-ireland","tag-marine-chemistry","tag-multidisciplinary","tag-rings-and-moons","tag-science","tag-space"],"_links":{"self":[{"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/posts\/527681","targetHints":{"allow":["GET"]}}],"collection":[{"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/posts"}],"about":[{"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/types\/post"}],"author":[{"embeddable":true,"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/users\/2"}],"replies":[{"embeddable":true,"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/comments?post=527681"}],"version-history":[{"count":0,"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/posts\/527681\/revisions"}],"wp:featuredmedia":[{"embeddable":true,"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/media\/527682"}],"wp:attachment":[{"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/media?parent=527681"}],"wp:term":[{"taxonomy":"category","embeddable":true,"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/categories?post=527681"},{"taxonomy":"post_tag","embeddable":true,"href":"https:\/\/www.newsbeep.com\/ie\/wp-json\/wp\/v2\/tags?post=527681"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}