The alkaline hydrothermal vent (AHV) hypothesis remains the most thermodynamically coherent framework linking Hadean geochemistry to extant biochemistry, yet the 2022–2025 literature has substantially revised it. Rather than a "naked mineral pore," the emerging model is a hybrid organic–inorganic compartment in which medium-chain fatty alcohols integrate into Fe(Ni)S/silicate micropores, enabling protocell boundaries within a chemiosmotic scaffold (Holler et al., 2023; Jordan et al., 2019). In parallel, the most rigorous phylogenomic reconstruction to date (Moody et al., 2024) places the Last Universal Common Ancestor (LUCA) at ~4.2 Ga with a ~2.75 Mb genome encoding ~2,600 proteins, a mature translation apparatus, a V/A-type rotary ATPase, a near-complete Wood–Ljungdahl carbon-fixation pathway, and a partial CRISPR–Cas interference system — implying LUCA was already embedded in a functional microbial ecosystem with now-extinct sister lineages. These findings force a reframing of the debate: the alkaline vent/metabolism-first camp (Martin, Russell, Lane, Moran) and the subaerial/genetics-first camp (Deamer, Damer, Sutherland, Mulkidjanian, Szostak) no longer represent mutually exclusive scenarios. Where consensus ends, this review flags the speculative residues explicitly.
Black smokers — hot (up to 400 °C), acidic (pH 2–3), metal-rich plumes at fast-spreading ridges — were once the canonical cradle of life but have been progressively abandoned because their temperatures hydrolyse organic polymers on millisecond timescales and their acidity precludes H₂-driven CO₂ reduction. The Lost City Hydrothermal Field, discovered in 2000 on the Atlantis Massif (Kelley et al., 2001, Nature), reframed the discussion. Lost City vents are driven not by magma but by serpentinization, the exergonic hydration of olivine and pyroxene:
(Mg,Fe)₂SiO₄ + H₂O → Mg₃Si₂O₅(OH)₄ (serpentine) + Mg(OH)₂ (brucite) + H₂ + heat
Serpentinization produces fluids at 40–90 °C, pH 9–11, saturated in H₂ (up to ~15 mM) and CH₄, emerging through porous Ca/Mg-carbonate–brucite chimneys up to 60 m tall, some persisting for >100,000 years (Martin & Russell, 2007; Kelley et al., 2005). Hadean oceans are modeled as slightly acidic (pH 5–6) and saturated in dissolved CO₂, because of the massive CO₂-rich atmosphere and absence of continental carbonate buffering (Russell, Hall, & Martin, 2010). The encounter between alkaline vent fluid and CO₂-rich ocean at the chimney wall therefore generates a proton-concentration differential of ~3–5 pH units across thin (initially ~1 µm) Fe(Ni)S–greenrust precipitates — a natural analog of the proton-motive force (PMF) that powers all modern cells (Lane & Martin, 2012, Cell, 151, 1406–1416).
Every extant organism stores usable energy in a transmembrane proton (or sodium) gradient of ~100–200 mV (ΔpH ≈ 1–2 units, ΔΨ ≈ 60–180 mV), which an F- or A/V-type rotary ATPase converts into ATP at a cost of ~70 kJ mol⁻¹ (Nicholls & Ferguson, 2013). The isomorphism between this architecture and the Lost City wall is the central claim of the Martin–Russell–Lane model: geochemistry did not merely supply ingredients; it supplied the thermodynamic topology of life itself (Martin & Russell, 2003, Philos. Trans. R. Soc. B, 358, 59–85; Sojo, Herschy, Whicher, Camprubí, & Lane, 2016, Astrobiology, 16, 181–197).
The physical bridge is Fe–S chemistry. Mackinawite (FeS), greigite (Fe₃S₄), violarite, awaruite (Ni₃Fe) and fougerite (green rust, [Fe²⁺Fe³⁺(OH)₂]) all host [4Fe–4S] and [Ni–Fe–S] cluster motifs whose redox potentials (−250 to −500 mV vs. SHE) overlap those of modern ferredoxins and of carbon-monoxide dehydrogenase/acetyl-CoA synthase (CODH/ACS) in methanogens and acetogens (Russell, 2023, Front. Microbiol., 14, 1145915). The empirical capstone came from Hudson, Furubayashi, Dias, Wang, McGlynn, Sojo, Lane, and colleagues (2020, PNAS, 117, 22873–22879), who used a microfluidic H₂/CO₂ reactor with freshly precipitated Fe(Ni)S barriers to demonstrate that a pH gradient alone drives reduction of CO₂ to formate, with isotopic labeling showing electron (not H₂) translocation across the mineral wall. Preiner et al. (2020, Nature Ecology & Evolution, 4, 534–542) extended this to yields of ~200 mM formate, ~100 µM acetate, ~10 µM pyruvate, plus methanol and methane over awaruite/magnetite/greigite at 100 °C — a geochemical Wood–Ljungdahl equivalent. Altair, Sojo, McGlynn, and Varela (2025, JACS, 147, 28674) scaled this to macroscopic reactors and showed that currents of 10 nA–10 µA across [Ni]FeS minerals suffice to drive both WLP steps (CO₂ → formic acid and C–C coupling to acetic acid). Weingart, Chen, Helmbrecht, Orsi, Braun, and Alim (2023, Science Advances, 9, eadg8931) visualized in real time, using a quasi-2D microfluidic cell, how finger-morphology precipitates maintain stable microscale pH gradients and accumulate dispersed particles — the compartmentalization function theoreticians had predicted.
A long-standing weakness of the AHV model was the mismatch between a thick (~1 µm) mineral wall and a 5-nm lipid bilayer — the geometry on which rotary ATPases evolved (Gogarten & Deamer, 2016, Nature Microbiology, 1, 16229). Three recent results converge on a resolution.
Holler, Bartlett, Löffler, Cartwright, and Hanczyc (2023, PNAS, 120, e2300491120) showed that chemical gardens — the classic silicate/metal-cation inorganic analogs of vent chimneys — do not form vesicles alone, nor does decanol alone, but chemical gardens grown in the presence of decanol integrate the fatty alcohol and template vesicle formation. The result is a hybrid organic–inorganic boundary in which silicate micropore walls are lined by, and eventually bleb off as, fatty-alcohol lamellae. Jordan, Rammu, Zheludev, Hartley, Maréchal, and Lane (2019, Nature Ecology & Evolution, 3, 1705–1714) had earlier demonstrated that mixed fatty acid/fatty alcohol/glycerol-monoester amphiphiles self-assemble into vesicles at 70 °C and alkaline pH, directly countering earlier objections that seawater ionic strength precludes lipid vesicles in vent settings. Barge's JPL group has systematically extended this through chemical-garden experiments with Fe/Mg silicates as analogs for vent and ocean-world chimneys (Barge et al., 2019, PNAS, 116, 4828–4833; Hooks et al., 2020, Langmuir, showing amino-acid incorporation; Marlin et al., 2024, Chem; Chavez et al., 2024, ACS Earth & Space Chemistry). Helmbrecht, Weingart, Klein, Braun, and Orsi (2023, Geobiology, 21, 78–93) further showed that amakinite → fougerite chimneys grown under anoxic ferruginous conditions concentrate RNA ~1,000-fold, addressing the dilution problem for informational polymers.
A caveat: Purvis, Šiller, Telling, and colleagues (2024, Communications Earth & Environment) initially reported abiotic synthesis of long-chain (C₁₈) fatty acids from H₂/bicarbonate/magnetite at 90 °C — a widely publicized result. The paper was retracted in August 2025 (Communications Earth & Environment, 6, 642) because GC-MS features identified by automated peak-fitting as fatty acids did not, on manual reinspection, match fatty-acid fragmentation patterns. The retraction removes a key claim but does not affect the vesicle-assembly results from exogenous amphiphiles.
Overall, the hybrid organic–inorganic reframing replaces the "mineral-only protocell" with a geochemically bounded compartment that can host, and eventually enclose and inherit, lipid membranes — converting the 1-µm wall into a scaffold for the 5-nm membrane rather than a competitor.
Of the six known biological CO₂-fixation routes, the reductive acetyl-CoA (Wood–Ljungdahl) pathway is the only one whose net reaction is exergonic under anaerobic conditions, the only one that is linear rather than cyclic, and the only one present in both archaea (methanogens) and bacteria (acetogens) with strictly homologous cofactors (Fuchs, 2011, Annu. Rev. Microbiol., 65, 631–658; Sousa et al., 2013). Its architecture maps onto the mineral inventory of serpentinizing vents:
Under vent-plausible conditions (70–100 °C, pH 9–10, 10–100 bar H₂), the standard free energies are favorable:
The first CO₂ reduction step is kinetically inhibited at standard conditions and only proceeds spontaneously under high H₂ partial pressure, alkaline pH, and transition-metal catalysis — exactly the vent conditions (Hudson et al., 2020; Preiner et al., 2020). The overall cellular biomass-synthesis reaction from CO₂ + H₂ remains exergonic across 25–125 °C (Amend & McCollom, 2009, ACS Symp. Ser., 1025, 63–94), meaning that anabolism itself could serve as an energy-harvesting reaction in an autotrophic acetogen, with no external reductant beyond geochemical H₂.
The Strasbourg group (Moran, Muchowska, Tüysüz) and the UCL group (Lane) have mapped non-enzymatic equivalents of substantial portions of core anabolism:
Nunes Palmeira, Colnaghi, Harrison, Pomiankowski, and Lane (2022, Proc. R. Soc. B, 289, 20221469) model a protocell in which branch-point fluxes in protometabolism confer heritable phenotypic variation without genes, provided that nucleotide cofactors feed back to catalyze CO₂ fixation itself. If autocatalysis rests on nucleotide synthesis alone, the system collapses; if it rests on nucleotide-catalyzed carbon fixation, protocell growth rate becomes a selectable trait. Harrison, Rammu, Liu, Halpern, Nunes Palmeira, and Lane (2023, Annu. Rev. Ecol. Evol. Syst., 54, 327–350) generalize this to "life as a guide to its own origins": the topology of autotrophic biosynthesis prefigures the universal core of metabolism, and protocell selection on growth can operate before any polymer-based heredity.
Harrison, Nunes Palmeira, Halpern, and Lane (2022, BBA-Bioenergetics, 1863, 148597) and Halpern, Bartsch, Ibrahim, Harrison, Ahn, Christodoulou, and Lane (2023, Life, 13, 1129) reinvigorated the stereochemical hypothesis for code origin. Mapping the MetaCyc autotrophic-biosynthesis network onto codon assignments, Harrison et al. showed the first codon letter tracks distance from CO₂ fixation (G < A < C < U), the anticodon middle base correlates with hydrophobicity, and third-position redundancy correlates with amino-acid size. Halpern et al. then used molecular dynamics and NMR for all 20 amino acids × 4 ribonucleotides × 3 charge states, finding that 95% of amino acids interact most strongly with at least one codonic or anticodonic base, and preference for the cognate anticodonic middle base exceeds 99% of randomized assignments. The interactions are weak (ΔΔG ~1.5 kcal mol⁻¹) but statistically real: random RNA sequences would template non-random peptides, seeding heritable information without a full translation apparatus.
The Szostak and Joyce programs have materially advanced the competing genetics-first hypothesis in 2022–2025. Papastavrou, Horning, and Joyce (2024, PNAS, 121, e2321592121) reported an RNA polymerase ribozyme propagating a hammerhead ribozyme across multiple generations with heritable variation — the first genuinely Darwinian RNA evolution — but copy fidelity remains ~90%, below the ~97% Eigen threshold needed for the replicase to replicate itself. Szostak's group has demonstrated in-situ activation of non-enzymatic RNA copying (Ding, Zhang, & Szostak, 2023, Nucleic Acids Res., 51, 6528–6539; Aitken, Wright, Radakovic, & Szostak, 2023, JACS, 145, 16142–16149), ribozyme-catalyzed RNA ligation in prebiotically plausible protocells (DasGupta, Zhang, Smela, & Szostak, 2023, Chem. Eur. J., 29, e202301376), rain-induced coacervate stabilization (Agrawal et al., 2024, Science Advances, 10, eadn9657), and selective non-enzymatic formation of biological RNA hairpins (Wu et al., 2025, Angew. Chem., 64, e202417370).
Classical objections to the RNA-first model persist: RNA's prebiotic synthesis requires narrow chemistries, its hydrolytic instability in Mg²⁺-rich conditions constrains ribozyme function, and its catalytic range is limited compared to protein. But the 2022–2025 convergence is striking: metabolism-first workers now show non-enzymatic nucleotide and ATP-proxy synthesis, while genetics-first workers embed ribozymes in fatty-acid protocells. The honest contemporary synthesis is that metabolism, membranes, and RNA coevolved in a single autotrophic protocell, with protometabolism supplying the chemical inventory that RNA later coded for, and weak biophysical amino acid–nucleotide affinities seeding the statistical templating of functional peptides (Harrison et al., 2023).
The landmark paper of the decade is Moody, Álvarez-Carretero, Mahendrarajah, Clark, Betts, Dombrowski, Szánthó, Boyle, Daines, Chen, Lane, Yang, Shields, Szöllősi, Spang, Pisani, Williams, Lenton, and Donoghue (2024, Nature Ecology & Evolution, 8, 1654–1666). Its principal findings:
Moody et al.'s age inference rests on cross-bracing of pre-LUCA paralogues — five pairs (F- vs. A/V-type ATPase catalytic and non-catalytic subunits, EF-Tu vs. EF-G, SRP vs. SRP-receptor, tyrosyl- vs. tryptophanyl-tRNA synthetases, leucyl- vs. valyl-tRNA synthetases) whose pre-LUCA duplication means LUCA appears twice on each gene tree. Constraining the mirrored nodes to the same age effectively doubles calibrations and dramatically narrows posterior intervals (developed in Mahendrarajah et al., 2023, Nature Communications, 14, 7456). Gene content inference relies on Amalgamated Likelihood Estimation (ALE), a probabilistic algorithm that reconciles gene-tree bootstrap distributions against a reference species tree (built from 57 ancient vertically evolving markers across 350 bacterial and 350 archaeal genomes; Moody et al., 2022, eLife, 11, e66695; Coleman et al., 2021, Science, 372, eabe0511), jointly modelling duplication, transfer, and loss (DTL). This directly addresses the central methodological critique of earlier LUCA inference.
Weiss, Sousa, Mrnjavac, Neukirchen, Roettger, Nelson-Sathi, and Martin (2016, Nature Microbiology, 1, 16116) clustered 6.1 million prokaryotic proteins into 286,514 families and applied nested filters requiring presence in both Archaea and Bacteria plus two-domain phyletic monophyly, yielding 355 LGT-resistant genes (3% of two-domain clusters) — the 97% rejection rate is itself evidence of pervasive ancient LGT. The 355-gene dataset depicted LUCA as anaerobic, thermophilic, H₂/CO₂-using via the Wood–Ljungdahl pathway, N₂-fixing, and Fe–S rich — compatible with a serpentinizing-vent habitat.
Three critiques followed:
Moody et al. resolve these by modelling LGT rather than filtering it out, propagating uncertainty across >9,000 KO families. Remarkably, the qualitative physiological picture converges with Weiss et al.'s: an anaerobic, H₂/CO₂-using Wood–Ljungdahl autotroph — but Moody's LUCA is seven-fold richer in gene content and no longer includes nitrogenase (consistent with Pi et al., 2022, Mol. Biol. Evol., placing N₂-fixation at the Last Bacterial Common Ancestor, not LUCA).
Moody et al.'s high-confidence reconstruction renders LUCA as follows:
Perhaps the most paradigm-shifting claim is that LUCA did not live alone. Three lines converge:
Multiple independent methods now agree that LUCA was an anaerobic, H₂/CO₂-using acetogen with Wood–Ljungdahl metabolism, a mature translation apparatus, rotary ATPase chemiosmosis, Fe–S biochemistry, and a cellular (not progenote) organization, and that LGT was rampant at and around LUCA. The ~2.5–3.0 Mb, ~2,600-gene estimate remains a Moody-2024-specific result whose LOESS extrapolation from modern genomes should not be treated as settled. The partial CRISPR–Cas claim is singular and invites replication. The 4.2 Ga point estimate is contingent on rejection of any LHB maximum constraint; reinstatement would push LUCA younger by 300–400 Myr. Membrane lipid chemistry, precise root topology (CPR/DPANN placement), and the single-cell-vs-population status of LUCA remain open.
AHV proponents must confront the fundamental thermodynamic asymmetry: proteins, nucleic acids, and oligosaccharides are condensation polymers whose backbones are formed by elimination of water, so by Le Chatelier's principle bulk water drives hydrolysis. Quantitatively:
Submerged vent chimneys continuously replenish fluid and cannot naturally reach low water activity in the bulk, which is what Damer and Deamer (2020) and Gogarten and Deamer (2016) identify as a fatal weakness of the AHV model for polymer synthesis. Ross (2018, Astrobiology) further argues that a pH gradient across a ~1 µm mineral wall cannot do meaningful thermodynamic work on polymerization without a chemiosmotic coupling machinery to concentrate the free energy into a specific bond.
AHV advocates reply on three fronts: (i) serpentinizing micropores may host structurally bound water of low activity analogous to dried films (Nascimento Vieira, Sousa, & Martin, 2020, FEBS Letters); (ii) CO₂ + H₂ → biomass is overall exergonic under alkaline conditions across 25–125 °C (Amend & McCollom, 2009); and (iii) acetyl phosphate (Pinna et al., 2022) supplies the ~45 kJ mol⁻¹ of transferable phosphorylation potential needed to drive condensation coupled to an energy-rich intermediate. Whether these solutions close the gap remains contested.
Damer and Deamer (2020, Astrobiology, 20, 429–452) offer the canonical modern synthesis. Fluctuating volcanic hot-spring pools (analog sites: Bumpass Hell, Mutnovsky, Yellowstone, Rotorua, Pilbara Dresser Formation) cycle through dry (condensation), moist (gel-phase sharing), and wet (vesicle budding) states. Monomers concentrate between multilamellar lipid bilayers during drying (reduced water activity drives condensation); products redistribute among protocells during moist phases; vesicles encapsulating polymer sets bud off during wet phases. Selection acts on progenote aggregates via shared combinatorial chemistry.
Key arguments:
Empirical wet–dry cycle demonstrations (2020–2025) have become increasingly robust:
The most productive prebiotic syntheses of the 2010s and 2020s — the Sutherland and Powner programs — are intrinsically surface chemistries requiring UV-B/C flux unavailable to submarine vents.
The most recent speculative extension is Mulkidjanian, Dibrova, and Bychkov (2025, Life, 15, 399), which proposes that the Moon-forming impact ejected moderately volatile elements (particularly Zn and K) onto Earth's protocrust, where geothermal vapor leached them into K⁺/Zn²⁺-rich pools. Metallic Zn and ⁴⁰K radiolysis would then drive CO₂/N₂ reduction; ZnS photocatalytic edifices would both shield and activate nascent RNA under high UV flux (quantum yield ~90% for H₂ production from aqueous ZnS). The cosmochemical estimate of ~10¹⁹ kg of metallic Zn is highly speculative and independent tests are lacking, but the model's appeal is that it simultaneously explains the K⁺/Zn²⁺ signature of cytoplasm and supplies an electron source without H₂.
Bruce Damer's programmatic extension of the Deamer framework treats progenote aggregates as the unit of selection — aggregated protocells sharing polymer products across permeable boundaries, with combinatorial selection acting on encapsulated polymer sets (Damer & Deamer, 2020). This framework has received partial experimental support (Hassenkam et al., 2020; Song et al., 2024) but the explicit "progenote combinatorial selection" component remains conceptual rather than fully demonstrated.
A convergence is emerging. Barge and Price (2022, Nature Geoscience) argued that shallow (<200 m) alkaline hydrothermal vents — exemplified by the Prony field in New Caledonia and the Strýtan field in Iceland — offer wet–dry cycling, tidal fluctuation, salinity variation, and UV exposure alongside serpentinization-driven H₂/alkaline chemistry. This hybrid setting combines the chemiosmotic logic of the AHV model with the polymerization conditions of the Damer–Deamer model.
The water problem is a real thermodynamic obstacle (~+3 kcal mol⁻¹ per bond) not seriously contested in principle. Cyanosulfidic chemistry reproducibly produces ribonucleotides, amino acids, and lipid precursors from plausible feedstocks. Wet–dry cycling reproducibly produces RNA-like polymers up to ~50 nt. Hadean subaerial volcanic land existed. These constitute the robust subaerial pillars. What remains speculative is whether those polymers are sequence-specific enough to bootstrap ribozymes, whether Mulkidjanian's post-impact Zn-metal layer existed at the proposed scale, whether AHV chemistry can yield polymers at all (yields remain sub-µM), and whether Hadean surface UV-B flux was high enough in practice (modelled but atmospheric-composition-dependent).
The 2020–2025 literature has quietly dissolved the traditional polarization between the alkaline vent and subaerial models, and between metabolism-first and genetics-first. Three observations drive the reframing.
First, LUCA is now reconstructed as a genomically modern, ecologically embedded prokaryote at ~4.2 Ga, not a progenote (Moody et al., 2024). This pushes the origin of cellular life far earlier than previously assumed and implies a substantial pre-LUCA phase of co-evolving lineages — the "ecosystem LUCA." The origin of life, in other words, was followed by hundreds of millions of years of microbial community evolution before the node ancestral to all extant cells crystallized.
Second, the alkaline vent model has been rescued from the naked-mineral-pore problem by the demonstration that fatty alcohols integrate into chemical-garden walls (Holler et al., 2023) and that mixed amphiphiles self-assemble into vesicles in vent-like conditions (Jordan et al., 2019), while pH-gradient-driven CO₂ reduction across Fe(Ni)S barriers is experimentally demonstrated at both microfluidic and macroscopic scales (Hudson et al., 2020; Preiner et al., 2020; Altair et al., 2025). The model now describes a hybrid organic–inorganic protocell within a chemiosmotic scaffold, not a speculative mineral-only compartment.
Third, non-enzymatic analogs of carbon fixation, the reverse Krebs cycle, amino-acid biosynthesis, pyrimidine nucleobase synthesis, ribonucleotide phosphorylation, and acetyl-phosphate-mediated ATP synthesis now span most of core anabolism (Moran, Muchowska, Tüysüz, Lane groups, 2022–2024). The "metabolic heredity" framework (Nunes Palmeira et al., 2022; Harrison et al., 2023) formalizes selection on growth rate before polymer-based genes, while weak but statistically real biophysical amino acid–nucleotide affinities (Halpern et al., 2023) show how random RNA would template non-random peptides, bridging metabolism and information without requiring a fully formed translation apparatus.
The unresolved speculative residues remain substantial. The water problem has not been decisively solved within the vent framework; the most successful RNA and peptide syntheses remain subaerial-compatible (Sutherland, Powner, Deamer). The CRISPR–Cas signal at LUCA, the ~2.6-kgene size, and the 4.2 Ga age all depend on modelling choices (LOESS extrapolation, rejection of an LHB constraint) whose robustness will be tested in the next generation of reconciliations. Membrane lipid chemistry of LUCA is still not resolved — the archaea-bacteria lipid divide remains the deepest unanswered question in deep phylogenomics.
The most defensible current model is a shallow alkaline hydrothermal setting with tidal and evaporative wet–dry cycling and occasional UV exposure (Barge & Price, 2022) — a geochemical niche that combines the chemiosmotic logic of Russell and Martin, the polymerization engine of Deamer and Damer, the photochemistry of Sutherland, and the ionic composition of Mulkidjanian. In this setting, geochemistry supplied the energy topology, the catalysts, and the first metabolic intermediates; hybrid organic-inorganic compartments supplied the boundaries; wet–dry cycles and nitrile-chemistry supplied the polymerization thermodynamics; and weak biophysical affinities between amino acids and nucleotides supplied the statistical template for coded information. LUCA emerged from this milieu not as a lonely founder but as the sole surviving lineage of a richer Hadean biosphere — a reminder that the origin of life is a problem in community ecology as much as in chemistry.
Agrawal, A., Radakovic, A., Vonteddu, A., Rizvi, S., & Szostak, J. W. (2024). Did the exposure of coacervate droplets to rain make them the first stable protocells? Science Advances, 10(15), eadn9657.
Aitken, H. R. M., Wright, T. H., Radakovic, A., & Szostak, J. W. (2023). Small-molecule organocatalysis of in-situ nucleotide activation and RNA copying. Journal of the American Chemical Society, 145(30), 16142–16149.
Altair, T., Sojo, V., McGlynn, S. E., & Varela, A. S. (2025). Electrochemical reduction of CO₂ to formic and acetic acid over synthetic [Ni]-FeS minerals: Implications for prebiotic Wood–Ljungdahl chemistry. Journal of the American Chemical Society, 147, 28674.
Amend, J. P., & McCollom, T. M. (2009). Energetics of biomolecule synthesis on early Earth. ACS Symposium Series, 1025, 63–94.
Amend, J. P., & Shock, E. L. (2001). Energetics of overall metabolic reactions of thermophilic and hyperthermophilic Archaea and Bacteria. FEMS Microbiology Reviews, 25(2), 175–243.
Andam, C. P., Williams, D., & Gogarten, J. P. (2015). Biased gene transfer mimics patterns created through shared ancestry. BMC Evolutionary Biology, 15, 70.
Barge, L. M., Flores, E., Baum, M. M., VanderVelde, D. G., & Russell, M. J. (2019). Redox and pH gradients drive amino acid synthesis in iron oxyhydroxide mineral systems. PNAS, 116(11), 4828–4833.
Barge, L. M., & Price, R. E. (2022). Diverse geochemical conditions for prebiotic chemistry in shallow-sea alkaline hydrothermal vents. Nature Geoscience, 15, 976–981.
Becker, S., Feldmann, J., Wiedemann, S., Okamura, H., Schneider, C., Iwan, K., Crisp, A., Rossa, M., Amatov, T., & Carell, T. (2019). Unified prebiotically plausible synthesis of pyrimidine and purine RNA ribonucleotides. Science, 366(6461), 76–82.
Benner, S. A., Kim, H.-J., & Carrigan, M. A. (2012). Asphalt, water, and the prebiotic synthesis of ribose, ribonucleosides, and RNA. Accounts of Chemical Research, 45(12), 2025–2034.
Berkemer, S. J., & McGlynn, S. E. (2020). A new analysis of archaea–bacteria domain separation: Variable phylogenetic distance and the tempo of early evolution. Molecular Biology and Evolution, 37(8), 2332–2340.
Betts, H. C., Puttick, M. N., Clark, J. W., Williams, T. A., Donoghue, P. C. J., & Pisani, D. (2018). Integrated genomic and fossil evidence illuminates life's early evolution and eukaryote origin. Nature Ecology & Evolution, 2(10), 1556–1562.
Beyazay, T., Belthle, K. S., Farès, C., Preiner, M., Moran, J., Martin, W. F., & Tüysüz, H. (2023). Ambient temperature CO₂ fixation to pyruvate and subsequently to citramalate over iron and nickel nanoparticles. Nature Communications, 14, 570.
Boehnke, P., & Harrison, T. M. (2016). Illusory late heavy bombardments. PNAS, 113(39), 10802–10806.
Coleman, G. A., Davín, A. A., Mahendrarajah, T. A., Szánthó, L. L., Spang, A., Hugenholtz, P., Szöllősi, G. J., & Williams, T. A. (2021). A rooted phylogeny resolves early bacterial evolution. Science, 372(6542), eabe0511.
Crapitto, A. J., Campbell, A., Harris, A. J., & Goldman, A. D. (2022). A consensus view of the proteome of the last universal common ancestor. Ecology and Evolution, 12(6), e8930.
Damer, B., & Deamer, D. (2020). The hot spring hypothesis for an origin of life. Astrobiology, 20(4), 429–452.
DasGupta, S., Zhang, S., Smela, M. P., & Szostak, J. W. (2023). RNA-catalysed RNA ligation within prebiotically plausible model protocells. Chemistry — A European Journal, 29, e202301376.
Dass, A. V., Wunnava, S., Langlais, J., von der Esch, B., Krusche, M., Ufer, L., Chrisam, N., Dubini, R. C. A., Rovó, P., Ochsenfeld, C., Braun, D., Schwierz, N., & Mutschler, H. (2023). RNA oligomerisation without added catalyst from 2′,3′-cyclic nucleotides by drying at air–water interfaces. ChemSystemsChem, 5(1), e202200026.
Delaye, L. (2024). A new view of the last universal common ancestor. Journal of Molecular Evolution, 92, 567–571.
Dherbassy, Q., Mayer, R. J., & Moran, J. (2024). Coenzymes in a pre-enzymatic metabolism. Science Advances, 10(38), eadr5357.
Dherbassy, Q., Mayer, R. J., Muchowska, K. B., & Moran, J. (2023). Metal-pyridoxal cooperativity in nonenzymatic transamination. Journal of the American Chemical Society, 145(24), 13357–13370.
Ding, D., Zhang, L., & Szostak, J. W. (2023). Enhanced nonenzymatic RNA copying with in-situ activation of short oligonucleotides. Nucleic Acids Research, 51, 6528–6539.
Djokic, T., Van Kranendonk, M. J., Campbell, K. A., Walter, M. R., & Ward, C. R. (2017). Earliest signs of life on land preserved in ca. 3.5 Ga hot spring deposits. Nature Communications, 8, 15263.
Fairchild, J., Islam, S., Singh, J., Bučar, D.-K., & Powner, M. W. (2024). Prebiotically plausible chemoselective pantetheine synthesis in water. Science, 383(6686), 911–918.
Foden, C. S., Islam, S., Fernández-García, C., Maugeri, L., Sheppard, T. D., & Powner, M. W. (2020). Prebiotic synthesis of cysteine peptides that catalyze peptide ligation in neutral water. Science, 370(6518), 865–869.
Fuchs, G. (2011). Alternative pathways of carbon dioxide fixation: Insights into the early evolution of life? Annual Review of Microbiology, 65, 631–658.
Gogarten, J. P., & Deamer, D. (2016). Is LUCA a thermophilic progenote? Nature Microbiology, 1, 16229.
Goldman, A. D., & Kaçar, B. (2023). Very early evolution from the perspective of microbial ecology. Journal of Molecular Evolution, 91, 683–692.
Green, N. J., Xu, J., & Sutherland, J. D. (2021). Illuminating life's origins: UV photochemistry in abiotic synthesis of biomolecules. Journal of the American Chemical Society, 143(19), 7219–7236.
Halpern, A., Bartsch, L. R., Ibrahim, K., Harrison, S. A., Ahn, M., Christodoulou, J., & Lane, N. (2023). Biophysical interactions underpin the emergence of information in the genetic code. Life, 13(5), 1129.
Harrison, S. A., Nunes Palmeira, R., Halpern, A., & Lane, N. (2022). A biophysical basis for the emergence of the genetic code in protocells. Biochimica et Biophysica Acta — Bioenergetics, 1863(8), 148597.
Harrison, S. A., Rammu, H., Liu, F., Halpern, A., Nunes Palmeira, R., & Lane, N. (2023). Life as a guide to its own origins. Annual Review of Ecology, Evolution, and Systematics, 54, 327–350.
Hassenkam, T., Damer, B., Mednick, G., & Deamer, D. (2020). AFM images of viroid-sized rings that self-assemble from mononucleotides through wet–dry cycling. Life, 10(12), 321.
Hassenkam, T., & Deamer, D. (2022). Visualizing RNA polymers produced by hot wet-dry cycling. Scientific Reports, 12, 10098.
Helmbrecht, V., Weingart, M., Klein, F., Braun, D., & Orsi, W. D. (2023). White and green rust chimneys accumulate RNA in a ferruginous chemical garden. Geobiology, 21, 78–93.
Herschy, B., Whicher, A., Camprubí, E., Watson, C., Dartnell, L., Ward, J., Evans, J. R. G., & Lane, N. (2014). An origin-of-life reactor to simulate alkaline hydrothermal vents. Journal of Molecular Evolution, 79, 213–227.
Holler, S., Bartlett, S., Löffler, R. J. G., Cartwright, J. H. E., & Hanczyc, M. M. (2023). Hybrid organic–inorganic structures trigger the formation of primitive cell-like compartments. PNAS, 120(37), e2300491120.
Hudson, R., de Graaf, R., Strandoo Rodin, M., Ohno, A., Lane, N., McGlynn, S. E., Yamada, Y. M. A., Nakamura, R., Barge, L. M., Braun, D., & Sojo, V. (2020). CO₂ reduction driven by a pH gradient. PNAS, 117(37), 22873–22879.
Jordan, S. F., Rammu, H., Zheludev, I. N., Hartley, A. M., Maréchal, A., & Lane, N. (2019). Promotion of protocell self-assembly from mixed amphiphiles at the origin of life. Nature Ecology & Evolution, 3, 1705–1714.
Kaur, H., Rauscher, S. A., Werner, E., Song, Y., Yi, J., Kazöne, W., Martin, W. F., Tüysüz, H., & Moran, J. (2024). A prebiotic Krebs cycle analog generates amino acids with H₂ and NH₃ over nickel. Chem, 10(5), 1528–1540.
Kelley, D. S., Karson, J. A., Blackman, D. K., Früh-Green, G. L., Butterfield, D. A., Lilley, M. D., Olson, E. J., Schrenk, M. O., Roe, K. K., Lebon, G. T., & Rivizzigno, P. (2001). An off-axis hydrothermal vent field near the Mid-Atlantic Ridge at 30°N. Nature, 412, 145–149.
Kelley, D. S., et al. (2005). A serpentinite-hosted ecosystem: The Lost City hydrothermal field. Science, 307, 1428–1434.
Lane, N. (2015). The vital question: Energy, evolution, and the origins of complex life. W. W. Norton.
Lane, N., & Martin, W. F. (2012). The origin of membrane bioenergetics. Cell, 151(7), 1406–1416.
Mahendrarajah, T. A., Moody, E. R. R., Schrempf, D., Szánthó, L. L., Dombrowski, N., Davín, A. A., Pisani, D., Donoghue, P. C. J., Szöllősi, G. J., Williams, T. A., & Spang, A. (2023). ATP synthase evolution on a cross-braced dated tree of life. Nature Communications, 14, 7456.
Martin, W. F. (2020). Older than genes: The acetyl-CoA pathway and origins. Frontiers in Microbiology, 11, 817.
Martin, W., & Russell, M. J. (2003). On the origins of cells. Philosophical Transactions of the Royal Society B, 358, 59–85.
Martin, W., & Russell, M. J. (2007). On the origin of biochemistry at an alkaline hydrothermal vent. Philosophical Transactions of the Royal Society B, 362, 1887–1925.
Milshteyn, D., Damer, B., Havig, J., & Deamer, D. (2018). Amphiphilic compounds assemble into membranous vesicles in hydrothermal hot spring water but not in seawater. Life, 8, 11.
Moody, E. R. R., Álvarez-Carretero, S., Mahendrarajah, T. A., Clark, J. W., Betts, H. C., Dombrowski, N., Szánthó, L. L., Boyle, R. A., Daines, S., Chen, X., Lane, N., Yang, Z., Shields, G. A., Szöllősi, G. J., Spang, A., Pisani, D., Williams, T. A., Lenton, T. M., & Donoghue, P. C. J. (2024). The nature of the last universal common ancestor and its impact on the early Earth system. Nature Ecology & Evolution, 8(9), 1654–1666.
Moody, E. R. R., Mahendrarajah, T. A., Dombrowski, N., Clark, J. W., Petitjean, C., Offre, P., Szöllősi, G. J., Spang, A., & Williams, T. A. (2022). An estimate of the deepest branches of the tree of life from ancient vertically evolving genes. eLife, 11, e66695.
Muchowska, K. B., Varma, S. J., & Moran, J. (2019). Synthesis and breakdown of universal metabolic precursors promoted by iron. Nature, 569, 104–107.
Muchowska, K. B., Varma, S. J., & Moran, J. (2020). Nonenzymatic metabolic reactions and life's origins. Chemical Reviews, 120(15), 7708–7744.
Mulkidjanian, A. Y., Bychkov, A. Y., Dibrova, D. V., Galperin, M. Y., & Koonin, E. V. (2012). Origin of first cells at terrestrial, anoxic geothermal fields. PNAS, 109(14), E821–E830.
Mulkidjanian, A. Y., Dibrova, D. V., & Bychkov, A. Y. (2025). Origin of the RNA World in cold Hadean geothermal fields enriched in zinc and potassium: Abiogenesis as a positive fallout from the Moon-forming impact? Life, 15(3), 399.
Nascimento Vieira, A., Kleinermanns, K., Martin, W. F., & Preiner, M. (2020). The ambivalent role of water at the origins of life. FEBS Letters, 594(17), 2717–2733.
Nunes Palmeira, R., Colnaghi, M., Harrison, S. A., Pomiankowski, A., & Lane, N. (2022). The limits of metabolic heredity in protocells. Proceedings of the Royal Society B, 289, 20221469.
Papastavrou, N., Horning, D. P., & Joyce, G. F. (2024). RNA-catalyzed evolution of catalytic RNA. PNAS, 121(11), e2321592121.
Patel, B. H., Percivalle, C., Ritson, D. J., Duffy, C. D., & Sutherland, J. D. (2015). Common origins of RNA, protein and lipid precursors in a cyanosulfidic protometabolism. Nature Chemistry, 7(4), 301–307.
Pi, H.-W., et al. (2022). Origin and evolution of nitrogen fixation in prokaryotes. Molecular Biology and Evolution, 39(9), msac181.
Pinna, S., Kunz, C., Halpern, A., Harrison, S. A., Jordan, S. F., Ward, J., Werner, F., & Lane, N. (2022). A prebiotic basis for ATP as the universal energy currency. PLoS Biology, 20(10), e3001437.
Powner, M. W., Gerland, B., & Sutherland, J. D. (2009). Synthesis of activated pyrimidine ribonucleotides in prebiotically plausible conditions. Nature, 459, 239–242.
Preiner, M., Igarashi, K., Muchowska, K. B., Yu, C., Varma, S. J., Kleinermanns, K., Nobu, M. K., Kamagata, Y., Tüysüz, H., Moran, J., & Martin, W. F. (2020). A hydrogen-dependent geochemical analogue of primordial carbon and energy metabolism. Nature Ecology & Evolution, 4, 534–542.
Ragsdale, S. W., & Pierce, E. (2008). Acetogenesis and the Wood–Ljungdahl pathway of CO₂ fixation. Biochimica et Biophysica Acta, 1784, 1873–1898.
Ross, D. S., & Deamer, D. (2016). Dry/wet cycling and the thermodynamics and kinetics of prebiotic polymer synthesis. Life, 6(3), 28.
Russell, M. J. (2023). A self-sustaining serpentinization mega-engine feeds the fougerite nanoengines implicated in the emergence of guided metabolism. Frontiers in Microbiology, 14, 1145915.
Russell, M. J., Hall, A. J., & Martin, W. (2010). Serpentinization as a source of energy at the origin of life. Geobiology, 8, 355–371.
Schuchmann, K., & Müller, V. (2014). Autotrophy at the thermodynamic limit of life: A model for energy conservation in acetogenic bacteria. Nature Reviews Microbiology, 12, 809–821.
Sojo, V., Herschy, B., Whicher, A., Camprubí, E., & Lane, N. (2016). The origin of life in alkaline hydrothermal vents. Astrobiology, 16(2), 181–197.
Song, Y., Hassenkam, T., Deamer, D., & Zare, R. N. (2024). Wet–dry cycles cause nucleic acid monomers to polymerize into long chains. PNAS, 121(49), e2412784121.
Sousa, F. L., Thiergart, T., Landan, G., Nelson-Sathi, S., Pereira, I. A. C., Allen, J. F., Lane, N., & Martin, W. F. (2013). Early bioenergetic evolution. Philosophical Transactions of the Royal Society B, 368, 20130088.
Spang, A., Mahendrarajah, T. A., Offre, P., & Stairs, C. W. (2022). Evolving perspective on the origin and diversification of cellular life and the virosphere. Genome Biology and Evolution, 14(6), evac034.
Szöllősi, G. J., Tannier, E., Lartillot, N., & Daubin, V. (2013). Lateral gene transfer from the dead. Systematic Biology, 62, 386–397.
Thauer, R. K., Jungermann, K., & Decker, K. (1977). Energy conservation in chemotrophic anaerobic bacteria. Bacteriological Reviews, 41, 100–180.
Thoma, B., & Powner, M. W. (2023). Selective synthesis of lysine peptides and the prebiotically plausible synthesis of catalytically active diaminopropionic acid peptide nitriles in water. Journal of the American Chemical Society, 145(5), 3121–3130.
Varma, S. J., Muchowska, K. B., Chatelain, P., & Moran, J. (2018). Native iron reduces CO₂ to intermediates and end-products of the acetyl-CoA pathway. Nature Ecology & Evolution, 2, 1019–1024.
Weingart, M., Chen, W., Helmbrecht, V., Orsi, W. D., Braun, D., & Alim, K. (2023). Visualization of mineral chimney formation and associated prebiotic pH gradients. Science Advances, 9, eadg8931.
Weiss, M. C., Preiner, M., Xavier, J. C., Zimorski, V., & Martin, W. F. (2018). The last universal common ancestor between ancient Earth chemistry and the onset of genetics. PLOS Genetics, 14(8), e1007518.
Weiss, M. C., Sousa, F. L., Mrnjavac, N., Neukirchen, S., Roettger, M., Nelson-Sathi, S., & Martin, W. F. (2016). The physiology and habitat of the last universal common ancestor. Nature Microbiology, 1, 16116.
Werner, E., Pinna, S., Mayer, R. J., & Moran, J. (2023). Metal/ADP complexes promote phosphorylation of ribonucleotides. Journal of the American Chemical Society, 145(39), 21630–21637.
Wolfe, J. M., & Fournier, G. P. (2018). Horizontal gene transfer constrains the timing of methanogen evolution. Nature Ecology & Evolution, 2, 897–903.
Xu, J., Tsanakopoulou, M., Magnani, C. J., Szabla, R., Šponer, J. E., Šponer, J., Góra, R. W., & Sutherland, J. D. (2022). Azoles as auxiliaries and intermediates in prebiotic nucleoside synthesis. Journal of the American Chemical Society, 144(42), 19447–19455.
Yi, J., Kaur, H., Kazöne, W., Rauscher, S. A., Gravillier, L.-A., Muchowska, K. B., & Moran, J. (2022). A nonenzymatic analog of pyrimidine nucleobase biosynthesis. Angewandte Chemie International Edition, 61, e202117211.
Zimmermann, J., Werner, E., Sodei, S., & Moran, J. (2024). Pinpointing conditions for a metabolic origin of life: Underlying mechanisms and the role of coenzymes. Accounts of Chemical Research, 57(20), 3032–3043.