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Reactive oxygen species, especially hydrogen peroxide, trigger rapid downward migration in diatom biofilms even in darkness and without needing the usual light-driven photoprotective cycle.

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2026-07-11 22:44 UTC pith:32HOOX46

load-bearing objection Solid experimental isolation of exogenous H2O2 as a darkness-sufficient trigger of reversible downward migration in a natural diatom biofilm, with a clean separation from xanthophyll photoprotection; the F0 proxy is the softest step, not a collapse of the claim. the 3 major comments →

arxiv 2607.03958 v1 pith:32HOOX46 submitted 2026-07-04 physics.bio-ph

Reactive oxygen species trigger downward vertical migration in diatom microphytobenthic biofilms as a strategy to cope with oxidative stress

Alexandre Desparmet , Bruno Jesus , Tony Robinet , Thierry Dufour , C\'edric Hubas This is my paper
classification physics.bio-ph
keywords diatom microphytobenthic biofilmsvertical migrationreactive oxygen specieshydrogen peroxidexanthophyll cyclephotophysiologyPleurosigma strigosumcold atmospheric plasma
verification ladder T0 review T1 audit T2 compute T3 formal T4 reserved

The pith

A machine-rendered reading of the paper's core claim, the machinery that carries it, and where it could break.

Intertidal diatom biofilms live under daily light swings that generate reactive oxygen species capable of damaging photosynthesis. Two well-known defenses are vertical migration into sediment and the xanthophyll cycle that dissipates excess energy. This paper asks whether ROS themselves are the signal that starts migration, and whether that signal is separate from the photoprotective pathway. Natural biofilms, dominated by the motile diatom Pleurosigma strigosum, were exposed either with sediment (migration free) or without it (migration blocked) to high light, hydrogen peroxide, or cold-atmospheric-plasma oxidative stress. Downward migration occurred under every oxidative treatment; the strongest, near-on/off response was produced by exogenous H2O2 in the dark, with little drop in photosynthetic efficiency and without a controlled non-photochemical-quenching response. High light alone engaged the xanthophyll cycle more strongly when cells could not migrate. The work therefore positions ROS sensing as an early-warning trigger that lets benthic diatoms leave harmful surface conditions before damage accumulates, and that can act independently of, or in synergy with, classical photoprotection.

Core claim

ROS, particularly H2O2, act as a sufficient trigger for rapid, reversible downward vertical migration in diatom-dominated microphytobenthic biofilms. Exogenous H2O2 applied in darkness elicits a strong motility response without requiring light or an effective Y(NPQ)/xanthophyll photoprotective response, indicating that the migration pathway can operate independently of the light-driven photoprotective cascade.

What carries the argument

ROS-triggered oxidotactic vertical migration: a rapid actin-myosin gliding response, scored by surface F0 biomass decline and recovery, that is elicited by both endogenous (high-light) and exogenous (H2O2, plasma) oxidative stress and that can be uncoupled from xanthophyll-cycle photoprotection.

Load-bearing premise

The large, reversible drop in surface fluorescence under 200 micromolar hydrogen peroxide is taken as a true ROS-signaling motility switch rather than non-specific toxicity or surface-cell damage, even though that concentration exceeds typical tidal-zone levels and intracellular calcium or motility-machinery readouts were not measured.

What would settle it

Apply the same 200 micromolar H2O2 treatment while blocking aquaporin-mediated H2O2 entry or calcium-channel activity; if the rapid F0 drop and subsequent recovery are abolished (or if intracellular ROS/Ca2+ sensors fail to rise before migration), the claimed ROS-signaling pathway fails.

Watch this falsifier — get emailed when new claim-graph text bears on it.

Editorial analysis

A structured set of objections, weighed in public.

Desk editor's note, referee report, simulated authors' rebuttal, and a circularity audit.

Referee Report

3 major / 5 minor

Summary. The manuscript tests whether reactive oxygen species (ROS), especially H2O2, act as a trigger for downward vertical migration in natural diatom-dominated microphytobenthic biofilms, and whether that motility can be engaged independently of xanthophyll-cycle photoprotection. Using paired sediment-associated (migration-permissive) and sediment-free (migration-restricted) communities, the authors apply high light, cold atmospheric plasma (composite RONS), and ~200 μM H2O2 matched to plasma-derived H2O2, against dark-adapted controls. Surface biomass is tracked by F0, photophysiology by PAM fluorescence, and metabolic impact by HPLC pigments, with community composition constrained by 18S amplicon sequencing (dominated by Pleurosigma strigosum). All three oxidative treatments induce reversible F0 declines interpreted as downward migration, strongest under exogenous H2O2; high light uniquely drives controlled de-epoxidation and Y(NPQ), amplified when migration is blocked, whereas exogenous ROS oxidize pigments more broadly with limited photosynthetic impairment and without effective Y(NPQ).

Significance. If the F0 kinetics truly report ROS-triggered gliding rather than non-specific surface loss, the work supplies a clear experimental bridge between environmental/metabolic oxidative stress and a major behavioral trait of epipelic diatoms, and it usefully separates motility from classical photoprotective engagement. Strengths include a coherent multi-stress design (natural high-light control, plasma composite stress, concentration-matched H2O2 positive control, dark exposures for exogenous ROS), the sediment-associated vs sediment-free contrast that exposes compensatory xanthophyll activation, and full open deposition of scripts, pigment chromatograms, and sequencing data (GitHub/Zenodo/ENA). The result would matter for intertidal biofilm ecology and for ROS as an early-warning signal in aquatic microbiomes, even if the proposed ROS–Ca2+–motility cascade (Fig. 6) remains a hypothesis for follow-up.

major comments (3)
  1. [§3.2, Fig. 2, Table 2] §3.2, Fig. 2, Table 2: The central claim that exogenous H2O2/plasma trigger physiological downward migration rests almost entirely on rapid, reversible surface F0 declines (−76% under 200 μM H2O2; recovery slope +116 F0 min⁻¹). Sediment-free F0 stability and high Fv/Fm usefully exclude wholesale bleaching and karyostrophy, but they do not exclude partial surface-cell detachment, EPS/adhesion failure, or selective loss of the most exposed cells under a supra-environmental dose. Without subsurface biomass profiles, direct motility assays, or microscopy of vertical redistribution, the F0-to-migration inference remains the least secure step. Strengthen this load-bearing link (e.g., post-stress vertical sectioning/F0 depth profiles, adhesion/EPS checks, or explicit dose–response below environmental ranges) or substantially qualify the claim.
  2. [§2.1; §4.2] §2.1 Oxidative treatments; §4.2: The authors themselves note that 200 μM H2O2 exceeds typical tidal-zone levels (citing values up to ~85 μM in precipitation and lower sediment photooxidation). Because the strongest behavioral response and the independence-from-photophysiology argument both come from this dose (and the matched plasma treatment), ecological transferability of the “on/off ROS trigger” needs either a lower-dose series that still elicits migration or a clearer framing that the result is a mechanistic positive control rather than a direct environmental simulation.
  3. [§3.3; §4.3; Fig. 3] §3.3–§4.3 and Fig. 3: Photophysiological parameters in sediment-associated biofilms are acknowledged to be compromised by migration (surface population change, possible residual senescent cells, NPQ underestimation). The claim that exogenous ROS induce migration “without” effective controlled Y(NPQ) therefore depends mainly on sediment-free communities, where migration cannot occur. Make this dependency explicit in the abstract/conclusion wording, and avoid treating sediment-associated NPQ/Y(NPQ) patterns under H2O2/plasma as evidence of pathway independence.
minor comments (5)
  1. [Fig. 6; §4.5] Fig. 6 and §4.5 present a putative ROS–calcium–motility cascade. The text correctly labels it hypothetical, but the figure and some summary sentences read more assertively than the data support. Soften figure title/legend language to “hypothetical model” throughout.
  2. [Table 1; §2.1] Table 1 and Methods: Plasma is a composite RONS stress (H2O2 plus NOx species). When attributing effects specifically to ROS/H2O2, consistently remind the reader that the H2O2 arm is the isolating control and that plasma alone cannot assign causality to H2O2.
  3. [§2.3] §2.3: Clarify timing of post-stress PAM measurements (high-light after dark re-adaptation vs H2O2/plasma immediately after dark exposure) earlier when interpreting ΔAfter–Before comparisons across treatments.
  4. [Summary; throughout] Minor text issues: “photo synthetic” / “underl ying” spacing artifacts in the Summary; ensure consistent units (μmol.photons.m−2.s−1 vs μmol photons m−2 s−1) and that Supplementary Tables 3–5 are fully cross-referenced where F0 and pigment statistics are claimed.
  5. [§3.1] Community characterization: 18S relative abundances are semi-quantitative and include possible planktonic eDNA; a brief caveat when attributing behavior primarily to P. strigosum would help non-specialist readers.

Circularity Check

0 steps flagged

Experimental intervention study with no derivation that reduces outputs to inputs by construction; H2O2 dose matching and standard F0/pigment proxies are design choices, not circular predictions.

full rationale

This paper is an empirical intervention study on natural microphytobenthic biofilms, not a first-principles derivation. The central claim—that exogenous ROS (especially H2O2) trigger rapid downward migration independently of controlled Y(NPQ)/xanthophyll photoprotection—is supported by comparative treatments (high-light, plasma-activated seawater, 200 μM H2O2) versus dark-adapted controls, with F0 as a standard surface-biomass proxy, HPLC pigment ratios (including de-epoxidation state), and PAM fluorescence parameters. The H2O2 concentration is set to match the ~209 μM produced by the 10-min plasma treatment for comparability, not fitted to force the migration outcome. No equation equates a claimed prediction to a fitted parameter; no uniqueness theorem or ansatz is imported via self-citation as a load-bearing premise; and standard proxies (F0, DES%, Fv/Fm, NPQ) are not redefined as the results they measure. Self-citations, if any, are not load-bearing for the trigger claim. Circularity score is therefore 0: the inference chain is experimental and externally falsifiable, even if the biological interpretation of F0 kinetics remains open to non-motility alternatives (a correctness/strength issue, not circularity).

Axiom & Free-Parameter Ledger

3 free parameters · 5 axioms · 1 invented entities

The central claim rests on standard experimental-biology domain assumptions (F0 as surface-biomass proxy; PAM-derived yields; HPLC pigment ratios; ROS as diffusible signals) plus a few design choices (stress doses and exposure times). No new physical entities are postulated as necessary for the result; the ROS–Ca2+–motility scheme in Fig. 6 is an interpretive model built from cited literature, not a required invented particle or force.

free parameters (3)
  • H2O2 exposure concentration
    Set to 200 μM to match measured plasma-activated seawater H2O2 (209 μM); chosen for comparability rather than field realism, and the migration magnitude depends on this dose.
  • High-light irradiance and duration
    1500 μmol photons m−2 s−1 for 30 min is an experimental choice representing intense natural light; response amplitude is dose- and time-dependent.
  • Plasma exposure geometry and duration
    10 min indirect plasma-activated seawater with 3 mm gap; composite RONS mix is instrument- and protocol-specific.
axioms (5)
  • domain assumption Dark-adapted minimum fluorescence F0 is a valid proxy for surface microalgal biomass and thus for vertical migration when bleaching and karyostrophy are ruled out.
    Used throughout §2.4 and Fig. 2/Table 2; supported by sediment-free F0 stability and Fv/Fm checks, but remains an indirect behavioral readout.
  • domain assumption Exogenously applied H2O2 can enter cells (e.g., via aquaporins) and act as a physiological signal at the applied dose.
    Invoked in §4.4 with citation to aquaporin literature; required to interpret H2O2 as a signaling trigger rather than only an external oxidant.
  • domain assumption De-epoxidation state and Y(NPQ) report engagement of the diadinoxanthin–diatoxanthin photoprotective pathway.
    Standard diatom photophysiology assumption used in §2.5 and §3.3 to claim pathway separation from migration.
  • domain assumption Sediment-free biofilm preparation preserves a representative motile surface community dominated by P. strigosum without major preparation artifacts.
    §2.1 community conditioning and §3.1 sequencing; needed to interpret impaired-migration responses as compensatory rather than community-shift artifacts.
  • standard math Statistical comparisons (Welch t-tests, Mann–Whitney, ANOSIM/SIMPER) on the reported n are sufficient to support treatment effects as stated.
    Applied in §2.6 and Results; small n for fluorescence (n=3) limits power but does not redefine the claim.
invented entities (1)
  • Putative ROS–calcium–motility signaling cascade (Fig. 6) no independent evidence
    purpose: Integrates observed H2O2-triggered migration with literature on Ca2+ channels and gliding machinery as a hypothetical mechanism.
    Not required to accept the behavioral result; presented as putative with grey dashed hypothetical links. No new molecule is measured or uniquely required by the data.

pith-pipeline@v1.1.0-grok45 · 25847 in / 3483 out tokens · 33749 ms · 2026-07-11T22:44:30.509302+00:00 · methodology

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read the original abstract

Diatom-dominated intertidal microphytobenthic biofilms experience daily fluctuations in irradiance, which can lead to oxidative stress within the photosynthetic apparatus through the production and accumulation of reactive oxygen species. To maintain photosynthetic efficiency, benthic diatoms have developed protective strategies, including mobilization of the antioxidant xanthophyll cycle and the ability to migrate vertically through sediments. However, mechanistic understanding of signaling pathways underlying migration remains poorly characterized. This study investigated the triggering effect of reactive oxygen species on behavioral and photophysiological responses through the analysis of lipophilic pigments and fluorescence parameters. To this end, two microphytobenthic communities, one with sediment allowing vertical migration and another without sediment restricting it, were exposed to irradiance, cold atmospheric plasma, and hydrogen peroxide stresses. Results showed a consistent downward migration response under all oxidative stresses, highlighting the key role of reactive oxygen species, especially hydrogen peroxide, in triggering this microphytobenthic behavior. Moreover, a difference was observed between the pathways involved in vertical migration and those underlying photoprotective responses. Hydrogen peroxide and cold atmospheric plasma stresses highlighted the necessity for substantial microphytobenthic migration, whereas irradiance induced a specific and controlled response involving engagement of the xanthophyll cycle, acting in synergy with the migration strategy by showing stronger activation when migration was impaired.

Figures

Figures reproduced from arXiv: 2607.03958 by Alexandre Desparmet, Bruno Jesus, C\'edric Hubas, Thierry Dufour, Tony Robinet.

Figure 2
Figure 2. Figure 2: Downward migration of microphytobenthos under the three oxidative stresses. Temporal monitoring of surface microalg [PITH_FULL_IMAGE:figures/full_fig_p006_2.png] view at source ↗

discussion (0)

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Reference graph

Works this paper leans on

97 extracted references · 83 canonical work pages · 1 internal anchor

  1. [5]

    The strongest downward migration occurred under exogenous stresses that minimally impacted photosynthetic efficiency, despite a broad an d untargeted oxidative stress on pigments

    Conclusion All results provide strong evidence for the key role of ROS, particularly H 2O2, as a critical signaling molecule triggering the rapid motile behavior in diatom -dominated microphytobenthic biofilms. The strongest downward migration occurred under exogenous stresses that minimally impacted photosynthetic efficiency, despite a broad an d untarge...

  2. [6]

    We are also thankful to Manon Leroux for her essential assistance in the laboratory during the experiment

    Acknowledgements Our thanks go to Johann Lavaud for his valuable discussions on photophysiology, and Cyril Noël for his assistance with the SAMBA workflow, which enabled the analysis of the metabarcoding data. We are also thankful to Manon Leroux for her essential assistance in the laboratory during the experiment. We acknowledge the chromatography and ma...

  3. [7]

    Author contributions AD (Conceptualization, Methodology, Investigation, Data curation, Formal analysis, Visualization, Writing —original draft), BJ (Data curation, Formal analysis, Validation, Writing —review & editing), TR (Data curation, Methodology, Formal analysis), TD (Reso urces, Methodology, Writing —review & editing), and CH (Supervision, Methodol...

  4. [8]

    Supplementary materials Supplementary material is available at The ISME Journal online

  5. [9]

    Conflicts of interest The authors declare no conflict of interest

  6. [10]

    Funding This work was funded by the Institut de l’Océan from the Sorbonne University alliance and the Muséum national d’Histoire naturelle

  7. [11]

    The datasets generated and analyzed during the current study are available in the Zenodo repository (https://doi.org/10.5281/zenodo.15835853)

    Data availability All scripts used to generate the figures and perform the analyses are available on GitHub ( https://github.com/adesparmet/ROS- Triggered-Diatom-Biofilm-Migration), and are also archived on Zenodo ( https://zenodo.org/records/17813206). The datasets generated and analyzed during the current study are available in the Zenodo repository (ht...

    work page doi:10.5281/zenodo.15835853 2026

  8. [12]

    The light field of microbenthic communities

    Kühl M, Jorgensen BB. The light field of microbenthic communities. Limnol Oceanogr 1994;39:1368–98. https://doi.org/10.4319/lo.1994.39.6.1368

    work page doi:10.4319/lo.1994.39.6.1368 1994

  9. [13]

    Microphytobenthos

    Underwood GJC. Microphytobenthos. Encyclopedia of Ocean Sciences. San Diego, CA, USA, 2001, 1770–7. https://doi.org/10.1006/rwos.2001.0213

    work page doi:10.1006/rwos.2001.0213 2001

  10. [14]

    A mini-review of the contribution of benthic microalgae to the ecology of the continental shelf in the South Atlantic bight

    Pinckney JL. A mini-review of the contribution of benthic microalgae to the ecology of the continental shelf in the South Atlantic bight. Estuaries Coast 2018;41:2070–8. https://doi.org/10.1007/s12237-018-0401-z

    work page doi:10.1007/s12237-018-0401-z 2018

  11. [15]

    Microphytobenthic biofilms

    Hubas C, Passarelli C, Paterson DM. Microphytobenthic biofilms. In (ed.), Mudflat Ecology. Cham, 2018, 63 –90. https://doi.org/10.1007/978-3-319- 99194-8_4

    doi:10.1007/978-3-319- 2018

  12. [16]

    Reactive oxygen species in plant signaling

    Waszczak C, Carmody M, Kangasjarvi J. Reactive oxygen species in plant signaling. Annu Rev Plant Biol 2018;69:209 –36. https://doi.org/10.1146/annurev- arplant-042817-040322

    doi:10.1146/annurev- 2018

  13. [17]

    The Molecular Life of Diatoms

    Falciatore A, Mock T (eds). The Molecular Life of Diatoms. Cham, 2022. https://doi.org/10.1007/978-3-030-92499-7

    work page doi:10.1007/978-3-030-92499-7 2022

  14. [18]

    Duchêne C, Bouly J-P, Pierella Karlusich JJ. et al. Diatom phytochromes integrate the underwater light spectrum to sense depth. Nature 2025;637:691–7. https://doi.org/10.1038/s41586-024-08301-3

    work page doi:10.1038/s41586-024-08301-3 2025

  15. [19]

    Light signaling in photosynthetic eukaryotes with ‘green’ and ‘red’ chloroplasts

    Lepetit B, Dietzel L. Light signaling in photosynthetic eukaryotes with ‘green’ and ‘red’ chloroplasts. Environ Exp Bot 2015;114:30 –47. https://doi.org/10.1016/j.envexpbot.2014.07.007

    work page doi:10.1016/j.envexpbot.2014.07.007 2015

  16. [20]

    ROS production and signalling in chloroplasts

    Foyer CH, Hanke G. ROS production and signalling in chloroplasts. Plant J 2022;111:642–61. https://doi.org/10.1111/tpj.15856

    work page doi:10.1111/tpj.15856 2022

  17. [21]

    Doose C, Oger C, Mas -Normand L. et al. Non -enzymatic oxylipin production in a mudflat microphytobenthic biofilm. Front Photobiol 2024;2:1441713. https://doi.org/10.3389/fphbi.2024.1441713

    work page doi:10.3389/fphbi.2024.1441713 2024

  18. [22]

    A new paradigm for the action of reactive oxygen species in the photoinhibition of photosystem II

    Nishiyama Y, Allakhverdiev SI, Murata N. A new paradigm for the action of reactive oxygen species in the photoinhibition of photosystem II. Biochim Biophys Acta Bioenerg 2006;1757:742 –9. https://doi.org/10.1016/j.bbabio.2006.05.013

    work page doi:10.1016/j.bbabio.2006.05.013 2006

  19. [23]

    Oxylipins and reactive carbonyls as regulators of the plant redox and reactive oxygen species network under stress

    Knieper M, Viehhauser A, Dietz K-J. Oxylipins and reactive carbonyls as regulators of the plant redox and reactive oxygen species network under stress. Antioxidants 2023;12:814. https://doi.org/10.3390/antiox12040814

    work page doi:10.3390/antiox12040814 2023

  20. [24]

    Mittler R, Vanderauwera S, Suzuki N. et al. ROS signaling? Trends Plant Sci 2011;16:300–9. https://doi.org/10.1016/j.tplants.2011.03.007

    work page doi:10.1016/j.tplants.2011.03.007 2011

  21. [25]

    Serôdio J, Ezequiel BA. et al. Efficiency of photoprotection in microphytobenthos. Aquat Microb Ecol 2012;16:161 –75. https://doi.org/10.3354/ame01591

    work page doi:10.3354/ame01591 2012

  22. [26]

    Photoprotection in Intertidal Benthic Diatoms

    Blommaert L. Photoprotection in Intertidal Benthic Diatoms. Ghent, Belgium. Faculty of Sciences, 2017, Ph.D. dissertation

    2017

  23. [27]

    Le rythme des marées chez les diatomées littorales

    Fauvel P, Bohn G. Le rythme des marées chez les diatomées littorales. CR Soc Biol 1907;62:121–3

    1907

  24. [28]

    The ups and downs of life in a benthic biofilm

    Consalvey M, Paterson DM, Underwood GJC. The ups and downs of life in a benthic biofilm. Diatom Res 2004;19:181 –202. https://doi.org/10.1080/0269249X.2004.9705870

    work page doi:10.1080/0269249x.2004.9705870 2004

  25. [29]

    Endogenous versus environmental control of vertical migration by intertidal benthic microalgae

    Coelho H, Vieira S, Serôdio J. Endogenous versus environmental control of vertical migration by intertidal benthic microalgae. Eur J Phycol 2011;46:271–81. https://doi.org/10.1080/09670262.2011.598242

    work page doi:10.1080/09670262.2011.598242 2011

  26. [30]

    Barnett A, Méléder V, Dupuy C. et al. The vertical migratory rhythm of intertidal microphytobenthos in sediment depends on the light photoperiod, intensity, and spectrum. Front Mar Sci 2020;7:212. https://doi.org/10.3389/fmars.2020.00212

    work page doi:10.3389/fmars.2020.00212 2020

  27. [31]

    Diatom Gliding Motility

    Cohn S, Manoylov K, Gordon R (eds). Diatom Gliding Motility. Hoboken, NJ, USA & Sons, 2021. https://doi.org/10.1002/9781119526483

    work page doi:10.1002/9781119526483 2021

  28. [32]

    Morelle J, Bastos A, Frankenbach S. et al. The photoprotective behavior of a motile benthic diatom as elucidated from the interplay between cell motility and physiological responses to a light micro-gradient using a novel experimental setup. Microb Ecol 2024;87:40. https://doi.org/10.1007/s00248-024-02354-7

    work page doi:10.1007/s00248-024-02354-7 2024

  29. [33]

    Poulsen NC, Spector I, Spurck TP. et al. Diatom gliding is the result of an actin-myosin motility system. Cell Motil Cytoskeleton 1999;44:23 –33. https://doi.org/10.1002/(SICI)1097-0169(199909)44:1<23::AID-CM2>3.0.CO;2-D

    work page doi:10.1002/(sici)1097-0169(199909)44:1 1999

  30. [34]

    Perkins R, Lavaud J, Serôdio J. et al. Vertical cell movement is a primary response of intertidal benthic biofilms to increasing light dose. Mar Ecol Prog Ser 2010;416:93–103. https://doi.org/10.3354/meps08787

    work page doi:10.3354/meps08787 2010

  31. [35]

    Travelling expenses

    Marques Da Silva J, Duarte B, Utkin AB. Travelling expenses. Front Mar Sci 2020;7:433. https://doi.org/10.3389/fmars.2020.00433

    work page doi:10.3389/fmars.2020.00433 2020

  32. [36]

    Suzuki S, Ota S, Yamagishi T. et al. Rapid transcriptomic and physiological changes in the freshwater pennate diatom Mayamaea pseudoterrestris in response to copper exposure. DNA Res 2022;29. https://doi.org/10.1093/dnares/dsac037

    work page doi:10.1093/dnares/dsac037 2022

  33. [37]

    Morelle J, Bastos A, Frankenbach S. et al. Inter -specific variability in photophysiological responses of epipelic diatoms to the actin inhibitor Latrunculin B. Diatom Res 2025;40:133 –46. https://doi.org/10.1080/0269249X.2024.2447333

    work page doi:10.1080/0269249x.2024.2447333 2025

  34. [38]

    Accelerated diversification is related to life history and locomotion in a hyperdiverse lineage of microbial eukaryotes (diatoms, Bacillariophyta)

    Nakov T, Beaulieu JM, Alverson AJ. Accelerated diversification is related to life history and locomotion in a hyperdiverse lineage of microbial eukaryotes (diatoms, Bacillariophyta). New Phytol 2018;219:462 –73. https://doi.org/10.1111/nph.15137

    work page doi:10.1111/nph.15137 2018

  35. [39]

    S Roy, CA Llewellyn, ES Egeland. et al. (eds). Phytoplankton Pigments, Chemotaxonomy and Applications in Oceanography. Cambridge, UK, 2011. https://doi.org/10.1017/CBO9780511732263

    work page doi:10.1017/cbo9780511732263 2011

  36. [40]

    Photosynthetic pigments in diatoms

    Kuczynska P, Jemiola -Rzeminska M, Strzalka K. Photosynthetic pigments in diatoms. Mar Drugs 2015;13:5847 –81. https://doi.org/10.3390/md13095847

    work page doi:10.3390/md13095847 2015

  37. [41]

    Regulation and function of xanthophyll cycle - dependent photoprotection in algae

    Goss R, Jakob T. Regulation and function of xanthophyll cycle - dependent photoprotection in algae. Photosynth Res 2010;106:103 –22. https://doi.org/10.1007/s11120-010-9536-x

    work page doi:10.1007/s11120-010-9536-x 2010

  38. [42]

    Xanthophyll cycle — a mechanism protecting plants against oxidative stress

    Latowski D, Kuczyńska P, Strzałka K. Xanthophyll cycle — a mechanism protecting plants against oxidative stress. Redox Rep 2011;16:78 –90. https://doi.org/10.1179/174329211X13020951739938

    work page doi:10.1179/174329211x13020951739938 2011

  39. [43]

    Bastos A, Morelle J, Frankenbach S. et al. Light response of karyostrophy in the benthic pennate diatom Pleurosigma strigosum (Bacillariophyceae)? J Phycol 2025;61:1140–52. https://doi.org/10.1111/jpy.70055

    work page doi:10.1111/jpy.70055 2025

  40. [44]

    Green - and blue-light- mediated chloroplast migration in the centric diatom Pleurosira laevis

    Furukawa T, Watanabe M, Shihira -Ishikawa I. Green - and blue-light- mediated chloroplast migration in the centric diatom Pleurosira laevis. Protoplasma 1998;203:214–20. https://doi.org/10.1007/BF01279479

    work page doi:10.1007/bf01279479 1998

  41. [45]

    An upright life -form of an epipelic motile diatom

    Jönsson B, Sundbäck K, Nilsson C. An upright life -form of an epipelic motile diatom. Eur J Phycol 1994;29:11 –5. https://doi.org/10.1080/09670269400650421

    work page doi:10.1080/09670269400650421 1994

  42. [46]

    Jesus B, Jauffrais T, Trampe E. et al. Microscale imaging sheds light on species-specific strategies for photo -regulation and photo -acclimation of microphytobenthic diatoms. Environ Microbiol 2023;25:3087 –103. https://doi.org/10.1111/1462-2920.16499

    work page doi:10.1111/1462-2920.16499 2023

  43. [47]

    De Tommasi E, Rea I, Ferrara MA. et al. Multiple -pathways light modulation in Pleurosigma strigosum bi -raphid diatom. Sci Rep 2024;14:6476. https://doi.org/10.1038/s41598-024-56206-y

    work page doi:10.1038/s41598-024-56206-y 2024

  44. [48]

    Nymark M, Valle KC, Brembu T. et al. An integrated analysis of molecular acclimation to high light in the marine diatom Phaeodactylum tricornutum. PLoS One 2009;4. https://doi.org/10.1371/journal.pone.0007743

    work page doi:10.1371/journal.pone.0007743 2009

  45. [49]

    Lepetit B, Volke D, Gilbert M. et al. Evidence for the existence of one antenna-associated, lipid -dissolved and two protein -bound pools of diadinoxanthin cycle pigments in diatoms. Plant Physiol 2010;154:1905 –

    2010

  46. [50]

    https://doi.org/10.1104/pp.110.166454

    work page doi:10.1104/pp.110.166454

  47. [51]

    Valle KC, Nymark M, Aamot I. et al. System responses to equal doses of photosynthetically usable radiation of blue, green, and red light in the marine diatom Phaeodactylum tricornutum. PLoS One 2014;9. https://doi.org/10.1371/journal.pone.0114211

    work page doi:10.1371/journal.pone.0114211 2014

  48. [52]

    Walpersdorf E, Kühl M, Elberling B. et al. In situ oxygen dynamics and carbon turnover in an intertidal sediment (Skallingen, Denmark). Mar Ecol Prog Ser 2017;566:49–65. https://doi.org/10.3354/meps12016

    work page doi:10.3354/meps12016 2017

  49. [53]

    Stoeck T, Bass D, Nebel M. et al. Multiple marker parallel tag environmental DNA sequencing reveals a highly complex eukaryotic community in marine anoxic water. Mol Ecol 2010;19:21 –31. https://doi.org/10.1111/j.1365-294X.2009.04480.x

    work page internal anchor Pith review Pith/arXiv arXiv doi:10.1111/j.1365-294x.2009.04480.x 2010

  50. [54]

    Estimating primary production rates from photosynthetic electron transport in estuarine microphytobenthos

    Barranguet KJ. Estimating primary production rates from photosynthetic electron transport in estuarine microphytobenthos. Mar Ecol Prog Ser 2000;204:39–52. https://doi.org/10.3354/meps204039

    work page doi:10.3354/meps204039 2000

  51. [55]

    A model for the relationship between light intensity and the rate of photosynthesis in phytoplankton

    Eilers PHC, Peeters JCH. A model for the relationship between light intensity and the rate of photosynthesis in phytoplankton. Ecol Model 1988;42:199–215. https://doi.org/10.1016/0304-3800(88)90057-9

    work page doi:10.1016/0304-3800(88)90057-9 1988

  52. [56]

    Modeling the irradiance dependency of the quantum efficiency of photosynthesis

    Silsbe GM, Kromkamp JC. Modeling the irradiance dependency of the quantum efficiency of photosynthesis. Limnol Oceanogr Methods 2012;10:645–52. https://doi.org/10.4319/lom.2012.10.645

    work page doi:10.4319/lom.2012.10.645 2012

  53. [57]

    Consalvey M, Perkins RG, Paterson DM. et al. PAM fluorescence. Diatom Res 2005;20:1–22. https://doi.org/10.1080/0269249X.2005.9705619

    work page doi:10.1080/0269249x.2005.9705619 2005

  54. [59]

    The use of HPLC pigment analysis to study microphytobenthos communities

    Brotas V, Plante-Cuny M-R. The use of HPLC pigment analysis to study microphytobenthos communities. Acta Oecol 2003;24 –15. https://doi.org/10.1016/S1146-609X(03)00013-4

    work page doi:10.1016/s1146-609x(03)00013-4 2003

  55. [60]

    Biases in bulk

    Van Der Loos LM, Nijland R. Biases in bulk. Mol Ecol 2021;30:3270–88. https://doi.org/10.1111/mec.15592

    work page doi:10.1111/mec.15592 2021

  56. [61]

    An untargeted metabolomic approach for microphytobenthic biofilms in intertidal mudflats

    Gaubert-Boussarie J, Prado S, Hubas C. An untargeted metabolomic approach for microphytobenthic biofilms in intertidal mudflats. Front Mar Sci 2020;7:250. https://doi.org/10.3389/fmars.2020.00250

    work page doi:10.3389/fmars.2020.00250 2020

  57. [62]

    What determines species composition in microphytobenthic biofilms

    Underwood GJC, Barnett M. What determines species composition in microphytobenthic biofilms. Royal Netherlands Academy of Arts and Sciences 2006;103:123–40

    2006

  58. [63]

    Méléder V, Rincé Y, Barillé L. et al. Spatiotemporal changes in microphytobenthos assemblages in a macrotidal flat (Bourgneuf Bay, France). J Phycol 2007;43:1177–90. https://doi.org/10.1111/j.1529-8817.2007.00423.x

    work page doi:10.1111/j.1529-8817.2007.00423.x 2007

  59. [64]

    The brown clock

    Farré EM. The brown clock. Physiol Plant 2020;169:430 –41. https://doi.org/10.1111/ppl.13104

    work page doi:10.1111/ppl.13104 2020

  60. [65]

    Prins A, Deleris P, Hubas C. et al. Effect of light intensity and light quality on diatom behavioral and physiological photoprotection. Front Mar Sci 2020;7:203. https://doi.org/10.3389/fmars.2020.00203

    work page doi:10.3389/fmars.2020.00203 2020

  61. [66]

    Schneider RJ, Roe KL, Hansel CM. et al. Species-level variability in extracellular production rates of reactive oxygen species by diatoms. Front Chem 2016;4:5. https://doi.org/10.3389/fchem.2016.00005

    work page doi:10.3389/fchem.2016.00005 2016

  62. [67]

    Richter PR, Streb C, Ntefidou M. et al. High light-induced sign change of gravitaxis in the flagellate euglena gracilis is mediated by reactive oxygen species. Acta Protozool 2003;42:197–204

    2003

  63. [68]

    Dynamics of UV-driven hydrogen peroxide formation on an intertidal sand flat

    Abele-Oeschger D, Tüg H, Röttgers R. Dynamics of UV-driven hydrogen peroxide formation on an intertidal sand flat. Limnol Oceanogr 1997;42:1406–15. https://doi.org/10.4319/lo.1997.42.6.1406

    work page doi:10.4319/lo.1997.42.6.1406 1997

  64. [69]

    Kottuparambil S, Shin W, Brown MT. et al. UV -B affects photosynthesis, ROS production and motility of the freshwater flagellate, Euglena agilis Carter. Aquat Toxicol 2012;122 –123:206–13. https://doi.org/10.1016/j.aquatox.2012.06.002

    work page doi:10.1016/j.aquatox.2012.06.002 2012

  65. [70]

    Reactive oxygen species in the world ocean and their impacts on marine ecosystems

    Morris JJ, Rose AL, Lu Z. Reactive oxygen species in the world ocean and their impacts on marine ecosystems. Redox Biol 2022;52:102285. https://doi.org/10.1016/j.redox.2022.102285

    work page doi:10.1016/j.redox.2022.102285 2022

  66. [71]

    Mizrachi A, Graff Van Creveld S, Shapiro OH. et al. Light -dependent single-cell heterogeneity in the chloroplast redox state regulates cell fate in a marine diatom. eLife 2019;8. https://doi.org/10.7554/eLife.47732

    work page doi:10.7554/elife.47732 2019

  67. [72]

    Lavaud J, Rousseau B, Van Gorkom HJ. et al. Influence of the diadinoxanthin pool size on photoprotection in the marine planktonic diatom Phaeodactylum tricornutum. Plant Physiol 2002;129:1398 –406. https://doi.org/10.1104/pp.002014

    work page doi:10.1104/pp.002014 2002

  68. [73]

    Waring J, Klenell M, Bechtold U. et al. Light -induced responses of oxygen photoreduction, reactive oxygen species production and scavenging in two diatom species. J Phycol 2010;46:1206 –17. https://doi.org/10.1111/j.1529-8817.2010.00919.x

    work page doi:10.1111/j.1529-8817.2010.00919.x 2010

  69. [74]

    UV- and salinity-induced oxidative effects in the marine diatom Cylindrotheca closterium during simulated emersion

    Rijstenbil JW. UV- and salinity-induced oxidative effects in the marine diatom Cylindrotheca closterium during simulated emersion. Mar Biol 2005;147:1063–73. https://doi.org/10.1007/s00227-005-0015-4

    work page doi:10.1007/s00227-005-0015-4 2005

  70. [75]

    Kalaji HM, Jajoo A, Oukarroum A. et al. Chlorophyll a fluorescence as a tool to monitor physiological status of plants under abiotic stress conditions. Acta Physiol Plant 2016;38:102. https://doi.org/10.1007/s11738-016- 2113-y

    work page doi:10.1007/s11738-016- 2016

  71. [76]

    Jesus B, Perkins R, Consalvey M. et al. Effects of vertical migrations by benthic microalgae on fluorescence measurements of photo -physiology. Mar Ecol Prog Ser 2006;315:55–66. https://doi.org/10.3354/meps315055

    work page doi:10.3354/meps315055 2006

  72. [77]

    Cartaxana P, Ruivo M, Hubas C. et al. Physiological versus behavioral photoprotection in intertidal epipelic and epipsammic benthic diatom communities. J Exp Mar Biol Ecol 2011;405:120 –7. https://doi.org/10.1016/j.jembe.2011.05.027

    work page doi:10.1016/j.jembe.2011.05.027 2011

  73. [78]

    Jesus B, Brotas V, Ribeiro L. et al. Adaptations of microphytobenthos assemblages to sediment type and tidal position. Cont Shelf Res 2009;29:1624–34. https://doi.org/10.1016/j.csr.2009.05.006

    work page doi:10.1016/j.csr.2009.05.006 2009

  74. [79]

    Barnett A, Méléder V, Blommaert L. et al. Growth form defines physiological photoprotective capacity in intertidal benthic diatoms. ISME J 2015;9:32–45. https://doi.org/10.1038/ismej.2014.105

    work page doi:10.1038/ismej.2014.105 2015

  75. [80]

    Cheng K, Li H, Laszakovits JR. et al. Probing the photochemical formation of hydroxyl radical from dissolved organic matter. Environ Sci Technol 2025;59:2245–56. https://doi.org/10.1021/acs.est.4c10348

    work page doi:10.1021/acs.est.4c10348 2025

  76. [81]

    Production of extracellular reactive oxygen species by phytoplankton

    Diaz JM, Plummer S. Production of extracellular reactive oxygen species by phytoplankton. J Plankton Res 2018;40:655 –66. https://doi.org/10.1093/plankt/fby039

    work page doi:10.1093/plankt/fby039 2018

  77. [82]

    Amorim DS, Amorim IS, Chisté RC. et al. Effects of cold plasma on chlorophylls, carotenoids, anthocyanins, and betalains. Food Res Int 2023;167:112593. https://doi.org/10.1016/j.foodres.2023.112593

    work page doi:10.1016/j.foodres.2023.112593 2023

  78. [83]

    Ataka R, Taniguchi T, Monde K. et al. Determination of N -centered stereochemistry in N22 -methylated chlorophyll -a derivatives and their epimer-dependent optical spectra. Chirality 2024;36. https://doi.org/10.1002/chir.23681

    work page doi:10.1002/chir.23681 2024

  79. [84]

    Carrara F, Sengupta A, Behrendt L. et al. Bistability in oxidative stress response determines the migration behavior of phytoplankton in turbulence. Proc Natl Acad Sci USA 2021;118. https://doi.org/10.1073/pnas.2005944118

    work page doi:10.1073/pnas.2005944118 2021

  80. [85]

    De Carpentier F, Maes A, Marchand CH. et al. How abiotic stress - induced socialization leads to the formation of massive aggregates in Chlamydomonas. Plant Physiol 2022;190:1927 –40. https://doi.org/10.1093/plphys/kiac321

    work page doi:10.1093/plphys/kiac321 2022

Showing first 80 references.