<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing with OASIS Tables v3.0 20080202//EN" "https://jats.nlm.nih.gov/nlm-dtd/publishing/3.0/journalpub-oasis3.dtd">
<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:oasis="http://docs.oasis-open.org/ns/oasis-exchange/table" xml:lang="en" dtd-version="3.0" article-type="research-article">
  <front>
    <journal-meta><journal-id journal-id-type="publisher">BG</journal-id><journal-title-group>
    <journal-title>Biogeosciences</journal-title>
    <abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">1726-4189</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/bg-23-3735-2026</article-id><title-group><article-title>The impact of large-scale macroalgae cultivation and harvesting strategies on the marine carbon dioxide removal efficacy and marine biogeochemistry</article-title><alt-title>Macroalgae cultivation and marine biogeochemistry</alt-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Anugerahanti</surname><given-names>Prima</given-names></name>
          <email>prianu@noc.ac.uk</email>
        <ext-link>https://orcid.org/0000-0002-5033-7934</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Palmiéri</surname><given-names>Julien</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-0226-5243</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Baker</surname><given-names>Chelsey A.</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-0840-2333</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Popova</surname><given-names>Ekaterina</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Yool</surname><given-names>Andrew</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-9879-2776</ext-link></contrib>
        <aff id="aff1"><label>1</label><institution>National Oceanography Centre, Joseph Proudman Building, 6 Brownlow Street, Liverpool L3 5DA, UK</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>National Oceanography Centre, European Way, Southampton SO14 3ZH, UK</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Prima Anugerahanti (prianu@noc.ac.uk)</corresp></author-notes><pub-date><day>5</day><month>June</month><year>2026</year></pub-date>
      
      <volume>23</volume>
      <issue>11</issue>
      <fpage>3735</fpage><lpage>3754</lpage>
      <history>
        <date date-type="received"><day>29</day><month>October</month><year>2025</year></date>
           <date date-type="rev-request"><day>1</day><month>December</month><year>2025</year></date>
           <date date-type="rev-recd"><day>24</day><month>April</month><year>2026</year></date>
           <date date-type="accepted"><day>4</day><month>May</month><year>2026</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2026 Prima Anugerahanti et al.</copyright-statement>
        <copyright-year>2026</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026.html">This article is available from https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026.pdf</self-uri>
      <abstract><title>Abstract</title>

      <p id="d2e126">The large-scale cultivation of macroalgae has been proposed as a marine carbon dioxide removal (mCDR) strategy, yet its efficiency and consequences for ocean biogeochemistry remain uncertain. Using a new macroalgae aquaculture module within an ocean biogeochemistry model, NEMO-MEDUSA, we investigate carbon removal potential and biogeochemical feedbacks under hypothetical global-scale macroalgae cultivation with varying harvest strategies, loss rates, and iron availability. Overall cultivation enhances air–sea <inline-formula><mml:math id="M1" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake by 11.0 <inline-formula><mml:math id="M2" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, but only <inline-formula><mml:math id="M3" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 27 % of macroalgal production results in additional <inline-formula><mml:math id="M4" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake. Furthermore, phytoplankton and zooplankton biomass is suppressed by almost 50 % and is geographically displaced by significant surface nutrient changes. Sinking of harvested biomass increases oxygen demand during remineralisation, leading to widespread oxygen depletion and the emergence of suboxic conditions at the seafloor in deposition regions. When macroalgal growth is not supplemented with iron micronutrient, its production declines sharply (<inline-formula><mml:math id="M5" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>74 %), revealing a significant limitation for large-scale feasibility. Collectively, our results reveal that large-scale macroalgal cultivation offers low mCDR potential, that it is both spatially extensive and locally intensive, and its unintended biogeochemical consequences can be substantial. Our findings highlight the urgent need to assess nutrient constraints and ecological trade-offs before considering this method as a viable large-scale mCDR strategy.</p>
  </abstract>
    
<funding-group>
<award-group id="gs1">
<funding-source>Natural Environment Research Council</funding-source>
<award-id>NE/W004895/1</award-id>
<award-id>NE/Y005589/1</award-id>
</award-group>
</funding-group>
</article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d2e194">Since the adoption of the Paris Agreement in 2015 and the publication of the IPCC Sixth Assessment Report, carbon dioxide removal (CDR) strategies have become increasingly recognised as essential to meet the global climate targets <xref ref-type="bibr" rid="bib1.bibx37" id="paren.1"/>. Among marine-based approaches, the cultivation of macroalgae (seaweed) has been suggested as a promising CDR strategy, due to its efficient uptake of dissolved inorganic carbon (DIC), facilitated by its high carbon-to-nitrogen (<inline-formula><mml:math id="M6" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula>) ratio <xref ref-type="bibr" rid="bib1.bibx26 bib1.bibx28 bib1.bibx68" id="paren.2"/>. In addition to this potential carbon sequestration, macroalgae can confer multiple ecological co-benefits depending on their deployment, including providing habitat for invertebrates and fish, mitigation of eutrophication, and shoreline protection <xref ref-type="bibr" rid="bib1.bibx9 bib1.bibx22 bib1.bibx25" id="paren.3"/>. Furthermore, macroalgae may act as a substitute for carbon-intensive industrial products, such as food, animal feed, and biofuels, and contribute indirectly to emission reductions <xref ref-type="bibr" rid="bib1.bibx24" id="paren.4"/>. These attributes have garnered interest in scaling macroalgae aquaculture offshore, which may increase its potential contribution to marine CDR <xref ref-type="bibr" rid="bib1.bibx53 bib1.bibx28" id="paren.5"/>.</p>
      <p id="d2e225">Relative to terrestrial agriculture, macroalgae aquaculture, which predominantly occurs in coastal zones, requires minimal fertiliser aside from iron (Fe) supplementation <xref ref-type="bibr" rid="bib1.bibx81" id="paren.6"/> and has lower associated emissions <xref ref-type="bibr" rid="bib1.bibx42" id="paren.7"/>. Current applications are predominantly toward food production <xref ref-type="bibr" rid="bib1.bibx52" id="paren.8"/> and producing biochar for agricultural use <xref ref-type="bibr" rid="bib1.bibx62" id="paren.9"/>. However, most of these uses return the carbon fixed by macroalgae to the atmosphere on timescales too short to be climate-relevant, so does not provide a long-term reservoir for carbon sequestration. In natural systems, a substantial fraction of macroalgal NPP is released as particulate and dissolved organic carbon (POC and DOC), as demonstrated by experimental and field-based estimates <xref ref-type="bibr" rid="bib1.bibx41 bib1.bibx17" id="paren.10"/>, which may either be consumed and/or respired or exported to the deep ocean or seafloor sediments where long-term storage may occur over climate-relevant timescales <xref ref-type="bibr" rid="bib1.bibx43" id="paren.11"/>. In contrast, aquaculture practices commonly involve harvesting the biomass at the end of the growing season or after it reaches a target biomass <xref ref-type="bibr" rid="bib1.bibx4" id="paren.12"/>. While cultivated macroalgae release DOC and POC during growth, the removal of standing biomass during harvest limits the macroalgae-sequestered carbon, than can be exported to the deep ocean, and therefore limiting the contribution to climate-relevant carbon sequestration <xref ref-type="bibr" rid="bib1.bibx35" id="paren.13"/>. To ensure durable storage of macroalgae carbon, active sinking of harvested biomass to the deep ocean has been proposed as a potential strategy <xref ref-type="bibr" rid="bib1.bibx28 bib1.bibx2" id="paren.14"/>, despite studies suggesting that replacing carbon intensive products, such as fertiliser and fuels, with macroalgae-based products would have greater potential for reducing atmospheric greenhouse gas than by sinking macroalgae for sequestration <xref ref-type="bibr" rid="bib1.bibx63 bib1.bibx16" id="paren.15"/>.</p>
      <p id="d2e259">There are two critical challenges in evaluating the efficacy of all mCDR methods: (1) quantifying additional atmospheric <inline-formula><mml:math id="M7" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake by the ocean relative to an established baseline, and (2) robustly estimating durability of additional carbon storage <xref ref-type="bibr" rid="bib1.bibx14" id="paren.16"/>. Macroalgae photosynthesis induces a local DIC deficit in seawater, shifting air–sea <inline-formula><mml:math id="M8" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> equilibrium, which promotes <inline-formula><mml:math id="M9" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake from the atmosphere <xref ref-type="bibr" rid="bib1.bibx35" id="paren.17"/>. The rate of equilibration depends on environmental conditions: weeks to months in coastal areas, several months to a year in open-ocean settings, and more than a year in high-latitude or ice-covered regions <xref ref-type="bibr" rid="bib1.bibx40 bib1.bibx5" id="paren.18"/>. If DIC-deficient water is rapidly subducted before atmospheric equilibration occurs, <inline-formula><mml:math id="M10" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> removal may not occur <xref ref-type="bibr" rid="bib1.bibx35 bib1.bibx5" id="paren.19"/>.</p>
      <p id="d2e319">When assessing the CDR potential of macroalgae cultivation, the minimum criterion is that, in utilising the resources and space of natural ecosystems, their activity sequesters more atmospheric <inline-formula><mml:math id="M11" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> than the background natural system. In an observational study, <xref ref-type="bibr" rid="bib1.bibx38" id="text.20"/> found that phytoplankton can be as effective at driving ocean uptake of <inline-formula><mml:math id="M12" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> as cultivated macroalgae. Since both macroalgae and phytoplankton rely on the same limiting resources, such as light, nitrogen (N), and phosphorus (P), intensive macroalgae aquaculture may reduce phytoplankton biomass through competitive exclusion <xref ref-type="bibr" rid="bib1.bibx15" id="paren.21"/>, potentially diminishing primary production of macroalgae and phytoplankton leading to an increase in outgassing <xref ref-type="bibr" rid="bib1.bibx11" id="paren.22"/>. On larger spatial scales, macroalgae nutrient uptake could alter regional nutrient fluxes. For example, in the Southern Ocean, large-scale cultivation may induce “nutrient trapping” <xref ref-type="bibr" rid="bib1.bibx79" id="paren.23"/>, a phenomenon where nutrient removal and transport to the deep ocean by macroalgae limits downstream phytoplankton production in the low latitudes because less nutrients are transported out of the Southern Ocean, as previously simulated in ocean Fe fertilisation experiments <xref ref-type="bibr" rid="bib1.bibx70" id="paren.24"/>.</p>
      <p id="d2e361">Prior to large-scale implementation of macroalgae cultivation, it is essential to estimate the potential CDR efficiency and assess its impact on marine biogeochemistry. Global ocean biogeochemical (OBGC) models provide a suitable scale for undertaking these assessments. Previous studies have estimated the potential of large-scale macroalgae CDR under different assumptions, such as different nutrient scenarios <xref ref-type="bibr" rid="bib1.bibx4 bib1.bibx11 bib1.bibx79" id="paren.25"/>. Large-scale simulations of macroalgae cultivation consistently show that nutrient limitations <xref ref-type="bibr" rid="bib1.bibx4" id="text.26"/> and competition with phytoplankton <xref ref-type="bibr" rid="bib1.bibx79 bib1.bibx11" id="text.27"/> can reduce macroalgae production, air–sea <inline-formula><mml:math id="M13" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux, and therefore CDR efficiency. Moreover, large-scale deployment could negatively impact ocean ecosystems; such as lowering phytoplankton productivity, expanding oxygen minimum zones <xref ref-type="bibr" rid="bib1.bibx79 bib1.bibx11" id="paren.28"/>, and can lead to net decreases in carbon sequestration <xref ref-type="bibr" rid="bib1.bibx5" id="paren.29"/>.</p>
      <p id="d2e391">To optimise large-scale offshore macroalgae CDR, it is important to assess constraints beyond nutrient limitation, such as considering other cultivation protocols, micronutrient fertlisation and limitation, biomass sinking, and remineralisation pathways as these affect CDR efficiency and may impede natural carbon cycling.  We address three key questions in this study: <list list-type="order"><list-item>
      <p id="d2e396">To what extent can macroalgae cultivation and harvesting lead to additional oceanic <inline-formula><mml:math id="M14" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake?</p></list-item><list-item>
      <p id="d2e411">How does large-scale macroalgae cultivation modify nutrient and light availability, and what are the resulting biogeochemical responses of phytoplankton and the biological pump?</p></list-item><list-item>
      <p id="d2e415">What are the consequences of remineralisation-induced deoxygenation at depth when we sink harvested macroalgae biomass?</p></list-item></list></p>
      <p id="d2e418">We investigate these questions using global simulations of an OBGC model that includes macroalgae in a new marine aquaculture submodel, simulated under the “near-present-day” period under observational forcing. This framework allows us to explore a range of cultivation and harvesting scenarios to assess mCDR efficacy and evaluate its wider impacts on marine ecosystems and processes.</p>
      <p id="d2e421">Our study extends previous global assessments of macroalgae cultivation (e.g. <xref ref-type="bibr" rid="bib1.bibx79" id="altparen.30"/>; <xref ref-type="bibr" rid="bib1.bibx4" id="altparen.31"/>; <xref ref-type="bibr" rid="bib1.bibx11" id="altparen.32"/>; <xref ref-type="bibr" rid="bib1.bibx80" id="altparen.33"/>; <xref ref-type="bibr" rid="bib1.bibx12" id="altparen.34"/>) by incorporating a suite of cultivation protocol experiments that explore the effects of harvest threshold, non-harvest loss, biomass extraction, nutrient supplementation, and limitation. This framework allows us to quantify how different modelling assumptions and protocols affect ocean biogeochemistry, including air–sea <inline-formula><mml:math id="M15" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux, nutrient distribution, seafloor oxygen, and phytoplankton NPP, providing a more comprehensive view of large-scale macroalgae cultivation in the global ocean.</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Method</title>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Ocean Model</title>
      <p id="d2e468">Our study uses a macroalgae model embedded within a coupled physical-biogeochemical ocean model. Ocean physics is represented by the Nucleus for European Modelling of the Ocean framework (NEMO; <xref ref-type="bibr" rid="bib1.bibx50" id="altparen.35"/>). NEMO is composed of an ocean general circulation model coupled to a separate sea-ice model, the Sea Ice modelling Integrated Initiative (SI3; <xref ref-type="bibr" rid="bib1.bibx73" id="altparen.36"/>). The NEMO domain used here (the extended ORCA1 grid; eORCA1) is at a horizontal resolution of approximately 1°, and uses a tripolar model grid and incorporates an equatorial band of enhanced resolution. Vertical space in NEMO is resolved into 75 <inline-formula><mml:math id="M16" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula>-levels that range in thickness from approximately 1 <inline-formula><mml:math id="M17" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> at the surface to approximately more than 200 <inline-formula><mml:math id="M18" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> at abyssal depths.</p>
      <p id="d2e500">Ocean biogeochemistry is represented by the Model of Ecosystem Dynamics, nutrient Utilisation, Sequestration and Acidification (MEDUSA; <xref ref-type="bibr" rid="bib1.bibx82" id="altparen.37"/>). MEDUSA is a dual size-class “intermediate complexity” ecosystem/biogeochemistry model that resolves nutrients and two size classes of phytoplankton, zooplankton, and detritus (NPZD) components. The smaller size class includes non-diatom phytoplankton, microzooplankton and small particles of slow-sinking detritus, while the larger size class includes diatoms, mesozooplankton and large particles of fast-sinking detritus (the latter modelled implicitly). The biogeochemical cycles of N, Fe, C, silicon, oxygen, and alkalinity are represented, coupled in both fixed and dynamic stoichiometric relationships.</p>
      <p id="d2e506">Air–sea <inline-formula><mml:math id="M19" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux in NEMO-MEDUSA is calculated prognostically using standard equations for the partition of dissolved inorganic carbon (DIC) into dissolved <inline-formula><mml:math id="M20" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> and carbonic acid, bicarbonate, and carbonate ions. Partitioning is a function of local temperature, salinity, DIC, and total alkalinity, and uses the MOCSY-2.0 carbonate chemistry routines <xref ref-type="bibr" rid="bib1.bibx55" id="paren.38"/>. The exchange of <inline-formula><mml:math id="M21" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> with the atmosphere is then calculated from the dissolved <inline-formula><mml:math id="M22" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> fraction, atmospheric <inline-formula><mml:math id="M23" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, temperature, salinity, and surface winds, following the gas transfer parameterisation of <xref ref-type="bibr" rid="bib1.bibx78" id="text.39"/>. Because gas transfer is finite and wind speed dependent, the rate at which surface ocean <inline-formula><mml:math id="M24" display="inline"><mml:mrow class="chem"><mml:mi>p</mml:mi><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> anomalies equilibrate with the atmosphere is limited, introducing a natural lag between biological carbon drawdown and atmospheric <inline-formula><mml:math id="M25" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake. A full description of MEDUSA can be found in <xref ref-type="bibr" rid="bib1.bibx82" id="text.40"/>, with the current version evaluated in <xref ref-type="bibr" rid="bib1.bibx83" id="text.41"/>.</p>
      <p id="d2e601">The ocean model is forced at its surface boundary using a climatologically-adjusted version of the ERA-5 atmospheric reanalysis <xref ref-type="bibr" rid="bib1.bibx32" id="paren.42"/>. Ocean physical fields and the biogeochemical fields of oxygen and nutrients (N and silicon) are initialised using the World Ocean Atlas 2023 <xref ref-type="bibr" rid="bib1.bibx61" id="paren.43"/>. As simulations are initialised from a time-point prior to the reference period of the GLODAP climatology <xref ref-type="bibr" rid="bib1.bibx46" id="paren.44"/>, dissolved inorganic carbon (DIC) and alkalinity fields are initialised from prior simulations of the model (in the UKESM1 framework; <xref ref-type="bibr" rid="bib1.bibx83" id="altparen.45"/>) to minimise bias in the <inline-formula><mml:math id="M26" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> air–sea flux due to model equilibration. For other plankton tracers we use uniform nominal values. Riverine nutrient inputs are not included in this configuration, so nutrient supply is limited to oceanic initial conditions and internal cycling.</p>
      <p id="d2e628">This initial state is then simulated for two cycles of the forcing period between 1976 to 2024 (2 <inline-formula><mml:math id="M27" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 49 years <inline-formula><mml:math id="M28" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 98 years). This is done so that surface biogeochemical properties reach quasi-equilibrium, minimising the drift during the experimental period when macroalgae are introduced (see Sect. <xref ref-type="sec" rid="Ch1.S2.SS2"/>). Note that, since DIC has a strong secular signal during the 1976–2024 period caused by rising atmospheric <inline-formula><mml:math id="M29" display="inline"><mml:mrow class="chem"><mml:mi>p</mml:mi><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>, its concentration is reset to the 1976 UKESM1 state at the beginning of each forcing cycle, and the rising atmospheric <inline-formula><mml:math id="M30" display="inline"><mml:mrow class="chem"><mml:mi>p</mml:mi><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> is still simulated during the simulation until the end of the cycle. The temperature increase within this simulated period is relatively small compared to late 21st-century warming used in other CDR studies (e.g. in <xref ref-type="bibr" rid="bib1.bibx58" id="altparen.46"/>) and therefore show limited effects on the macroalgae and other biogeochemical tracers.</p>
</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>Macroalgae Module</title>
      <p id="d2e685">mCDR by macroalgae is simulated by adding a new aquaculture module to MEDUSA. This module effectively represents macroalgae as being cultivated on a fabricated floating structure that is anchored and immobile <xref ref-type="bibr" rid="bib1.bibx64 bib1.bibx79" id="paren.47"/>. As such, macroalgae is located on the ocean model's grid but it is not advected by ocean currents, although it interacts with MEDUSA's advected passive tracers. We assume that macroalgae cultivation started from the last 20 years of the control run (2004–2023), starting from a small initial biomass of 0.01 <inline-formula><mml:math id="M31" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>. This initial value is consistent with other biological tracers in MEDUSA and serves as an “inoculation”, after which biomass grows in response to temperature, light and nutrient limitation.</p>
      <p id="d2e711">Fundamentally, the macroalgae growth and loss processes are similar to those of phytoplankton with the key differences subsequently outlined. Unlike MEDUSA's phytoplankton, modelled macroalgae biomass is static within the model grid and has a restricted vertical extent, by default from approximately 5 to 30 <inline-formula><mml:math id="M32" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> in the water column (10 model levels) following previous modelling studies <xref ref-type="bibr" rid="bib1.bibx79 bib1.bibx80" id="paren.48"/> and offshore cultivation experiment <xref ref-type="bibr" rid="bib1.bibx86 bib1.bibx71" id="paren.49"/>. This depth range is intended to represent large-scale offshore cultivation and differs from nearshore farming practices, where shallower deployments are common (e.g. <xref ref-type="bibr" rid="bib1.bibx18 bib1.bibx65 bib1.bibx36" id="altparen.50"/>). In an initial simulation, we found negligible differences between cultivating macroalgae between 1–30 and 5–30 <inline-formula><mml:math id="M33" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> (results not shown).  Functionally, macroalgae growth is loosely based on that of MEDUSA's phytoplankton, with separate chlorophyll (Chl) and N biomass, and with its rate governed by temperature and the availability of light and dissolved inorganic nitrogen (DIN) and iron (DFe). No external nutrient inputs or pulsed fertilisation are applied. As a result macroalgae compete directly with phytoplankton for the same nutrient resources. Fig. <xref ref-type="fig" rid="F1"/> presents a schematic of the macroalgae model and its relationships with MEDUSA's dissolved tracers.</p>

      <fig id="F1"><label>Figure 1</label><caption><p id="d2e744">Schematic diagram of the macroalgae model. Macroalgae is grown 5 <inline-formula><mml:math id="M34" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> below the surface, consuming C, N, and Fe. In the model, we limit the macroalgae growth to 30 <inline-formula><mml:math id="M35" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula>. Seaweed cultivation includes non-harvest loss and harvesting, where the latter is directly sunk to the seafloor, contributing to seafloor DIN, DIC, DFe, and Oxygen.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f01.png"/>

        </fig>

      <p id="d2e770">In terms of loss processes, while the simulated macroalgae incur non-harvest losses, they are assumed not to be explicitly grazed by zooplankton. However, their cultivation involves active harvesting, the precise mechanism and fate of which forms part of the experimental design described below.</p>
      <p id="d2e773">Following the work of <xref ref-type="bibr" rid="bib1.bibx4" id="text.51"/>, we use multiple species in our macroalgae module with: two cold water species (<italic>Macrocystis</italic> and <italic>Saccharina</italic>) and two warm water species (<italic>Sargassum</italic> and <italic>Euchema</italic>). These taxa span a board range of thermal niches and cultivation context: cold water species usually occupies high latitude nutrient-rich upwelling system, commonly found in the North Atlantic aquaculture industry <xref ref-type="bibr" rid="bib1.bibx75" id="paren.52"/>, while warm water species are predominantly cultivated in tropical and subtropical regions <xref ref-type="bibr" rid="bib1.bibx31 bib1.bibx51" id="paren.53"/>. All four are represented in identical functional from, with a number of shared parameters, but each species has different values for certain key parameters. These include optimal temperature range (between <inline-formula><mml:math id="M36" display="inline"><mml:mrow><mml:msubsup><mml:mi>T</mml:mi><mml:mtext>opt</mml:mtext><mml:mn mathvariant="normal">1</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M37" display="inline"><mml:mrow><mml:msubsup><mml:mi>T</mml:mi><mml:mtext>opt</mml:mtext><mml:mn mathvariant="normal">2</mml:mn></mml:msubsup></mml:mrow></mml:math></inline-formula>), half-saturation constant for DIN uptake (<inline-formula><mml:math id="M38" display="inline"><mml:mrow><mml:msub><mml:mi>k</mml:mi><mml:mtext>DIN</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula>), and <inline-formula><mml:math id="M39" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio (<inline-formula><mml:math id="M40" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="normal">Θ</mml:mi><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) adopted from <xref ref-type="bibr" rid="bib1.bibx4" id="text.54"/>. Although these taxa differ in their real-world cultivation methods and physiological traits, they are represented using a common idealised cultivation framework to isolate first-order biogeochemical effects at large spatial scales. We acknowledge that growth rates, carbon content, and other physiological traits vary considerably across and within taxa in reality <xref ref-type="bibr" rid="bib1.bibx66" id="paren.55"/>, but our choice rather reflects a functional representation designed to capture first-order biogeochemical effects at global scale. Our analysis therefore focuses on the sensitivity of large-scale biogeochemical responses to macroalgal cultivation.</p>
      <p id="d2e870">Macroalgae growth is limited by temperature (<inline-formula><mml:math id="M41" display="inline"><mml:mrow><mml:msub><mml:mi>T</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>), light (<inline-formula><mml:math id="M42" display="inline"><mml:mrow><mml:msub><mml:mi>L</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>), and nutrient (<inline-formula><mml:math id="M43" display="inline"><mml:mrow><mml:msub><mml:mi>N</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>), and its losses comprises of non-harvest loss <inline-formula><mml:math id="M44" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Loss</mml:mi><mml:mi mathvariant="normal">noharv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> and harvesting loss <inline-formula><mml:math id="M45" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Loss</mml:mi><mml:mi mathvariant="normal">harv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>. Therefore the change in each macroalgae type, <inline-formula><mml:math id="M46" display="inline"><mml:mrow><mml:msub><mml:mi>M</mml:mi><mml:mi mathvariant="normal">n</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> is calculated by:

            <disp-formula id="Ch1.E1" content-type="numbered"><label>1</label><mml:math id="M47" display="block"><mml:mrow><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mi mathvariant="normal">d</mml:mi><mml:msub><mml:mi>M</mml:mi><mml:mi mathvariant="normal">n</mml:mi></mml:msub></mml:mrow><mml:mrow><mml:mi mathvariant="normal">d</mml:mi><mml:mi>t</mml:mi></mml:mrow></mml:mfrac></mml:mstyle><mml:mo>=</mml:mo><mml:mrow class="chem"><mml:mi mathvariant="normal">Growth</mml:mi></mml:mrow><mml:mo>-</mml:mo><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Loss</mml:mi><mml:mi mathvariant="normal">noharv</mml:mi></mml:msub></mml:mrow><mml:mo>-</mml:mo><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Loss</mml:mi><mml:mi mathvariant="normal">harv</mml:mi></mml:msub></mml:mrow></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e981">The temperature limitation term is similar to a Gaussian probability curve with flat peak, adopted from <xref ref-type="bibr" rid="bib1.bibx4" id="text.56"/>:

                <disp-formula specific-use="align" content-type="numbered"><mml:math id="M48" display="block"><mml:mtable displaystyle="true"><mml:mlabeledtr id="Ch1.E2"><mml:mtd><mml:mtext>2</mml:mtext></mml:mtd><mml:mtd><mml:mstyle class="stylechange" displaystyle="true"/></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:msub><mml:mi>T</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mi>exp⁡</mml:mi><mml:mfenced open="(" close=")"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>T</mml:mi><mml:mo>-</mml:mo><mml:msub><mml:mi>T</mml:mi><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>,</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mtext>opt</mml:mtext></mml:mrow></mml:msub><mml:msup><mml:mo>)</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:mfenced><mml:mo>,</mml:mo><mml:mtext>    </mml:mtext><mml:mi>T</mml:mi><mml:mo>&lt;</mml:mo><mml:msub><mml:mi>T</mml:mi><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>,</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mtext>opt</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mlabeledtr id="Ch1.E3"><mml:mtd><mml:mtext>3</mml:mtext></mml:mtd><mml:mtd><mml:mstyle class="stylechange" displaystyle="true"/></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:msub><mml:mi>T</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mi>exp⁡</mml:mi><mml:mfenced close=")" open="("><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub><mml:mo>(</mml:mo><mml:mi>T</mml:mi><mml:mo>-</mml:mo><mml:msub><mml:mi>T</mml:mi><mml:mrow><mml:mn mathvariant="normal">2</mml:mn><mml:mo>,</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mtext>opt</mml:mtext></mml:mrow></mml:msub><mml:msup><mml:mo>)</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:mfenced><mml:mo>,</mml:mo><mml:mtext>    </mml:mtext><mml:mi>T</mml:mi><mml:mo>&gt;</mml:mo><mml:msub><mml:mi>T</mml:mi><mml:mrow><mml:mn mathvariant="normal">2</mml:mn><mml:mo>,</mml:mo><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mtext>opt</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mlabeledtr id="Ch1.E4"><mml:mtd><mml:mtext>4</mml:mtext></mml:mtd><mml:mtd><mml:mstyle displaystyle="true" class="stylechange"/></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:msub><mml:mi>T</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn><mml:mo>,</mml:mo><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:msub><mml:mi>T</mml:mi><mml:mtext>opt</mml:mtext></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr></mml:mtable></mml:math></disp-formula></p>
      <p id="d2e1157">Where <inline-formula><mml:math id="M49" display="inline"><mml:mrow><mml:msub><mml:mi>T</mml:mi><mml:mtext>opt</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula> is a 5 <inline-formula><mml:math id="M50" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">°</mml:mi><mml:mi mathvariant="normal">C</mml:mi></mml:mrow></mml:math></inline-formula> optimal temperature range for each macroalgae group. <inline-formula><mml:math id="M51" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M52" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> are adjusted near the lower (<inline-formula><mml:math id="M53" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) and upper temperature limits (<inline-formula><mml:math id="M54" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) to reach zero, respectively. The temperature limitation function acts as a smooth growth filter rather than a binary on-off switch. As such, growth declines gradually as temperature departs from the optimal range for each represented taxon, ensuring more realistic transitions across latitude and season.</p>
      <p id="d2e1226">Light limitation is formulated in the same way as in phytoplankton:

            <disp-formula id="Ch1.E5" content-type="numbered"><label>5</label><mml:math id="M55" display="block"><mml:mrow><mml:msub><mml:mi>L</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mi mathvariant="italic">μ</mml:mi><mml:msub><mml:mi>T</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:msub><mml:mover accent="true"><mml:mi mathvariant="italic">α</mml:mi><mml:mo mathvariant="normal" stretchy="false">^</mml:mo></mml:mover><mml:mi mathvariant="normal">m</mml:mi></mml:msub><mml:mi>I</mml:mi></mml:mrow><mml:mrow><mml:msup><mml:mfenced close=")" open="("><mml:mrow><mml:mo>(</mml:mo><mml:mi mathvariant="italic">μ</mml:mi><mml:msub><mml:mi>T</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:msup><mml:mo>)</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>+</mml:mo><mml:msubsup><mml:mover accent="true"><mml:mi mathvariant="italic">α</mml:mi><mml:mo stretchy="false" mathvariant="normal">^</mml:mo></mml:mover><mml:mi mathvariant="normal">m</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msubsup><mml:msup><mml:mi>I</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:mfenced><mml:mn mathvariant="normal">0.5</mml:mn></mml:msup></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1299">Where <inline-formula><mml:math id="M56" display="inline"><mml:mi>I</mml:mi></mml:math></inline-formula> is the irradiance and <inline-formula><mml:math id="M57" display="inline"><mml:mover accent="true"><mml:mi mathvariant="italic">α</mml:mi><mml:mo mathvariant="normal" stretchy="false">^</mml:mo></mml:mover></mml:math></inline-formula> is the initial slope of photosynthesis-irradiance curve, so that macroalgae with a high chlorophyll content have an elevated response to irradiance.</p>
      <p id="d2e1319">Macroalgae compete with phytoplankton to obtain nutrients (<inline-formula><mml:math id="M58" display="inline"><mml:mi>n</mml:mi></mml:math></inline-formula>), and nutrient uptake (<inline-formula><mml:math id="M59" display="inline"><mml:mrow><mml:msub><mml:mi>n</mml:mi><mml:mtext>up</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula>) is formulated using Michaelis–Menten kinetics:

            <disp-formula id="Ch1.E6" content-type="numbered"><label>6</label><mml:math id="M60" display="block"><mml:mrow><mml:msub><mml:mi>n</mml:mi><mml:mtext>up</mml:mtext></mml:msub><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mi>n</mml:mi><mml:mrow><mml:msub><mml:mi>k</mml:mi><mml:mi mathvariant="normal">m</mml:mi></mml:msub><mml:mo>+</mml:mo><mml:mi>n</mml:mi></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1366">Unlike the default formula for phytoplankton, in the macroalgae default simulation, we assume that Fe is supplemented (<inline-formula><mml:math id="M61" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Fe</mml:mi><mml:mi mathvariant="normal">up</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M62" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> =1) and only DIN is the limiting nutrient. However, we allow multiple nutrient limitation, by using the Liebig's law of minimum, described below (<inline-formula><mml:math id="M63" display="inline"><mml:mrow><mml:msub><mml:mi>N</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>):

            <disp-formula id="Ch1.E7" content-type="numbered"><label>7</label><mml:math id="M64" display="block"><mml:mrow><mml:msub><mml:mi>N</mml:mi><mml:mi mathvariant="normal">L</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mtext>min</mml:mtext><mml:mo>(</mml:mo><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">DIN</mml:mi><mml:mi mathvariant="normal">up</mml:mi></mml:msub></mml:mrow><mml:mo>,</mml:mo><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Fe</mml:mi><mml:mi mathvariant="normal">up</mml:mi></mml:msub></mml:mrow><mml:mo>)</mml:mo></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1430">In addition to harvesting, simulated macroalgae experience non-harvest losses, which represents biomass losses outside deliberate harvesting, such as physical erosion, aging, fragmentation, and implicit grazing processes. Non-harvesting loss are transferred directly to the slow detrital pool, where they are remineralised according to the standard MEDUSA slow sinking detritus formulation. As a result, processes such as direct DOC release during production are implicitly subsumed within the detrital pathway rather than treated explicitly:

            <disp-formula id="Ch1.E8" content-type="numbered"><label>8</label><mml:math id="M65" display="block"><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Loss</mml:mi><mml:mi mathvariant="normal">noharv</mml:mi></mml:msub></mml:mrow><mml:mo>=</mml:mo><mml:mi>m</mml:mi><mml:msub><mml:mn mathvariant="normal">2</mml:mn><mml:mtext>max</mml:mtext></mml:msub><mml:msub><mml:mi>M</mml:mi><mml:mi mathvariant="normal">n</mml:mi></mml:msub></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1458">Where <inline-formula><mml:math id="M66" display="inline"><mml:mrow><mml:mi>m</mml:mi><mml:msub><mml:mn mathvariant="normal">2</mml:mn><mml:mtext>max</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula> is the rate of mortality.</p>
      <p id="d2e1474">For harvesting, once total integrated macroalgae biomass at a grid point has reached the target harvesting threshold (<inline-formula><mml:math id="M67" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">TW</mml:mi></mml:mrow></mml:math></inline-formula>), it is harvested over the course of a single simulated day, using a linear loss term and parameter <inline-formula><mml:math id="M68" display="inline"><mml:mrow><mml:mi>m</mml:mi><mml:msub><mml:mn mathvariant="normal">1</mml:mn><mml:mtext>max</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula>. Otherwise, the macroalgae continues to grow.

            <disp-formula id="Ch1.E9" content-type="numbered"><label>9</label><mml:math id="M69" display="block"><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">Loss</mml:mi><mml:mi mathvariant="normal">harv</mml:mi></mml:msub></mml:mrow><mml:mo>=</mml:mo><mml:mi>m</mml:mi><mml:msub><mml:mn mathvariant="normal">1</mml:mn><mml:mtext>max</mml:mtext></mml:msub><mml:msub><mml:mi>M</mml:mi><mml:mi mathvariant="normal">n</mml:mi></mml:msub></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1524">In the default case, harvested macroalgae biomass is assumed to be baled-up and immediately sunk to the seafloor in the same water column, so that the associated carbon biomass is durably stored and isolated from the surface ocean and atmosphere. This process is formulated similarly to that of fast-sinking organic detritus in MEDUSA (see Sect. 2.3.10 of <xref ref-type="bibr" rid="bib1.bibx82" id="altparen.57"/>), but with a different remineralisation length-scale, and without the influence of any biomineral ballasting. The value of the length-scale is set to considerably deeper than the seafloor (40 <inline-formula><mml:math id="M70" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">km</mml:mi></mml:mrow></mml:math></inline-formula>) to represent this active baling and deposition that aims to ensure that the harvested biomass reaches the seafloor does not undergo substantial decay. The organic material reaching the seafloor in this way is added to MEDUSA's existing benthic reservoirs and is remineralised from these in the same manner as material arriving via the natural biological pump (i.e. slow remineralisation to DIN, DIC and DFe with the associated consumption of oxygen). This approach avoids cultivated macroalgae biomass undergoing remineralisation in the upper water column and increases the efficiency with which fixed <inline-formula><mml:math id="M71" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> is stored in the ocean (and removed from exchange with the atmosphere). Parameter values used in these equations are described in Table <xref ref-type="table" rid="T1"/>.</p>

<table-wrap id="T1" specific-use="star"><label>Table 1</label><caption><p id="d2e1554">Parameter table for different macroalgae species. The different macroalgae parameters are based on <xref ref-type="bibr" rid="bib1.bibx4" id="text.58"/> (Supplementary material). <inline-formula><mml:math id="M72" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio is based on the initial cell quota of different macroalgae species from the same study. We apply a common set of physiological parameters across macroalgal taxa in order to focus on first-order biogeochemical responses rather than species-specific physiological variability.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="7">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left" colsep="1"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:colspec colnum="7" colname="col7" align="right"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Parameter</oasis:entry>
         <oasis:entry colname="col2">Symbol</oasis:entry>
         <oasis:entry colname="col3">Units</oasis:entry>
         <oasis:entry colname="col4"><italic>Saccharina</italic></oasis:entry>
         <oasis:entry colname="col5"><italic>Sargassum</italic></oasis:entry>
         <oasis:entry colname="col6"><italic>Eucheuma</italic></oasis:entry>
         <oasis:entry colname="col7"><italic>Macrocystis</italic></oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">DIN half-saturation uptake constant</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M73" display="inline"><mml:mrow><mml:msub><mml:mi>k</mml:mi><mml:mtext>DIN</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M74" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4">2.00</oasis:entry>
         <oasis:entry colname="col5">2.95</oasis:entry>
         <oasis:entry colname="col6">5.60</oasis:entry>
         <oasis:entry colname="col7">10.13</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Optimum Temperature 1</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M75" display="inline"><mml:mrow><mml:msub><mml:mi>T</mml:mi><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>,</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mtext>opt</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M76" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">°</mml:mi><mml:mi mathvariant="normal">C</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4">10.0</oasis:entry>
         <oasis:entry colname="col5">20.5</oasis:entry>
         <oasis:entry colname="col6">22.5</oasis:entry>
         <oasis:entry colname="col7">13.0</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Optimum Temperature 2</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M77" display="inline"><mml:mrow><mml:msub><mml:mi>T</mml:mi><mml:mrow><mml:mn mathvariant="normal">2</mml:mn><mml:mo>,</mml:mo><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mtext>opt</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M78" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">°</mml:mi><mml:mi mathvariant="normal">C</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4">15.0</oasis:entry>
         <oasis:entry colname="col5">25.5</oasis:entry>
         <oasis:entry colname="col6">27.5</oasis:entry>
         <oasis:entry colname="col7">18.0</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Biomass <inline-formula><mml:math id="M79" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M80" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="normal">Θ</mml:mi><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M81" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>(</mml:mo><mml:mi mathvariant="normal">mol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:msup><mml:mo>)</mml:mo><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4">26.8</oasis:entry>
         <oasis:entry colname="col5">34.3</oasis:entry>
         <oasis:entry colname="col6">34.3</oasis:entry>
         <oasis:entry colname="col7">26.8</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Maximum growth rate</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M82" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">μ</mml:mi><mml:mi mathvariant="normal">m</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M83" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">d</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry namest="col4" nameend="col7" align="center">0.2 <xref ref-type="bibr" rid="bib1.bibx85 bib1.bibx4" id="paren.59"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Chl <inline-formula><mml:math id="M84" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> specific initial slope of <inline-formula><mml:math id="M85" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>-<inline-formula><mml:math id="M86" display="inline"><mml:mi>I</mml:mi></mml:math></inline-formula> curve</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M87" display="inline"><mml:mover accent="true"><mml:mrow><mml:msub><mml:mi mathvariant="italic">α</mml:mi><mml:mi mathvariant="normal">m</mml:mi></mml:msub></mml:mrow><mml:mo stretchy="false" mathvariant="normal">^</mml:mo></mml:mover></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"/>
         <oasis:entry namest="col4" nameend="col7" align="center">15 <xref ref-type="bibr" rid="bib1.bibx27" id="paren.60"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">DFe half-saturation uptake constant</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M88" display="inline"><mml:mrow><mml:msub><mml:mi>k</mml:mi><mml:mtext>DFe</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M89" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">Fe</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry namest="col4" nameend="col7" align="center">1.5 <inline-formula><mml:math id="M90" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup><xref ref-type="bibr" rid="bib1.bibx57" id="paren.61"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Biomass <inline-formula><mml:math id="M92" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">Fe</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M93" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="normal">Θ</mml:mi><mml:mrow class="chem"><mml:mi mathvariant="normal">Fe</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M94" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">mol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mo>(</mml:mo><mml:mi mathvariant="normal">mol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">Fe</mml:mi><mml:msup><mml:mo>)</mml:mo><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry namest="col4" nameend="col7" align="center">5.08 <inline-formula><mml:math id="M95" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup><xref ref-type="bibr" rid="bib1.bibx67" id="paren.62"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Non-harvest loss</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M97" display="inline"><mml:mrow><mml:mi>m</mml:mi><mml:msub><mml:mn mathvariant="normal">2</mml:mn><mml:mtext>max</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M98" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">d</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry namest="col4" nameend="col7" align="center">0.025 <xref ref-type="bibr" rid="bib1.bibx4" id="paren.63"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Adjusted coefficient lower <inline-formula><mml:math id="M99" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> limit</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M100" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"/>
         <oasis:entry namest="col4" nameend="col7" align="center">0.03 <xref ref-type="bibr" rid="bib1.bibx4" id="paren.64"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Adjusted coefficient upper <inline-formula><mml:math id="M101" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> limit</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M102" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">β</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"/>
         <oasis:entry namest="col4" nameend="col7" align="center">0.1 <xref ref-type="bibr" rid="bib1.bibx4" id="paren.65"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Fraction of macroalgae that are harvested</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M103" display="inline"><mml:mrow><mml:mi>m</mml:mi><mml:msub><mml:mn mathvariant="normal">1</mml:mn><mml:mtext>max</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M104" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">d</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry namest="col4" nameend="col7" align="center">0.9 </oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Target harvesting threshold</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M105" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">TH</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M106" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry namest="col4" nameend="col7" align="center">400 <xref ref-type="bibr" rid="bib1.bibx7 bib1.bibx77" id="paren.66"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Carbon content of dry macroalgae</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M107" display="inline"><mml:mrow><mml:msub><mml:mi>C</mml:mi><mml:mtext>DW</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"/>
         <oasis:entry namest="col4" nameend="col7" align="center">0.3 <xref ref-type="bibr" rid="bib1.bibx20" id="paren.67"/></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M108" display="inline"><mml:mrow><mml:mrow class="chem"><mml:mi mathvariant="normal">Wet</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">weight</mml:mi></mml:mrow><mml:mo>:</mml:mo><mml:mrow class="chem"><mml:mi mathvariant="normal">Dry</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">Weight</mml:mi></mml:mrow></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M109" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">WW</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">DW</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3"/>
         <oasis:entry namest="col4" nameend="col7" align="center">9.33 <xref ref-type="bibr" rid="bib1.bibx74" id="paren.68"/></oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

      <p id="d2e2344">Note that MEDUSA resolves biological respiration and remineralisation for both phytoplankton and macroalgae at every depth level and timestep. Air–sea <inline-formula><mml:math id="M110" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux is therefore driven by the net change in surface DIC after all biological production, community respiration, grazing, and remineralisation processes operate, rather than by NPP alone. The net ecosystem production (NEP) signal is thus an emergent property of the model's coupled biological and chemical state.</p>
      <p id="d2e2358">After initialisation and a spin-up period (see Sect. <xref ref-type="sec" rid="Ch1.S2.SS1"/>), we run the control model (NEMO-MEDUSA only) using ERA-5 forcing from 1976 until 2024. We assume that macroalgae cultivation started from the last 20 years of the control run (2004–2023). We will focus on quantifying <inline-formula><mml:math id="M111" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> sequestration, biomass harvested, and assessing how harvesting strategy would affect ocean biogeochemistry including nutrient cycles (N and Fe), phytoplankton net primary production (NPP) and plankton biomass, carbonate chemistry (air–sea <inline-formula><mml:math id="M112" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux), seafloor oxygen, and carbon pool distributions. The results shown in this manuscript are the average from 2015–2024.</p>
</sec>
<sec id="Ch1.S2.SS3">
  <label>2.3</label><title>Experimental Design</title>
      <p id="d2e2393">To assess the biogeochemical impacts of large-scale macroalgae cultivation, we compare a baseline MEDUSA simulation with one incorporating dynamic macroalgae growth (MEDUSA <inline-formula><mml:math id="M113" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae). Building on this framework, we conduct four sensitivity experiments to explore key intervention strategies and test modelling uncertainties: <list list-type="order"><list-item>
      <p id="d2e2405">varying the biomass threshold for harvesting and sinking</p></list-item><list-item>
      <p id="d2e2409">increasing non-harvest loss</p></list-item><list-item>
      <p id="d2e2413">extracting harvested biomass from the ocean system entirely rather than sinking it</p></list-item><list-item>
      <p id="d2e2417">applying Fe limitation.</p></list-item></list></p>
      <p id="d2e2420">These experiments and shorthands are summarised in Table <xref ref-type="table" rid="T2"/>.</p>

<table-wrap id="T2" specific-use="star"><label>Table 2</label><caption><p id="d2e2428">Experiment summary table. Macroalgae experiments are initialised from the Control simulation in 2004 and simulated for 20 years.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="3">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Experiment</oasis:entry>
         <oasis:entry colname="col2">Simulation description</oasis:entry>
         <oasis:entry colname="col3">Run period</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">Control</oasis:entry>
         <oasis:entry colname="col2">NEMO-MEDUSA</oasis:entry>
         <oasis:entry colname="col3">1976–2023</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Default (Harvest 400)</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M114" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae, harvest at 400 <inline-formula><mml:math id="M115" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, Fe fertilised</oasis:entry>
         <oasis:entry colname="col3">2004–2023</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Harvest 200</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M116" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae, harvest at 200 <inline-formula><mml:math id="M117" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, Fe fertilised</oasis:entry>
         <oasis:entry colname="col3">2004–2023</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Harvest 800</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M118" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae, harvest at 800 <inline-formula><mml:math id="M119" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, Fe fertilised</oasis:entry>
         <oasis:entry colname="col3">2004–2023</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">High Loss</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M120" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae, non harvesting loss <inline-formula><mml:math id="M121" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 0.05 <inline-formula><mml:math id="M122" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">d</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, Fe fertilised</oasis:entry>
         <oasis:entry colname="col3">2004–2023</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Extraction</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M123" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae, no biomass sinking, Fe fertilised</oasis:entry>
         <oasis:entry colname="col3">2004–2023</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Fe limitation</oasis:entry>
         <oasis:entry colname="col2"><inline-formula><mml:math id="M124" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> Macroalgae, with Fe limitation</oasis:entry>
         <oasis:entry colname="col3">2004–2023</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

      <p id="d2e2663">In all experiments we define the target biomass as the sum of local integrated macroalgae biomass in <inline-formula><mml:math id="M125" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>. The default target biomass is 400 <inline-formula><mml:math id="M126" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> (500 <inline-formula><mml:math id="M127" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">g</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">dry</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">weight</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>(</mml:mo><mml:mi mathvariant="normal">DW</mml:mi><mml:mo>)</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>), resulting in two harvests a year on average based on studies by <xref ref-type="bibr" rid="bib1.bibx7 bib1.bibx77" id="text.69"/>, although warmer macroalgae species can be harvested up to 12 times a year <xref ref-type="bibr" rid="bib1.bibx72" id="paren.70"/>. To assess the sensitivity of target biomass in harvesting, we vary the weight to 200 and 800 <inline-formula><mml:math id="M128" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">N</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>. Macroalgae are not harvested until they reach these target biomasses, so will continue to grow until they do (and will continue to experience non-harvest losses). Harvesting involves decreasing the biomass of macroalgae by 90 % during the harvest day. Since Fe is one of the limiting nutrients for macroalgae, we prescribe <inline-formula><mml:math id="M129" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">Fe</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio of 5.1 <inline-formula><mml:math id="M130" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup> and half saturation constant of 1.5 <inline-formula><mml:math id="M132" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup> based on laboratory studies <xref ref-type="bibr" rid="bib1.bibx67 bib1.bibx57" id="paren.71"/>. However, in the default simulation (as well as all other experiments except for Fe limitation), we assume that macroalgae are only limited by DIN. This corresponds to an assumption that Fe is supplemented, following established farming practices <xref ref-type="bibr" rid="bib1.bibx81" id="text.72"/>, and that it is at non-limiting (replete) concentrations. When macroalgae is being remineralised, the added Fe will also be released.</p>
      <p id="d2e2824">To determine whether CDR is efficient within a certain region, we use two different CDR efficiency metrics; efficiency calculated using macroalgae NPP (<inline-formula><mml:math id="M134" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) and harvested macroalgae biomass (<inline-formula><mml:math id="M135" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>). <inline-formula><mml:math id="M136" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> can be calculated as the proportion between macroalgae NPP and the additional air–sea <inline-formula><mml:math id="M137" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux within each grid cell (CDR flux) <xref ref-type="bibr" rid="bib1.bibx11" id="paren.73"/>:

            <disp-formula id="Ch1.E10" content-type="numbered"><label>10</label><mml:math id="M138" display="block"><mml:mtable class="split" rowspacing="0.2ex" displaystyle="true" columnalign="right left"><mml:mtr><mml:mtd><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow><mml:mo>=</mml:mo></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mspace linebreak="nobreak" width="0.25em"/><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msub><mml:mi mathvariant="normal">flux</mml:mi><mml:mi mathvariant="normal">macroalgae</mml:mi></mml:msub></mml:mrow><mml:mo>-</mml:mo><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msub><mml:mi mathvariant="normal">flux</mml:mi><mml:mi mathvariant="normal">control</mml:mi></mml:msub></mml:mrow></mml:mrow><mml:mrow class="chem"><mml:mi mathvariant="normal">Macroalgae</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd/><mml:mtd><mml:mrow><mml:mo>×</mml:mo><mml:mn mathvariant="normal">100</mml:mn><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="italic">%</mml:mi></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula></p>

      <fig id="F2" specific-use="star"><label>Figure 2</label><caption><p id="d2e2960">Macroalgae Biomass <bold>(a)</bold>, NPP <bold>(b)</bold>, harvest <bold>(c)</bold>, and percentage of how much NPP are harvested <bold>(d)</bold> from the default experiment averaged between 2015–2024.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f02.png"/>

        </fig>

      <p id="d2e2981">If annual average macroalgal NPP associated carbon is equivalent to the annual average of additional atmospheric <inline-formula><mml:math id="M139" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake within a grid cell then the CDR efficiency would be 100 %, as it is assumed that the macroalgae-induced DIC deficit within the grid cell has driven the <inline-formula><mml:math id="M140" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake. The calculation of <inline-formula><mml:math id="M141" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> is therefore a net measure, incorporating macroalgal production, phytoplankton feedback, and air–sea carbon equilibration collectively <xref ref-type="bibr" rid="bib1.bibx11" id="paren.74"/>. It should be noted that we use annual average macroalgae NPP, to account for macroalgae seasonality, whereas the modelling study by <xref ref-type="bibr" rid="bib1.bibx11" id="altparen.75"/> uses prescribed maximal macroalgal NPP.</p>
      <p id="d2e3028">To calculate how much carbon is assumed to be durably stored, we calculate CDR efficiency using the proportion between harvested macroalgae biomass and the CDR flux <xref ref-type="bibr" rid="bib1.bibx80 bib1.bibx54" id="paren.76"/>:

            <disp-formula id="Ch1.E11" content-type="numbered"><label>11</label><mml:math id="M142" display="block"><mml:mtable class="split" rowspacing="0.2ex" displaystyle="true" columnalign="right left"><mml:mtr><mml:mtd><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow><mml:mo>=</mml:mo></mml:mrow></mml:mtd><mml:mtd><mml:mrow><mml:mspace width="0.25em" linebreak="nobreak"/><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msub><mml:mi mathvariant="normal">flux</mml:mi><mml:mi mathvariant="normal">macroalgae</mml:mi></mml:msub></mml:mrow><mml:mo>-</mml:mo><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msub><mml:mi mathvariant="normal">flux</mml:mi><mml:mi mathvariant="normal">control</mml:mi></mml:msub></mml:mrow></mml:mrow><mml:mrow class="chem"><mml:mi mathvariant="normal">Macroalgae</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">Harvest</mml:mi></mml:mrow></mml:mfrac></mml:mstyle></mml:mrow></mml:mtd></mml:mtr><mml:mtr><mml:mtd/><mml:mtd><mml:mrow><mml:mo>×</mml:mo><mml:mn mathvariant="normal">100</mml:mn><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="italic">%</mml:mi></mml:mrow></mml:mtd></mml:mtr></mml:mtable></mml:math></disp-formula></p>

      <fig id="F3" specific-use="star"><label>Figure 3</label><caption><p id="d2e3110">Global macroalgae cultivation and its influence on air–sea carbon flux across all experiments averaged between 2015–2024. <bold>(a)</bold> Absolute values of macroalgae primary production, phytoplankton primary production, the resulting enhancement in air–sea <inline-formula><mml:math id="M143" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> exchange (CDR flux), and harvest in <inline-formula><mml:math id="M144" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>. <bold>(b)</bold> CDR efficiency and harvest expressed as a proportion of macroalgae NPP (<inline-formula><mml:math id="M145" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>). Dark green bar shows the decline of phytoplankton net primary production.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f03.png"/>

        </fig>

</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
<sec id="Ch1.S3.SS1">
  <label>3.1</label><title>Seaweed Production and Harvesting</title>
      <p id="d2e3188">In the default simulation, macroalgae occupies 51.7 <inline-formula><mml:math id="M146" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>6</sup> <inline-formula><mml:math id="M148" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">km</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>, accounting for approximately 14.35 % of the global ocean surface. Macroalgae were primarily distributed across nitrogen-rich surface waters and major upwelling zones, including the Equatorial Pacific, North Pacific, North Atlantic, Southern Ocean, and coastal regions such as Chile, Argentina and Namibia (Fig. <xref ref-type="fig" rid="F2"/>a). The average annual macroalgae NPP of 111.1 <inline-formula><mml:math id="M149" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">g</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> (40.34 <inline-formula><mml:math id="M150" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>) was similar in magnitude to the average phytoplankton NPP from the control run (47.7 <inline-formula><mml:math id="M151" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>). The most dominant macroalgae species are <italic>Saccharina</italic> and <italic>Sargassum</italic> in high and low latitudes, respectively (see Fig. S1 in the Supplement), with the former contributing the most to macroalgae NPP (36.11 <inline-formula><mml:math id="M152" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>). The annual harvest yield that is sunk to the deep ocean reached 12.5 <inline-formula><mml:math id="M153" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> (135.59 <inline-formula><mml:math id="M154" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">tDW</mml:mi></mml:mrow></mml:math></inline-formula> (tonne Dry Weight) <inline-formula><mml:math id="M155" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">km</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>), 31.1 % of the macroalgae NPP, with a high proportion harvested in the North Atlantic, Southern Ocean, and along the coast of Chile (Figs. <xref ref-type="fig" rid="F2"/>b–d and <xref ref-type="fig" rid="F3"/>). The spatial patterns of macroalgae, NPP, and nutrients shown here are emergent outcomes of open-ocean biogeochemical conditions within a coarse-resolution global model, and should be interpreted as indicating potential cultivation areas and not as plausible regions for macroalgae farms. Many regions identified as productive for macroalgae would present extreme technological, logistical, and societal challenges.</p>
      <p id="d2e3372">Different cultivation protocols and model assumptions affect macroalgae outcomes. Varying the harvesting threshold changes NPP magnitude while preserving the general spatial pattern (Figs. S2 and S3 in the Supplement). Harvest 800 led to less harvested biomass (<inline-formula><mml:math id="M156" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>8.7 %), higher area of coverage (+6.28 %), and higher macroalgae NPP (+49 %). Harvest 200 decreases macroalgae coverage (<inline-formula><mml:math id="M157" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>5.43 %), reduced total macroalgae NPP (<inline-formula><mml:math id="M158" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>30 %), and modestly raised the harvested biomass (+13 %) due to more frequent harvest (see Fig. <xref ref-type="fig" rid="F3"/>). This resulted in a higher harvest efficiency (i.e., harvested biomass as a fraction of macroalgae NPP, see Fig. S3b in the Supplement) compared to the default simulation (Harvest 400).  Other protocols also affected macroalgae production, Fig. <xref ref-type="fig" rid="F3"/>. Extraction shows slight decreases of macroalgae NPP and area (by <inline-formula><mml:math id="M159" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1.37 %). Introducing Fe limitation (i.e. stopping Fe supplementation), dramatically suppressed macroalgae growth and harvest was almost entirely eliminated (<inline-formula><mml:math id="M160" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>99.2 %), and the cultivated area shrank by 47.95 %. Macroalgae NPP also declined by 72.8 %, especially in regions where Fe concentration is low, such as the Equatorial and North Pacific (see Fig. S2f in the Supplement). Higher non-harvesting loss also significantly reduced macroalgae area (44.6 %), NPP and harvest.</p>

      <fig id="F4" specific-use="star"><label>Figure 4</label><caption><p id="d2e3417"><inline-formula><mml:math id="M161" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux for <bold>(a)</bold> the control and <bold>(b)</bold> the difference between control and default macroalgae simulations. We also calculate the proportion of additional <inline-formula><mml:math id="M162" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux and macroalgae NPP, expressed as <inline-formula><mml:math id="M163" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f04.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS2">
  <label>3.2</label><title>mCDR Efficiency and <inline-formula><mml:math id="M164" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> Flux</title>
      <p id="d2e3484">Under default simulation conditions assuming no Fe limitation, global-scale macroalgae cultivation enhanced net air–sea <inline-formula><mml:math id="M165" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux by approximately 11.0 <inline-formula><mml:math id="M166" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> (equivalent to 40.3 <inline-formula><mml:math id="M167" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>; Fig. <xref ref-type="fig" rid="F3"/>), while without macroalgae, <inline-formula><mml:math id="M168" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux is only 1.4 <inline-formula><mml:math id="M169" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, indicating a substantial contribution to marine carbon dioxide removal (mCDR). The increase in <inline-formula><mml:math id="M170" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux under the default simulation is sufficient to limit warming to <inline-formula><mml:math id="M171" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>2 °C (between 0.7–3.6 <inline-formula><mml:math id="M172" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> <xref ref-type="bibr" rid="bib1.bibx23 bib1.bibx37" id="paren.77"/>). Although 31.1 % of primary production is being harvested and sunk, the <inline-formula><mml:math id="M173" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> (the proportion of additional <inline-formula><mml:math id="M174" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux to macroalgae NPP) was only 27.3 % but <inline-formula><mml:math id="M175" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> was 87.9 %. <inline-formula><mml:math id="M176" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> also varied regionally. High-efficiency zones (<inline-formula><mml:math id="M177" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M178" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 50 %) were simulated along the Southern Ocean, while low-efficiency areas (<inline-formula><mml:math id="M179" display="inline"><mml:mo lspace="0mm">&lt;</mml:mo></mml:math></inline-formula> 10 %) emerged in the central Equatorial Pacific, off the western coast of North Japan, and offshore South Africa (Fig. <xref ref-type="fig" rid="F4"/>c). Areas off southern coast of Chile, exhibited net outgassing due to surface DIC accumulation (Fig. <xref ref-type="fig" rid="F4"/>b). Since additional <inline-formula><mml:math id="M180" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux often occurs outside harvesting area (Fig. <xref ref-type="fig" rid="F2"/>c and Fig. <xref ref-type="fig" rid="F4"/>b), we do not show efficiency maps for <inline-formula><mml:math id="M181" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">Harv</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>.</p>
      <p id="d2e3737">The mCDR efficiency is sensitive to cultivation protocol. Harvest 200 slightly decreased the <inline-formula><mml:math id="M182" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux, improved the <inline-formula><mml:math id="M183" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> to 38.4 % (Macroalgae NPP of 27.95 <inline-formula><mml:math id="M184" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, CDR flux of 10.73 <inline-formula><mml:math id="M185" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>), but slightly lowers <inline-formula><mml:math id="M186" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> to 86.1 %, due to lower CDR flux. Whereas Harvest 800 reduced the CDR flux to 9.6 <inline-formula><mml:math id="M187" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> and lowered <inline-formula><mml:math id="M188" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> to 16.0 %, but slight increases <inline-formula><mml:math id="M189" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> to 89.4 %. In all threshold scenarios, the Equatorial Pacific consistently showed the lowest <inline-formula><mml:math id="M190" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, whilst modest improvements occurred in the Southern Ocean under the low-threshold scenario (Fig. S4b in the Supplement).</p>
      <p id="d2e3882">Further amendments to the cultivation strategies and modelling assumptions also influenced mCDR outcomes, Fig. <xref ref-type="fig" rid="F3"/>. The High Loss experiment increases macroalgae NPP to 60.18 <inline-formula><mml:math id="M191" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> because of the regenerated nutrients from non-harvesting loss, but reduced global <inline-formula><mml:math id="M192" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux into the ocean to 7.3 <inline-formula><mml:math id="M193" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, because there are less macroalgae harvested and sunk, which reduces <inline-formula><mml:math id="M194" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> to 16.0 %, but slightly increases <inline-formula><mml:math id="M195" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> to 88.4 %. Extraction increased the global <inline-formula><mml:math id="M196" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux by 0.13 <inline-formula><mml:math id="M197" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, because of less remineralisation in shallow waters, which induced <inline-formula><mml:math id="M198" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux. This protocol also has the highest <inline-formula><mml:math id="M199" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> of 91.7 % due to higher CDR flux. Simulating realistic Fe limitation constraints on macroalgae growth causes outgassing by contributing 0.2 <inline-formula><mml:math id="M200" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> to the atmosphere, indicating that scaled-up macroalgae cultivation without Fe fertilisation is not an effective mCDR technique (negative CDR efficiency, <inline-formula><mml:math id="M201" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1.6 %).</p>

<table-wrap id="T3" specific-use="star"><label>Table 3</label><caption><p id="d2e4051">Summary of absolute concentrations and percentage changes across macroalgae cultivation protocols, relative to the control simulation, for DIN and Fe at the surface and seafloor, Oxygen at the seafloor, as well as integrated phytoplankton NPP, surface phytoplankton, and zooplankton biomass.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="8">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:colspec colnum="4" colname="col4" align="left"/>
     <oasis:colspec colnum="5" colname="col5" align="left"/>
     <oasis:colspec colnum="6" colname="col6" align="left"/>
     <oasis:colspec colnum="7" colname="col7" align="left"/>
     <oasis:colspec colnum="8" colname="col8" align="left"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1"/>
         <oasis:entry colname="col2">control</oasis:entry>
         <oasis:entry colname="col3">Harvest 200</oasis:entry>
         <oasis:entry colname="col4">Default</oasis:entry>
         <oasis:entry colname="col5">Harvest 800</oasis:entry>
         <oasis:entry colname="col6">High Loss</oasis:entry>
         <oasis:entry colname="col7">Extraction</oasis:entry>
         <oasis:entry colname="col8">Fe limitation</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">Surface DIN (<inline-formula><mml:math id="M202" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">6.19</oasis:entry>
         <oasis:entry colname="col3">3.08</oasis:entry>
         <oasis:entry colname="col4">2.90</oasis:entry>
         <oasis:entry colname="col5">2.85</oasis:entry>
         <oasis:entry colname="col6">3.56</oasis:entry>
         <oasis:entry colname="col7">2.89</oasis:entry>
         <oasis:entry colname="col8">9.11</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M203" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M204" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>50.3</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M205" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>53.1</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M206" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>53.9</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M207" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>42.5</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M208" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>53.3</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M209" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>47.2</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Surface DFe (<inline-formula><mml:math id="M210" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">0.60 <inline-formula><mml:math id="M211" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col3">1.10 <inline-formula><mml:math id="M213" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col4">1.14 <inline-formula><mml:math id="M215" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col5">1.22 <inline-formula><mml:math id="M217" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col6">0.85 <inline-formula><mml:math id="M219" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col7">1.13 <inline-formula><mml:math id="M221" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col8">0.42 <inline-formula><mml:math id="M223" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M225" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M226" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>83.6</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M227" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>90.3</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M228" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>102.8</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M229" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>41.1</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M230" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>89.1</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M231" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>30.7</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Seafloor DIN (<inline-formula><mml:math id="M232" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">30.31</oasis:entry>
         <oasis:entry colname="col3">31.01</oasis:entry>
         <oasis:entry colname="col4">30.96</oasis:entry>
         <oasis:entry colname="col5">30.86</oasis:entry>
         <oasis:entry colname="col6">30.76</oasis:entry>
         <oasis:entry colname="col7">29.81</oasis:entry>
         <oasis:entry colname="col8">30.17</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M233" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M234" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>2.3</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M235" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>2.1</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M236" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>1.8</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M237" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>1.5</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M238" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1.7</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M239" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>0.5</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Seafloor DFe (<inline-formula><mml:math id="M240" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">0.55 <inline-formula><mml:math id="M241" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col3">1.61 <inline-formula><mml:math id="M243" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col4">1.59 <inline-formula><mml:math id="M245" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col5">1.42 <inline-formula><mml:math id="M247" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col6">0.56 <inline-formula><mml:math id="M249" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col7">0.55 <inline-formula><mml:math id="M251" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
         <oasis:entry colname="col8">0.55 <inline-formula><mml:math id="M253" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>−3</sup></oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M255" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M256" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>195.1</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M257" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>191.7</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M258" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>159.5</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M259" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>3.0</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M260" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>1.3</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M261" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>1.6</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Seafloor oxygen (<inline-formula><mml:math id="M262" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">205.5</oasis:entry>
         <oasis:entry colname="col3">162.6</oasis:entry>
         <oasis:entry colname="col4">163.7</oasis:entry>
         <oasis:entry colname="col5">168.8</oasis:entry>
         <oasis:entry colname="col6">176.6</oasis:entry>
         <oasis:entry colname="col7">205.8</oasis:entry>
         <oasis:entry colname="col8">205.1</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1"><inline-formula><mml:math id="M263" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M264" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>20.9</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M265" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>20.3</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M266" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>17.9</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M267" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>14.1</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M268" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula>0.2</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M269" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>0.2</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Phytoplankton (<inline-formula><mml:math id="M270" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">0.37</oasis:entry>
         <oasis:entry colname="col3">0.22</oasis:entry>
         <oasis:entry colname="col4">0.20</oasis:entry>
         <oasis:entry colname="col5">0.19</oasis:entry>
         <oasis:entry colname="col6">0.24</oasis:entry>
         <oasis:entry colname="col7">0.20</oasis:entry>
         <oasis:entry colname="col8">0.27</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M271" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M272" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>40.7</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M273" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>45.4</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M274" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>48.1</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M275" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>35.8</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M276" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>46.0</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M277" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>26.1</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Zooplankton (<inline-formula><mml:math id="M278" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mmol</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">0.26</oasis:entry>
         <oasis:entry colname="col3">0.13</oasis:entry>
         <oasis:entry colname="col4">0.14</oasis:entry>
         <oasis:entry colname="col5">0.16</oasis:entry>
         <oasis:entry colname="col6">0.17</oasis:entry>
         <oasis:entry colname="col7">0.13</oasis:entry>
         <oasis:entry colname="col8">0.19</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M279" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M280" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.6</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M281" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>47.2</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M282" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>36.8</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M283" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>34.5</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M284" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>48.1</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M285" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>27.8</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Phytoplankton NPP (<inline-formula><mml:math id="M286" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col2">47.7</oasis:entry>
         <oasis:entry colname="col3">26.0</oasis:entry>
         <oasis:entry colname="col4">24.0</oasis:entry>
         <oasis:entry colname="col5">23.0</oasis:entry>
         <oasis:entry colname="col6">29.5</oasis:entry>
         <oasis:entry colname="col7">23.8</oasis:entry>
         <oasis:entry colname="col8">31.7</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1"><inline-formula><mml:math id="M287" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula> (%)</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"><inline-formula><mml:math id="M288" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>45.5</oasis:entry>
         <oasis:entry colname="col4"><inline-formula><mml:math id="M289" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.8</oasis:entry>
         <oasis:entry colname="col5"><inline-formula><mml:math id="M290" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>51.9</oasis:entry>
         <oasis:entry colname="col6"><inline-formula><mml:math id="M291" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>38.2</oasis:entry>
         <oasis:entry colname="col7"><inline-formula><mml:math id="M292" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>50.2</oasis:entry>
         <oasis:entry colname="col8"><inline-formula><mml:math id="M293" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>33.6</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

</sec>
<sec id="Ch1.S3.SS3">
  <label>3.3</label><title>Impact of Macroalgae Cultivation on Ocean Biogeochemistry</title>
      <p id="d2e5286">Macroalgae cultivation had significant effects on surface nutrient concentrations and oxygen levels, primarily in areas of cultivation and harvesting, summarised in Table <xref ref-type="table" rid="T3"/>. In the default simulation, surface DIN declined by 53.1 %. This decline was co-located with high macroalgae productivity. Because of the assumption of Fe supplementation, surface DFe increases by 90.26 % from non-harvesting loss. Furthermore, to grow macroalgae ubiquitously in the default run, on average 1.4 <inline-formula><mml:math id="M294" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">Fe</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> was added to the ocean. The increased DFe occurs in areas occupied by macroalgae.  DIN and DFe accumulation was also found near the seafloor at deposition sites, reflecting remineralisation of sunken biomass. Due to high <inline-formula><mml:math id="M295" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">Fe</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio of macroalgae, the increase in seafloor DFe can reach up to 3<inline-formula><mml:math id="M296" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> the control simulation.</p>
      <p id="d2e5330">Altering the harvest threshold had a significant effect on surface nutrients (Table <xref ref-type="table" rid="T3"/> and Fig. S5a–h in the Supplement). Harvest 200 shows slightly weaker DIN drawdown (<inline-formula><mml:math id="M297" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>50.3 %), while Harvest 800 showed a slightly stronger DIN drawdown (<inline-formula><mml:math id="M298" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>53.9 %). These experiments also produce higher and lower surface DFe compared to the control simulation (83.56 % and 102.84 % more DFe for Harvest 200 and Harvest 800, respectively).  Among the alternative cultivation protocols, higher non-harvesting loss rate leads to less DIN decline (<inline-formula><mml:math id="M299" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>42.5 %) and lower DFe increase (41.05 %). In contrast, the Extraction experiment removed the benthic DIN enrichment entirely, and shows a slight benthic DFe increase, while having similar surface impacts to the default run. For the Fe limited experiment, the surface DIN and DFe increases and decreases by 47.2 % and <inline-formula><mml:math id="M300" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>30.7 %, respectively. At the seafloor, the DIN concentration declines by 0.48 % because of the lack of harvest but DFe increases by 1.64 % due to high <inline-formula><mml:math id="M301" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">Fe</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> ratio.</p>

      <fig id="F5" specific-use="star"><label>Figure 5</label><caption><p id="d2e5378">Changes in nutrients distribution in the surface between control and the default macroalgae simulation. <bold>(a)</bold>, <bold>(d)</bold>, and <bold>(g)</bold> show DIN, DFe, and seafloor oxygen from the control simulation, respectively, while from the default simulations are shown in <bold>(b)</bold>, <bold>(e)</bold>, and <bold>(h)</bold>. The difference between the two simulations are shown in <bold>(c)</bold>, <bold>(f)</bold>, and <bold>(i)</bold>.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f05.png"/>

        </fig>

      <p id="d2e5416">Oxygen concentrations were affected by both macroalgae growth and biomass disposal. At the seafloor, oxygen losses reached 20.3 % on average globally, but can reach 70 % decline in the harvesting and deposition areas (see Fig. <xref ref-type="fig" rid="F2"/>c). These areas can also become suboxic (Fig. <xref ref-type="fig" rid="F5"/>k), which covers 7.9 % of the seafloor, compared to 0.5 % in the control simulation.</p>
      <p id="d2e5423">Impact on oxygen concentration is closely linked with harvesting threshold (Table <xref ref-type="table" rid="T3"/> and Figs. S5o–u and S6 in the Supplement). Harvest 200 led to greater seafloor oxygen loss. Whereas Harvest 800 reduced oxygen depletion at the seafloor due to less deposition. Similarly, High Loss shows lower oxygen loss in the deep ocean, as less macroalgae is harvested (see Fig. <xref ref-type="fig" rid="F3"/>). In contrast, the Extraction experiment eliminated benthic oxygen loss. The inclusion of Fe limitation also shows very low benthic oxygen less compared to other experiments owing to suppressed harvest and sinking (Fig. S6f and g in the Supplement).</p>

      <fig id="F6" specific-use="star"><label>Figure 6</label><caption><p id="d2e5432">Plankton biomass and phytoplankton NPP and how it changes after macroalgae cultivation. The top, middle, and bottom rows show phytoplankton biomass, zooplankton biomass, and phytoplankton NPP. Control simulations are shown in <bold>(a)</bold>, <bold>(d)</bold>, and <bold>(g)</bold>, while <bold>(b)</bold>, <bold>(e)</bold>, and <bold>(h)</bold> shows the default macroalgae simulation. The difference between the two simulations are shown in <bold>(c)</bold>, <bold>(f)</bold>, and <bold>(i)</bold>.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f06.png"/>

        </fig>

      <p id="d2e5469">Large-scale macroalgae cultivation substantially altered surface nutrient availability and light conditions, leading to pronounced changes in the phytoplankton NPP. Note that these changes reflect competition for nutrients and light as represented in the model and provide an indicative estimate of the potential biogeochemical response, rather than a detailed prediction of realised ecosystem behaviour.  In the default simulation, phytoplankton NPP fell by 49.78 % (<inline-formula><mml:math id="M302" display="inline"><mml:mo lspace="0mm">-</mml:mo></mml:math></inline-formula>24.81 <inline-formula><mml:math id="M303" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>), accompanied by declines of in phytoplankton biomass (45.40 %) and zooplankton biomass (47.16 %), with the largest reductions in nutrient-rich and upwelling regions (Fig. <xref ref-type="fig" rid="F6"/>b, e, and h). There are also areas where both phytoplankton and zooplankton concentrations are increasing, such as the polar Southern Ocean (Fig. <xref ref-type="fig" rid="F6"/>d, e, and f), which may occur due to the increase in DFe concentration (Fig. <xref ref-type="fig" rid="F5"/>d–f). Adjusting harvesting threshold modified the severity but not the direction of these changes (Fig. 7c, d, j, and k in the Supplement): Harvest 200 slightly alleviated phytoplankton biomass and NPP losses but intensified zooplankton declines, while Harvest 800 worsened phytoplankton and NPP reductions but not as much zooplankton decline as the default run.</p>
      <p id="d2e5505">Other modelling considerations, such as imposing higher non-harvest loss would reduce less phytoplankton biomass, NPP, and zooplankton biomass. Extraction had little effect on plankton dynamics compared to the default, suggesting that surface processes dominate short-term responses. Imposing Fe limitation sharply constrained macroalgal growth and partly restored surface nutrients, especially DIN, yet phytoplankton and zooplankton showed net decline (Table <xref ref-type="table" rid="T3"/>), although regions such as the Indian, Atlantic, and parts of North Pacific did exhibit increased phytoplankton and zooplankton concentrations (Fig. S7 in the Supplement). These results underscore that ecosystem-wide impacts persist even when macroalgae productivity is minimal.</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
      <p id="d2e5519">Our study extends previous large-scale macroalgal cultivation modelling by focusing on cultivation protocols, such as harvesting and extracting biomass, physiological constraints, and biogeochemical feedbacks using the NEMO–MEDUSA ocean biogeochemistry model.  This approach allowed us to assess not only the theoretical mCDR potential of large-scale macroalgae cultivation, but also the ecological trade-offs associated with different deployment strategies and assumptions.</p>
      <p id="d2e5522">Our default simulation is broadly consistent with previous observations and models. Simulated macroalgae NPP (111.1 <inline-formula><mml:math id="M304" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">g</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>) falls within the observed global range of 91–522 <inline-formula><mml:math id="M305" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">g</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> (<xref ref-type="bibr" rid="bib1.bibx33 bib1.bibx25 bib1.bibx56" id="altparen.78"/>. The simulated harvesting yield of 135.6 <inline-formula><mml:math id="M306" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">tDW</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">km</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, is slightly smaller than previous modelling study <xref ref-type="bibr" rid="bib1.bibx79" id="paren.79"/>, yet remains within the observational range <xref ref-type="bibr" rid="bib1.bibx60" id="paren.80"/>. Macroalgae biomass hotspots align with previously reported patterns from models <xref ref-type="bibr" rid="bib1.bibx4 bib1.bibx79" id="paren.81"><named-content content-type="pre">e.g.</named-content></xref> and the estimated suitable ocean area for macroalgae cultivation, (51.7 <inline-formula><mml:math id="M307" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 10<sup>6</sup> <inline-formula><mml:math id="M309" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">km</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> of the ocean) agrees with previous estimates using <inline-formula><mml:math id="M310" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">N</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">P</mml:mi></mml:mrow></mml:math></inline-formula> ratios and Earth system models <xref ref-type="bibr" rid="bib1.bibx28 bib1.bibx79" id="paren.82"/>. These agreements show that our simulation captures the key first-order biogeochemical drivers relevant to large-scale macroalgae cultivation.</p>
      <p id="d2e5658">From the default simulation, global macroalgae cultivation has the potential to increase <inline-formula><mml:math id="M311" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake by 11.0 <inline-formula><mml:math id="M312" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> exceeding the amount required to align with 2 <inline-formula><mml:math id="M313" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">°</mml:mi><mml:mi mathvariant="normal">C</mml:mi></mml:mrow></mml:math></inline-formula> climate pathway (0.7–3.6 <inline-formula><mml:math id="M314" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> by mid-century <xref ref-type="bibr" rid="bib1.bibx24 bib1.bibx37" id="altparen.83"/>). This magnitude is higher than the earlier global cultivation study of <xref ref-type="bibr" rid="bib1.bibx79" id="text.84"/> (3.63 <inline-formula><mml:math id="M315" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>), which may be due to no phosphate limitation and Fe supplementation in our simulation. Relative to macroalgal NPP the increase in <inline-formula><mml:math id="M316" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake remains modest (<inline-formula><mml:math id="M317" display="inline"><mml:mo lspace="0mm">∼</mml:mo></mml:math></inline-formula> 27 %, see Fig. <xref ref-type="fig" rid="F3"/>), and is lower than a modelling cultivation study within EEZs (58 %; <xref ref-type="bibr" rid="bib1.bibx11" id="altparen.85"/>). It is important to note that the 27 % CDR efficiency already accounts for the displacement of phytoplankton productivity. <inline-formula><mml:math id="M318" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> measures the fraction of macroalgal NPP that results in additional <inline-formula><mml:math id="M319" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake by the ocean, relative to a control simulation without macroalgae (Eq. <xref ref-type="disp-formula" rid="Ch1.E10"/>, following <xref ref-type="bibr" rid="bib1.bibx11" id="altparen.86"/>). The 50 % suppression of phytoplankton NPP reported in Sect. 3.3 is therefore not an additional loss to be subtracted from the 27 % figure, it is already embedded within it. In other words, macroalgae cultivation under the default scenario replaces roughly half of the natural biological pump while delivering only 27 % additional CDR, highlighting the modest net climate benefit relative to the scale of ecosystem disruption. This reflects the strong nutrient constraints on biotic mCDR.</p>
      <p id="d2e5805">Although up to 87.9 % of the <inline-formula><mml:math id="M320" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake in our simulations is associated with harvested and sunk biomass, only 31 % of macroalgal production is actually harvested, indicating that most fixed carbon is lost than durably stored. While our idealised experiments allow macroalgae cultivation over large oceanic areas, such configuration should be viewed as upper-bound scenarios. When cultivation is restricted to shelf-sea area (depth <inline-formula><mml:math id="M321" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 200m, Fig. S8 in the Supplement), simulated <inline-formula><mml:math id="M322" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake is reduced to 0.81 <inline-formula><mml:math id="M323" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> using default simulation condition, higher than that in previous EEZ study (0.21 <inline-formula><mml:math id="M324" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, <xref ref-type="bibr" rid="bib1.bibx11" id="altparen.87"/>). This also suggests that constraining the spatial extent of cultivation affects the magnitude, rather than direction of biogeochemical response. As with other biotic CDR approaches, the net removal ultimately depends on both biogeochemical constraints and equilibration timescales governing air–sea <inline-formula><mml:math id="M325" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> exchange.</p>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>Biogeochemical Impacts</title>
      <p id="d2e5900">The enhanced <inline-formula><mml:math id="M326" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake in the default scenario was accompanied by major biogeochemical side-effects. Macroalgae cultivation reduced global phytoplankton NPP and zooplankton biomass by almost half the control model, which is caused by DIN depletion in surface waters. This decline exceeded 50 %, consistent with nutrient robbing seen in previous modelling studies <xref ref-type="bibr" rid="bib1.bibx1 bib1.bibx15 bib1.bibx79" id="paren.88"/>. However it is also important to note that the macroalgae submodule includes only a simple loss term and does not explicitly resolve grazing or other higher trophic food-web feedbacks (e.g. in <xref ref-type="bibr" rid="bib1.bibx79 bib1.bibx80" id="altparen.89"/> zooplankton grazing is explicitly represented). Additionally, MEDUSA only represents two phytoplankton functional types, which do not fully capture the diversity of nutrient acquisition strategies and ecological responses present in natural systems. As a result, the simulated reduction in phytoplankton NPP and zooplankton biomass reported here may represent an overestimation, because explicit herbivory on macroalgae could recycle nutrients and reduce the impact on phytoplankton NPP, as illustrated in the High Loss experiment (see Fig. 3). A more diverse representation of phytoplankton functional types could also alter the magnitude and spatial pattern of NPP responses. This makes the simulated phytoplankton response in this study a first-order biogeochemical estimate of potential competition effects.</p>
      <p id="d2e5920">Sustaining large-scale global macroalgal cultivation required Fe supplementation, which drove a <inline-formula><mml:math id="M327" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 90 % increase in surface DFe and up to 3<inline-formula><mml:math id="M328" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> increase at the seafloor due to the remineralisation of biomass deposition. Despite the DFe enrichment phytoplankton NPP biomass declined by nearly half (see Fig. <xref ref-type="fig" rid="F3"/>, Table <xref ref-type="table" rid="T3"/>) due to macronutrient robbing by macroalgae. An ocean Fe fertilisation modelling study also showed large-scale Fe fertilisation caused enhanced consumption of major nutrients in surface waters, and also reduces nutrients availability for lower latitudes, which offsets overall NPP <xref ref-type="bibr" rid="bib1.bibx70" id="paren.90"/>. This suggests that mCDR approaches can risk redistributing ecological pressure, while also suppressing additional carbon removal from phytoplankton.</p>
      <p id="d2e5944">Deep-sea biogeochemistry will also be altered due to macroalgal deposition. In the default simulation, deep-sea oxygen losses can reach suboxic levels in deposition zones (Fig. <xref ref-type="fig" rid="F5"/>j-l) which has been a concern in previous studies <xref ref-type="bibr" rid="bib1.bibx79 bib1.bibx48 bib1.bibx19" id="paren.91"/>. Severe deoxygenation would favour smaller species, reduces large predators and bioturbation, and triggers faunal emergence or habitat avoidance, especially in the coastal benthos <xref ref-type="bibr" rid="bib1.bibx48 bib1.bibx47" id="paren.92"/>, which may alter natural carbon sequestration in the deep sea environment. Deposition also increases DIN at the seafloor (Table <xref ref-type="table" rid="T3"/>) and the introduction of nutrients to the oligotrophic seafloor may also alter benthic species interactions <xref ref-type="bibr" rid="bib1.bibx48" id="paren.93"/>.  We note that these results are not a detailed characterisation of benthic ecosystem impacts. MEDUSA's representation is intentionally simple, and a full assessment of seafloor community responses, including benthic respiration dynamics and faunal impacts, would require dedicated benthic ecological models. However, these results may be used to identify large-scale signals and pattens that would motivate future benthic studies.</p>
</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>Model Sensitivities </title>
      <p id="d2e5968">To examine how these impacts depend on cultivation and modelling design, we tested alternative harvesting threshold, extraction, loss rates, and nutrient limitations. Harvest timing and threshold can affect the optimal yield and capital expenditure <xref ref-type="bibr" rid="bib1.bibx7" id="paren.94"/>. Lowering the harvest threshold improved CDR flux and efficiencies, both <inline-formula><mml:math id="M329" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mi mathvariant="normal">effHarv</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M330" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CDR</mml:mi><mml:mrow><mml:mi mathvariant="normal">eff</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">NPP</mml:mi></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, while slightly reducing biogeochemical disruption, and with similar CDR flux as the default run. A field study has showed that a more frequent harvest without re-seeding, would increase yield per meter growth line and reduced cultivation cost <xref ref-type="bibr" rid="bib1.bibx7 bib1.bibx8" id="paren.95"/>. However, it also resulted in a greater decline in zooplankton biomass than phytoplankton biomass (Table <xref ref-type="table" rid="T3"/>) indicating potential downstream effects of macroalgae cultivation on higher trophic levels and, potentially, fisheries <xref ref-type="bibr" rid="bib1.bibx30 bib1.bibx44" id="paren.96"/>. Higher thresholds allow more macroalgae biomass to accumulate ahead of harvesting, leading to an increase in macroalgal NPP. However, as more of this NPP is lost to non-harvesting losses in the model, the associated CDR efficiency is actually lower compared to other harvest experiments (Fig. <xref ref-type="fig" rid="F3"/>). Further, by increasing the quantity of harvested material transferred to depth, higher thresholds additionally cause greater seafloor oxygen depletion, as well as a larger reduction in the NPP of phytoplankton (Table <xref ref-type="table" rid="T3"/>, Fig. <xref ref-type="fig" rid="F5"/>k).</p>
      <p id="d2e6016">Other modelling and cultivation protocols can also significantly affect macroalgae production and CDR capacity. When non-harvesting loss is doubled, the macroalgae NPP is reduced by 2.20 <inline-formula><mml:math id="M331" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> and its CDR flux, by 3.7 <inline-formula><mml:math id="M332" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>, making it less efficient (Fig.  <xref ref-type="fig" rid="F3"/>a). High-loss experiments also lose less surface DIN compared to the default run because of higher remineralisation near the surface. In farmed macroalgae, non-harvesting loss due to falloff and frond erosion can reach more than 10 % of its growth rate <xref ref-type="bibr" rid="bib1.bibx84" id="paren.97"/>. This simulation explores how <inline-formula><mml:math id="M333" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux and ocean biogeochemistry are affected when non-harvesting loss consume more of macroalgal biomass. When harvested biomass is extracted rather than sunk to the deep ocean, CDR efficiencies become slightly higher than the default simulation because of the lack of carbon leaking when depositing macroalgae at shallower depths. This simulation also shows minimal impacts at the seafloor due to the absence of deposition (Table <xref ref-type="table" rid="T3"/>).</p>
      <p id="d2e6078">The CDR capacity of macroalgae is lower when Fe is not supplemented, since most areas of the open ocean will not be suitable for growing macroalgae <xref ref-type="bibr" rid="bib1.bibx57" id="paren.98"/>. When Fe limitation is implemented, macroalgae NPP collapsed with a 73.7 % decrease in magnitude, aligning with recent work by <xref ref-type="bibr" rid="bib1.bibx12" id="text.99"/>, and simulate a net increase in <inline-formula><mml:math id="M334" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux to the atmosphere of 0.2 <inline-formula><mml:math id="M335" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>. However, due to the positive bias in Fe concentration in MEDUSA, compared to the major decline of macroalgae harvest in the Southern Ocean that is simulated in <xref ref-type="bibr" rid="bib1.bibx12" id="text.100"/>, the decline in this study is not as dramatic. The Fe limitation experiment also simulated 30.7 % less DFe concentrations in surface waters than the control model (Table <xref ref-type="table" rid="T3"/>), further depressing phytoplankton NPP in the Southern Ocean, Equatorial Pacific, and subpolar North Atlantic. This simulation emphasises that large-scale macroalgal CDR depends critically on Fe supplementation and the enhanced <inline-formula><mml:math id="M336" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake simulated under Fe-replete conditions depends on nutrient subsidies.</p>

      <fig id="F7" specific-use="star"><label>Figure 7</label><caption><p id="d2e6138">Large variation in <inline-formula><mml:math id="M337" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake within macroalgae cultivation areas. <bold>(a)</bold> Model grid cells are sorted by their quantitative contributions to additional <inline-formula><mml:math id="M338" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake and then the associated cultivation area and <inline-formula><mml:math id="M339" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake are accumulated. <bold>(b)</bold> Geographical distribution of cultivation area which contributes the most to additional <inline-formula><mml:math id="M340" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>. Deep red regions are those that contribute most to <inline-formula><mml:math id="M341" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake, while deep blue regions contribute least.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/3735/2026/bg-23-3735-2026-f07.png"/>

        </fig>

</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>Challenges for Implementation and Uncertainties</title>
      <p id="d2e6217">In carbon dioxide removal, additionality denotes that an mCDR strategy must demonstrate more <inline-formula><mml:math id="M342" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake relative to what would occur without it which will require model assessments due to the global scale of impacts and potential feedbacks between reservoirs <xref ref-type="bibr" rid="bib1.bibx6" id="paren.101"/>. Assessing the additionality of macroalgae-based mCDR can be challenging because its <inline-formula><mml:math id="M343" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake enhancement is tightly coupled to the redistribution of nutrients and phytoplankton NPP loss within the ocean. In our simulation large-scale cultivation suppressed phytoplankton NPP both within and beyond cultivation areas (Figs. <xref ref-type="fig" rid="F6"/> and S7 in the Supplement), and this can complicate the measurement of net carbon removed <xref ref-type="bibr" rid="bib1.bibx6" id="paren.102"/>. In terms of macroalgal cultivation, even with Fe supplementation, the increase of macroalgae biomass is at the expense of natural biological carbon export <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx11 bib1.bibx79" id="paren.103"/>. According to the framework of <xref ref-type="bibr" rid="bib1.bibx6" id="text.104"/> macroalgae cultivation may show even smaller additionality since the carbon cost for setting up offshore macroalgae farm can be more than half of the potential CDR <xref ref-type="bibr" rid="bib1.bibx21" id="paren.105"/>.</p>
      <p id="d2e6260">Compared to abiotic approaches, such as coastal ocean alkalinity enhancement (OAE), our large-scale macroalgal cultivation uptake is less spatially concentrated. Using the same modelling framework, <xref ref-type="bibr" rid="bib1.bibx58" id="text.106"/> reported that 13.8 % of the shelf accounts for 50 % of the total extra <inline-formula><mml:math id="M344" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux, with warmer areas contributing higher <inline-formula><mml:math id="M345" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake, indicating that CDR flux is dominated by a few high efficiency regions. In contrast, our default macroalgae simulation indicates that 28.9 % of the cultivation area is required to reach the same proportion (Fig. <xref ref-type="fig" rid="F7"/>a). The regions driving this uptake coincide with areas of high macroalgae NPP and CDR flux (Figs. <xref ref-type="fig" rid="F2"/>b and <xref ref-type="fig" rid="F4"/>b) particularly subpolar Southern Ocean and coast of Chile, as shown in Fig. <xref ref-type="fig" rid="F7"/>b, reflecting a more widespread uptake pattern. This difference arises from the tight nutrient constrain in macroalgae-cultivation, while in OAE, additional <inline-formula><mml:math id="M346" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake is dependent on temperature, that enhances dissolution rate and the geography of the shelf.  Furthermore, the rate of air–sea <inline-formula><mml:math id="M347" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> gas transfer, parameterised following <xref ref-type="bibr" rid="bib1.bibx78" id="text.107"/> as a function of local wind speed, introduces a natural lag between biological carbon drawdown and atmospheric <inline-formula><mml:math id="M348" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake. DIC deficits that persist beyond a single growing season continue to drive <inline-formula><mml:math id="M349" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> uptake as surface waters are re-exposed to the atmosphere through seasonal mixed layer dynamics and circulation. This explains the regional variation in CDR efficiency shown in Fig. 4c. In the Equatorial Pacific, relatively high macroalgal NPP is associated with lower CDR efficiency. In contrast, the deeper mixed layer in the Southern Ocean keeps low-DIC water in contact with the atmosphere for longer, resulting in higher efficiency. Although regions where high cumulative <inline-formula><mml:math id="M350" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> share, CDR flux, and macroalgae NPP are located in the Southern Ocean, this area may not be suitable. Large-scale offshore macroalgae deployment would require extensive floating infrastructure, long-distance operations, and resilience to extreme wave and weather conditions <xref ref-type="bibr" rid="bib1.bibx39 bib1.bibx45" id="paren.108"/>. Such engineering and logistical considerations are not represented in the present model and remain outside the scope of this study.</p>
      <p id="d2e6359">Macroalgal extraction and product substitution can offer co-benefits such as substituting high-emission products with seaweed-based alternative. However, life-cycle analyses show that processing macroalgal products can offset much of the theoretical climate advantage <xref ref-type="bibr" rid="bib1.bibx39" id="paren.109"/>. A recent economic model estimates that product substitution yields a net profit of <inline-formula><mml:math id="M351" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> USD 50 <inline-formula><mml:math id="M352" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">per</mml:mi><mml:mspace linebreak="nobreak" width="0.25em"/><mml:mi mathvariant="normal">t</mml:mi><mml:mspace linebreak="nobreak" width="0.25em"/><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> avoided, compared to a cost of USD 480 <inline-formula><mml:math id="M353" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">per</mml:mi><mml:mspace linebreak="nobreak" width="0.25em"/><mml:mi mathvariant="normal">t</mml:mi><mml:mspace width="0.25em" linebreak="nobreak"/><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> for deep-ocean deposition <xref ref-type="bibr" rid="bib1.bibx24" id="paren.110"/>, and product substitution could cut regional emissions by up to 13 %, whereas direct sequestration contributes relatively little <xref ref-type="bibr" rid="bib1.bibx16" id="paren.111"/>. Together these studies highlight the importance of robust monitoring, reporting, and verification frameworks that can verify additionality and durable sequestration.</p>
      <p id="d2e6413">Natural climate solutions, such as saltmarshes, mangroves, and seagrass ecosystems or coastal macroalgal cultivation, can deliver both carbon sequestration and a range of co-benefits, including coastal protection, nutrient retention, and biodiversity support <xref ref-type="bibr" rid="bib1.bibx49 bib1.bibx59" id="paren.112"/>. These systems provide relatively direct and more verifiable carbon storage through sediment burial, while also helping mitigate eutrophication. Such coastal approaches may offer more tractable mCDR deployment, even though at smaller scale. These findings reinforce that no single approach is likely to deliver the required scale of carbon removal.</p>
      <p id="d2e6420">Several model limitations highlight the need for cautious interpretation of the quantitative estimates. Although MEDUSA captures the patterns and concentration of macronutrients fairly well <xref ref-type="bibr" rid="bib1.bibx83" id="paren.113"/>, the model tends to overestimates DFe concentrations in regions known for persistent Fe limitation <xref ref-type="bibr" rid="bib1.bibx69" id="paren.114"/>. Additionally, our macroalgae module lacks variable stoichiometry <xref ref-type="bibr" rid="bib1.bibx4" id="paren.115"/>, DOC release <xref ref-type="bibr" rid="bib1.bibx56" id="paren.116"/>, and explicit macroalgal grazing and erosion <xref ref-type="bibr" rid="bib1.bibx79 bib1.bibx80 bib1.bibx4" id="paren.117"/>. The model also does not represent calcifying epibionts that commonly colonise sargassum, and their calcification can offset CDR <xref ref-type="bibr" rid="bib1.bibx5" id="paren.118"/>. Furthermore, MEDUSA does not resolve coccolithophores phytoplankton types, meaning that potential reduction in calcite ballasting due to the reduction in phytoplankton concentration, are not captured. These omissions would act to further reduce CDR efficiency.</p>
      <p id="d2e6442">The absence of riverine nutrient inputs and coastal-shelf processes also prevents realistic simulation of Sargassum dynamics <xref ref-type="bibr" rid="bib1.bibx76" id="paren.119"/> and likely underestimates the productive potential of warm-water cultivation in regions such as Southeast and East Asia where terrestrial nutrient subsidies support high macroalgal biomass in nearshore systems. Although the four taxa represented in this study span a range of thermal niches, cold and warm water taxa are associated with different cultivation systems, whereby warm water species are usually farmed using single step methods through vegetative propagation and low-cost fixed or floating raft systems in nearshore environments <xref ref-type="bibr" rid="bib1.bibx10" id="paren.120"/> while cold water species require multi-step propagation from spores, hatchery-based seeding, and deployment on longlines system <xref ref-type="bibr" rid="bib1.bibx13 bib1.bibx8" id="paren.121"/>. These differences are not captured in this idealised framework and may mean that the modelled growth potential presented here translates differently to real-world aquaculture output depending on the region in question.</p>
      <p id="d2e6454">Although MEDUSA includes deoxygenation and acidification processes, the simplicity of its benthic ecosystem model cannot represent the impacts of such mechanisms on the seafloor ecosystems. While some processes may be partially represented within MEDUSA’s remineralisation scheme, a targeted evaluation of acidification dynamics was beyond the scope of this study. Our results should be interpreted as a first order approximation of the biogeochemical consequences of large-scale macroalgae cultivation, and that representing these additional processes in future model developments will be important in estimating macroalgal CDR potential</p>
      <p id="d2e6457">Taken together, our results reinforce growing concerns that large-scale macroalgae cultivation may offer limited net climate benefits when ecological externalities and feedbacks are considered <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx29 bib1.bibx34" id="paren.122"/>. Despite its theoretical potential, macroalgal CDR appears constrained by nutrient bottlenecks, competition with natural carbon pumps, and low CDR capacity compared to macroalgal NPP. These findings point to the need for regionally tailored deployment, modelling and observation monitoring strategies, and explicit additionality accounting if macroalgal cultivation is to play a role in future climate mitigation portfolios.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusions</title>
      <p id="d2e6473">This study provides a global-scale assessment of macroalgae cultivation within a coupled ocean biogeochemical model. Our simulations highlight the potential for carbon sequestration, while also revealing significant trade-offs, as well as sensitivities to model assumptions and different cultivation strategies. Given the uncertainties and model limitations, the results presented here should be interpreted as a first-order approximation of the biogeochemical consequences of large-scale macroalgae cultivation. Overall, our findings suggest that solely relying on large-scale macroalgae cultivation is unlikely to provide a viable mCDR solution, and that mitigation efforts will likely need to be distributed across different mCDR strategies, such as OAE and coastal ecosystem restoration. The key findings of this study are summarised as follows: <list list-type="bullet"><list-item>
      <p id="d2e6478">This study presents a global scale implementation of a macroalgae cultivation module within a coupled OBGC model, and evaluates the carbon removal potential and biogeochemical impacts of this activity under multiple cultivation strategies and assumptions.</p></list-item><list-item>
      <p id="d2e6482">We find that while macroalgae cultivation can enhance the net air–sea <inline-formula><mml:math id="M354" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux into the ocean by up to 11.0 <inline-formula><mml:math id="M355" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> when supplemented with Fe, the associated CDR efficiency is a modest 27.3 % relative to the macroalgae productivity driving it.</p></list-item><list-item>
      <p id="d2e6517">However, these fluxes come at the cost of substantial alterations to ocean biogeochemistry and ecosystems. In the default case, macroalgal growth reduces surface DIN concentrations by 53.1 %, resulting in suppression of phytoplankton primary production by 49.8 %, and global reductions in phytoplankton and zooplankton biomass, by 45.4 % and 47.2 %, respectively.</p></list-item><list-item>
      <p id="d2e6521">When macroalgae are harvested and deposited, seafloor oxygen will decline globally by 20.3 % and cause suboxic conditions in 7.9 % of the seafloor, compared to 0.5 % in the control model, especially within the deposition zones.</p></list-item><list-item>
      <p id="d2e6525">The magnitude and spatial footprint of these impacts are strongly modulated by cultivation protocol, with more frequent harvesting improving CDR efficiency but intensifying the simulated oxygen depletion at depth caused by macroalgae biomass dumping.</p></list-item><list-item>
      <p id="d2e6529">Exploration of other cultivation protocols and model assumptions, finds significant changes in macroalgae production and <inline-formula><mml:math id="M356" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> flux. For example, Fe limitation of macroalgae growth actually causes outgassing by contributing 0.2 <inline-formula><mml:math id="M357" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">Pg</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">C</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">yr</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula> to the atmosphere, indicating that scaled-up macroalgae cultivation without Fe fertilisation is not an effective mCDR technique.</p></list-item></list></p>
</sec>

      
      </body>
    <back><notes notes-type="codedataavailability"><title>Code and data availability</title>

      <p id="d2e6567">The FORTRAN code for MEDUSA-Macroalgae, instructions for its use, and model output is archived in the Zenodo repository at <ext-link xlink:href="https://doi.org/10.5281/zenodo.20154696" ext-link-type="DOI">10.5281/zenodo.20154696</ext-link> <xref ref-type="bibr" rid="bib1.bibx3" id="paren.123"/>.</p>
  </notes><app-group>
        <supplementary-material position="anchor"><p id="d2e6576">The supplement related to this article is available online at <inline-supplementary-material xlink:href="https://doi.org/10.5194/bg-23-3735-2026-supplement" xlink:title="pdf">https://doi.org/10.5194/bg-23-3735-2026-supplement</inline-supplementary-material>.</p></supplementary-material>
        </app-group><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d2e6585">EKP, AY, and CAB acquired funding. PA, AY, and JP developed the macroalgae module. PA and AY conceptualised the study. PA performed the simulations, model output analysis, writing the first draft, and produced the figures. All authors contributed to ideas, participated in discussions of the results, and editing of the manuscript.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d2e6591">The contact author has declared that none of the authors has any competing interests.</p>
  </notes><notes notes-type="disclaimer"><title>Disclaimer</title>

      <p id="d2e6597">Publisher's note: Copernicus Publications remains neutral with regard to jurisdictional claims made in the text, published maps, institutional affiliations, or any other geographical representation in this paper. The authors bear the ultimate responsibility for providing appropriate place names. Views expressed in the text are those of the authors and do not necessarily reflect the views of the publisher.</p>
  </notes><ack><title>Acknowledgements</title><p id="d2e6603">The authors are grateful to the National Oceanography Centre's High-Performance Computing (HPC) system, which facilitated model simulation and analysis. All authors are funded by UK Research and Innovation (UKRI) under National Capability Science Multi-Centre (NCSMC) funding for Atlantic Climate and Environment Strategic Science (AtlantiS; NE/Y005589/1). Additionally, Prima Anugerahanti, Andrew Yool, Ekaterina Popova, and Julien Palmiéri are funded under NCSMC funding for Future Impacts, Risks, and Mitigation Actions in a changing Earth system project (TerraFIRMA LTSM; NE/W004895/1). The Python code used to generate figures in this manuscript was written with the help of ChatGPT.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d2e6608">This research has been supported by the National Environment Research Council (NERC) (grant nos. NE/W004895/1 and NE/Y005589/1).The article processing charges for this open-access publication were covered by the National Oceanography Centre.</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d2e6619">This paper was edited by Koji Suzuki and reviewed by Gregory Nishihara, John Gallagher, and one anonymous referee.</p>
  </notes><ref-list>
    <title>References</title>

      <ref id="bib1.bibx1"><label>Aldridge et al.(2021)Aldridge, Mooney, Dabrowski, and Capuzzo</label><mixed-citation>Aldridge, J., Mooney, K., Dabrowski, T., and Capuzzo, E.: Modelling effects of seaweed aquaculture on phytoplankton and mussel production. Application to Strangford Lough (Northern Ireland), Aquaculture, 536, 736400, <ext-link xlink:href="https://doi.org/10.1016/j.aquaculture.2021.736400" ext-link-type="DOI">10.1016/j.aquaculture.2021.736400</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx2"><label>Alevizos and Barillé(2023)</label><mixed-citation>Alevizos, E. and Barillé, L.: Global ocean spatial suitability for macroalgae offshore cultivation and sinking, Front. Mar. Sci., 10, 1320642, <ext-link xlink:href="https://doi.org/10.3389/fmars.2023.1320642" ext-link-type="DOI">10.3389/fmars.2023.1320642</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx3"><label>Anugerahanti et al.(2026)Anugerahanti, Palmieri, and Yool</label><mixed-citation>Anugerahanti, P., Palmieri, J., and Yool, A.: The Impact of Large-Scale Macroalgae Cultivation and Harvesting Strategies on the Marine Carbon Dioxide Removal Efficacy and Marine Biogeochemistry – Supplementary materials, Zenodo [code and data set], <ext-link xlink:href="https://doi.org/10.5281/zenodo.20154696" ext-link-type="DOI">10.5281/zenodo.20154696</ext-link>, 2026.</mixed-citation></ref>
      <ref id="bib1.bibx4"><label>Arzeno-Soltero et al.(2023)Arzeno-Soltero, Saenz, Frieder, Long, DeAngelo, Davis, and Davis</label><mixed-citation>Arzeno-Soltero, I. B., Saenz, B. T., Frieder, C. A., Long, M. C., DeAngelo, J., Davis, S. J., and Davis, K. A.: Large global variations in the carbon dioxide removal potential of seaweed farming due to biophysical constraints, Commun. Earth Environ., 4, 185, <ext-link xlink:href="https://doi.org/10.1038/s43247-023-00833-2" ext-link-type="DOI">10.1038/s43247-023-00833-2</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx5"><label>Bach et al.(2021)Bach, Tamsitt, Gower, Hurd, Raven, and Boyd</label><mixed-citation>Bach, L. T., Tamsitt, V., Gower, J., Hurd, C. L., Raven, J. A., and Boyd, P. W.: Testing the climate intervention potential of ocean afforestation using the Great Atlantic Sargassum Belt, Nat. Commun., 12, 2556, <ext-link xlink:href="https://doi.org/10.1038/s41467-021-22837-2" ext-link-type="DOI">10.1038/s41467-021-22837-2</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx6"><label>Bach et al.(2024)Bach, Vaughan, Law, and Williamson</label><mixed-citation>Bach, L. T., Vaughan, N. E., Law, C. S., and Williamson, P.: Implementation of marine CO2 removal for climate mitigation: The challenges of additionality, predictability, and governability, Elem. Sci. Anth., 12, 00034, <ext-link xlink:href="https://doi.org/10.1525/elementa.2023.00034" ext-link-type="DOI">10.1525/elementa.2023.00034</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bibx7"><label>Bak et al.(2018)Bak, Mols-Mortensen, and Gregersen</label><mixed-citation>Bak, U. G., Mols-Mortensen, A., and Gregersen, O.: Production method and cost of commercial-scale offshore cultivation of kelp in the Faroe Islands using multiple partial harvesting, Algal Res., 33, 36–47, <ext-link xlink:href="https://doi.org/10.1016/j.algal.2018.05.001" ext-link-type="DOI">10.1016/j.algal.2018.05.001</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bibx8"><label>Bak et al.(2020)Bak, Gregersen, and Infante</label><mixed-citation>Bak, U. G., Gregersen, O., and Infante, J.: Technical challenges for offshore cultivation of kelp species: lessons learned and future directions, Bot. Mar., 63, 341–353, <ext-link xlink:href="https://doi.org/10.1515/bot-2019-0005" ext-link-type="DOI">10.1515/bot-2019-0005</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bibx9"><label>Barrett et al.(2022)Barrett, Theuerkauf, Rose, Alleway, Bricker, Parker, Petrolia, and Jones</label><mixed-citation>Barrett, L. T., Theuerkauf, S. J., Rose, J. M., Alleway, H. K., Bricker, S. B., Parker, M., Petrolia, D. R., and Jones, R. C.: Sustainable growth of non-fed aquaculture can generate valuable ecosystem benefits, Ecosyst. Serv., 53, 101396, <ext-link xlink:href="https://doi.org/10.1016/j.ecoser.2021.101396" ext-link-type="DOI">10.1016/j.ecoser.2021.101396</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx10"><label>Behera et al.(2022)Behera, Vadodariya, Veeragurunathan, Sigamani, Moovendhan, Srinivasan, Kolandhasamy, and Ingle</label><mixed-citation>Behera, D. P., Vadodariya, V., Veeragurunathan, V., Sigamani, S., Moovendhan, M., Srinivasan, R., Kolandhasamy, P., and Ingle, K. N.: Seaweeds cultivation methods and their role in climate mitigation and environmental cleanup, Total Environment Research Themes, 3–4, 100016, <ext-link xlink:href="https://doi.org/10.1016/j.totert.2022.100016" ext-link-type="DOI">10.1016/j.totert.2022.100016</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx11"><label>Berger et al.(2023)Berger, Kwiatkowski, Ho, and Bopp</label><mixed-citation>Berger, M., Kwiatkowski, L., Ho, D. T., and Bopp, L.: Ocean dynamics and biological feedbacks limit the potential of macroalgae carbon dioxide removal, Environ. Res. Lett., 18, 024039, <ext-link xlink:href="https://doi.org/10.1088/1748-9326/acb06e" ext-link-type="DOI">10.1088/1748-9326/acb06e</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx12"><label>Berger et al.(2025)Berger, Kwiatkowski, Bopp, and Ho</label><mixed-citation>Berger, M., Kwiatkowski, L., Bopp, L., and Ho, D. T.: Efficacy of seaweed-based carbon dioxide removal reduced by iron limitation and nutrient competition with phytoplankton, CDRxiv [Preprints], <ext-link xlink:href="https://doi.org/10.70212/cdrxiv.2025385.v1" ext-link-type="DOI">10.70212/cdrxiv.2025385.v1</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx13"><label>Boderskov et al.(2023)Boderskov, Rasmussen, and Bruhn</label><mixed-citation>Boderskov, T., Rasmussen, M. B., and Bruhn, A.: Upscaling cultivation of <italic>Saccharina latissima</italic> on net or line systems; comparing biomass yields and nutrient extraction potentials, Front. Mar. Sci., 10, 992179, <ext-link xlink:href="https://doi.org/10.3389/fmars.2023.992179" ext-link-type="DOI">10.3389/fmars.2023.992179</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx14"><label>Boyd and Vivian(2019)</label><mixed-citation> Boyd, P. and Vivian, C. M. G., e.: GESAMP “High level review of a wide range of proposed marine geoengineering techniques”, Tech. rep., Joint Group of Experts on the Scientific Aspects of Marine Environmental Protection, 2019.</mixed-citation></ref>
      <ref id="bib1.bibx15"><label>Boyd et al.(2022)Boyd, Bach, Hurd, Paine, Raven, and Tamsitt</label><mixed-citation>Boyd, P. W., Bach, L. T., Hurd, C. L., Paine, E., Raven, J. A., and Tamsitt, V.: Potential negative effects of ocean afforestation on offshore ecosystems, Nat. Ecol. Evol., 6, 675–683, <ext-link xlink:href="https://doi.org/10.1038/s41559-022-01722-1" ext-link-type="DOI">10.1038/s41559-022-01722-1</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx16"><label>Bullen et al.(2024)Bullen, Driscoll, Burt, Stephens, Hessing-Lewis, and Gregr</label><mixed-citation>Bullen, C. D., Driscoll, J., Burt, J., Stephens, T., Hessing-Lewis, M., and Gregr, E. J.: The potential climate benefits of seaweed farming in temperate waters, Sci. Rep., 14, 15021, <ext-link xlink:href="https://doi.org/10.1038/s41598-024-65408-3" ext-link-type="DOI">10.1038/s41598-024-65408-3</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bibx17"><label>Chen et al.(2020)Chen, Li, Zhang, He, Shi, Jiao, and Zhang</label><mixed-citation>Chen, J., Li, H., Zhang, Z., He, C., Shi, Q., Jiao, N., and Zhang, Y.: DOC dynamics and bacterial community succession during long-term degradation of <italic>Ulva prolifera</italic> and their implications for the legacy effect of green tides on refractory DOC pool in seawater, Water Res., 185, 116268, <ext-link xlink:href="https://doi.org/10.1016/j.watres.2020.116268" ext-link-type="DOI">10.1016/j.watres.2020.116268</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bibx18"><label>Choi et al.(2025)Choi, Lee, Jung, Lee, Han, Hyeon, and Choi</label><mixed-citation>Choi, D., Lee, H.-G., Jung, Y.-H., Lee, D.-W., Han, J., Hyeon, J.-Y., and Choi, Y.-U.: Pilot-Scale Cultivation of Seaweed (<italic>Undaria pinnatifida</italic>) Along an Offshore Wind Farm in Southwestern Korea, J. Mar. Sci. Eng., 13, <ext-link xlink:href="https://doi.org/10.3390/jmse13050882" ext-link-type="DOI">10.3390/jmse13050882</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx19"><label>Chopin et al.(2024)Chopin, Costa-Pierce, Troell, Hurd, Costello, Backman, Buschmann, Cuhel, Duarte, Gröndahl, Heasman, Haroun, Johansen, Jueterbock, Lench, Lindell, Pavia, Ricart, Sundell, and Yarish</label><mixed-citation>Chopin, T., Costa-Pierce, B. A., Troell, M., Hurd, C. L., Costello, M. J., Backman, S., Buschmann, A. H., Cuhel, R., Duarte, C. M., Gröndahl, F., Heasman, K., Haroun, R. J., Johansen, J., Jueterbock, A., Lench, M., Lindell, S., Pavia, H., Ricart, A. M., Sundell, K. S., and Yarish, C.: Deep-ocean seaweed dumping for carbon sequestration: Questionable, risky, and not the best use of valuable biomass, One Earth, 7, S2590332224000356, <ext-link xlink:href="https://doi.org/10.1016/j.oneear.2024.01.013" ext-link-type="DOI">10.1016/j.oneear.2024.01.013</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bibx20"><label>Chung et al.(2011)Chung, Beardall, Mehta, Sahoo, and Stojkovic</label><mixed-citation>Chung, I. K., Beardall, J., Mehta, S., Sahoo, D., and Stojkovic, S.: Using marine macroalgae for carbon sequestration: a critical appraisal, J. Appl. Phycol., 23, 877–886, <ext-link xlink:href="https://doi.org/10.1007/s10811-010-9604-9" ext-link-type="DOI">10.1007/s10811-010-9604-9</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bibx21"><label>Coleman et al.(2022)Coleman, Dewhurst, Fredriksson, St. Gelais, Cole, MacNicoll, Laufer, and Brady</label><mixed-citation>Coleman, S., Dewhurst, T., Fredriksson, D. W., St. Gelais, A. T., Cole, K. L., MacNicoll, M., Laufer, E., and Brady, D. C.: Quantifying baseline costs and cataloging potential optimization strategies for kelp aquaculture carbon dioxide removal, Front. Marine Sci., 9, <ext-link xlink:href="https://doi.org/10.3389/fmars.2022.966304" ext-link-type="DOI">10.3389/fmars.2022.966304</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx22"><label>Corrigan et al.(2022)Corrigan, Brown, Ashton, Smale, and Tyler</label><mixed-citation>Corrigan, S., Brown, A. R., Ashton, I. G. C., Smale, D. A., and Tyler, C. R.: Quantifying habitat provisioning at macroalgal cultivation sites, Rev. Aquacult., 14, 1671–1694, <ext-link xlink:href="https://doi.org/10.1111/raq.12669" ext-link-type="DOI">10.1111/raq.12669</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx23"><label>DeAngelo et al.(2021)DeAngelo, Azevedo, Bistline, Clarke, Luderer, Byers, and Davis</label><mixed-citation>DeAngelo, J., Azevedo, I., Bistline, J., Clarke, L., Luderer, G., Byers, E., and Davis, S. J.: Energy systems in scenarios at net-zero CO2 emissions, Nat. Commun., 12, 6096, <ext-link xlink:href="https://doi.org/10.1038/s41467-021-26356-y" ext-link-type="DOI">10.1038/s41467-021-26356-y</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx24"><label>DeAngelo et al.(2022)DeAngelo, Saenz, Arzeno-Soltero, Frieder, Long, Hamman, Davis, and Davis</label><mixed-citation>DeAngelo, J., Saenz, B. T., Arzeno-Soltero, I. B., Frieder, C. A., Long, M. C., Hamman, J., Davis, K. A., and Davis, S. J.: Economic and biophysical limits to seaweed farming for climate change mitigation, Nat. Plants, 9, 45–57, <ext-link xlink:href="https://doi.org/10.1038/s41477-022-01305-9" ext-link-type="DOI">10.1038/s41477-022-01305-9</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx25"><label>Duarte et al.(2017)Duarte, Wu, Xiao, Bruhn, and Krause-Jensen</label><mixed-citation>Duarte, C. M., Wu, J., Xiao, X., Bruhn, A., and Krause-Jensen, D.: Can Seaweed Farming Play a Role in Climate Change Mitigation and Adaptation?, Front. Mar. Sci., 4, <ext-link xlink:href="https://doi.org/10.3389/fmars.2017.00100" ext-link-type="DOI">10.3389/fmars.2017.00100</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bibx26"><label>Duarte et al.(2021)Duarte, Bruhn, and Krause-Jensen</label><mixed-citation>Duarte, C. M., Bruhn, A., and Krause-Jensen, D.: A seaweed aquaculture imperative to meet global sustainability targets, Nat. Sustain., 5, 185–193, <ext-link xlink:href="https://doi.org/10.1038/s41893-021-00773-9" ext-link-type="DOI">10.1038/s41893-021-00773-9</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx27"><label>Enríquez et al.(1996)Enríquez, Duarte, Sand-Jensen, and Nielsen</label><mixed-citation>Enríquez, S., Duarte, C. M., Sand-Jensen, K., and Nielsen, S. L.: Broad-scale comparison of photosynthetic rates across phototrophic organisms, Oecologia, 108, 197–206, <ext-link xlink:href="https://doi.org/10.1007/BF00334642" ext-link-type="DOI">10.1007/BF00334642</ext-link>, 1996.</mixed-citation></ref>
      <ref id="bib1.bibx28"><label>Froehlich et al.(2019)Froehlich, Afflerbach, Frazier, and Halpern</label><mixed-citation>Froehlich, H. E., Afflerbach, J. C., Frazier, M., and Halpern, B. S.: Blue Growth Potential to Mitigate Climate Change through Seaweed Offsetting, Curr. Biol., 29, 3087–3093.e3, <ext-link xlink:href="https://doi.org/10.1016/j.cub.2019.07.041" ext-link-type="DOI">10.1016/j.cub.2019.07.041</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bibx29"><label>Gallagher et al.(2022)Gallagher, Shelamoff, and Layton</label><mixed-citation>Gallagher, J. B., Shelamoff, V., and Layton, C.: Seaweed ecosystems may not mitigate CO2 emissions, ICES J. Mar. Sci., 79, 585–592, <ext-link xlink:href="https://doi.org/10.1093/icesjms/fsac011" ext-link-type="DOI">10.1093/icesjms/fsac011</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx30"><label>Guibourd de Luzinais et al.(2023)Guibourd de Luzinais, du Pontavice, Reygondeau, Barrier, Blanchard, Bornarel, Büchner, Cheung, Eddy, Everett, Guiet, Harrison, Maury, Novaglio, Petrik, Steenbeek, Tittensor, and Gascuel</label><mixed-citation>Guibourd de Luzinais, V., du Pontavice, H., Reygondeau, G., Barrier, N., Blanchard, J. L., Bornarel, V., Büchner, M., Cheung, W. W. L., Eddy, T. D., Everett, J. D., Guiet, J., Harrison, C. S., Maury, O., Novaglio, C., Petrik, C. M., Steenbeek, J., Tittensor, D. P., and Gascuel, D.: Trophic amplification: A model intercomparison of climate driven changes in marine food webs, PLOS ONE, 18, 1–23, <ext-link xlink:href="https://doi.org/10.1371/journal.pone.0287570" ext-link-type="DOI">10.1371/journal.pone.0287570</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx31"><label>Hayashi et al.(2017)Hayashi, Reis, Dos Santos, Castelar, Robledo, De Vega, Msuya, Eswaran, Yasir, Ali, and Hurtado</label><mixed-citation>Hayashi, L., Reis, R. P., Dos Santos, A. A., Castelar, B., Robledo, D., De Vega, G. B., Msuya, F. E., Eswaran, K., Yasir, S. M., Ali, M. K. M., and Hurtado, A. Q.: The Cultivation of Kappaphycus and Eucheuma in Tropical and Sub-Tropical Waters, in: Tropical Seaweed Farming Trends, Problems and Opportunities, edited by: Hurtado, A. Q., Critchley, A. T., and Neish, I. C., Springer International Publishing, Cham, pp. 55–90, <ext-link xlink:href="https://doi.org/10.1007/978-3-319-63498-2_4" ext-link-type="DOI">10.1007/978-3-319-63498-2_4</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bibx32"><label>Hersbach et al.(2020)Hersbach, Bell, Berrisford, Hirahara, Horányi, Muñoz-Sabater, Nicolas, Peubey, Radu, Schepers, Simmons, Soci, Abdalla, Abellan, Balsamo, Bechtold, Biavati, Bidlot, Bonavita, De Chiara, Dahlgren, Dee, Diamantakis, Dragani, Flemming, Forbes, Fuentes, Geer, Haimberger, Healy, Hogan, Hólm, Janisková, Keeley, Laloyaux, Lopez, Lupu, Radnoti, de Rosnay, Rozum, Vamborg, Villaume, and Thépaut</label><mixed-citation>Hersbach, H., Bell, B., Berrisford, P., Hirahara, S., Horányi, A., Muñoz-Sabater, J., Nicolas, J., Peubey, C., Radu, R., Schepers, D., Simmons, A., Soci, C., Abdalla, S., Abellan, X., Balsamo, G., Bechtold, P., Biavati, G., Bidlot, J., Bonavita, M., De Chiara, G., Dahlgren, P., Dee, D., Diamantakis, M., Dragani, R., Flemming, J., Forbes, R., Fuentes, M., Geer, A., Haimberger, L., Healy, S., Hogan, R. J., Hólm, E., Janisková, M., Keeley, S., Laloyaux, P., Lopez, P., Lupu, C., Radnoti, G., de Rosnay, P., Rozum, I., Vamborg, F., Villaume, S., and Thépaut, J.-N.: The ERA5 global reanalysis, Q. J. Roy. Meteor. Soc., 146, 1999–2049, <ext-link xlink:href="https://doi.org/10.1002/qj.3803" ext-link-type="DOI">10.1002/qj.3803</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bibx33"><label>Hurd et al.(2014)Hurd, Harrison, Bischof, and Lobban</label><mixed-citation>Hurd, C. L., Harrison, P. J., Bischof, K., and Lobban, C. S.: Seaweed Ecology and Physiology, Cambridge University Press, Cambridge, ISBN 9781139192637, <ext-link xlink:href="https://doi.org/10.1017/CBO9781139192637" ext-link-type="DOI">10.1017/CBO9781139192637</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bibx34"><label>Hurd et al.(2022)Hurd, Law, Bach, Britton, Hovenden, Paine, Raven, Tamsitt, and Boyd</label><mixed-citation>Hurd, C. L., Law, C. S., Bach, L. T., Britton, D., Hovenden, M., Paine, E. R., Raven, J. A., Tamsitt, V., and Boyd, P. W.: Forensic carbon accounting: Assessing the role of seaweeds for carbon sequestration, J. Phycol., 58, 347–363, <ext-link xlink:href="https://doi.org/10.1111/jpy.13249" ext-link-type="DOI">10.1111/jpy.13249</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx35"><label>Hurd et al.(2023)Hurd, Gattuso, and Boyd</label><mixed-citation>Hurd, C. L., Gattuso, J., and Boyd, P. W.: Air-sea carbon dioxide equilibrium: Will it be possible to use seaweeds for carbon removal offsets?, J. Phycol., 60, jpy.13405, <ext-link xlink:href="https://doi.org/10.1111/jpy.13405" ext-link-type="DOI">10.1111/jpy.13405</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx36"><label>Hwang et al.(2018)Hwang, Liu, Lee, Ha, and Park</label><mixed-citation>Hwang, E. K., Liu, F., Lee, K. H., Ha, D. S., and Park, C. S.: Comparison of the cultivation performance between Korean (Sugwawon No. 301) and Chinese strains (Huangguan No. 1) of kelp <italic>Saccharina japonica</italic> in an aquaculture farm in Korea, Algae, 33, 101–108, <ext-link xlink:href="https://doi.org/10.4490/algae.2018.33.2.4" ext-link-type="DOI">10.4490/algae.2018.33.2.4</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bibx37"><label>IPCC(2022)</label><mixed-citation>IPCC: Summary for Policymakers, Cambridge University Press, 3–48, <ext-link xlink:href="https://doi.org/10.1017/9781009157896.001" ext-link-type="DOI">10.1017/9781009157896.001</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx38"><label>Jiang et al.(2013)Jiang, Fang, Mao, Han, and Wang</label><mixed-citation>Jiang, Z., Fang, J., Mao, Y., Han, T., and Wang, G.: Influence of Seaweed Aquaculture on Marine Inorganic Carbon Dynamics and Sea-air <inline-formula><mml:math id="M358" display="inline"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">CO</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula> Flux, J. World Aquacult. Soc., 44, 133–140, <ext-link xlink:href="https://doi.org/10.1111/jwas.12000" ext-link-type="DOI">10.1111/jwas.12000</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bibx39"><label>Jiao et al.(2025)Jiao, Feng, Li, and Tian</label><mixed-citation>Jiao, T., Feng, E. Y., Li, Y., and Tian, Y.: Carbon dioxide removal dilemma of macroalgae products: Evidence from carbon footprint and profitability, J. Clean. Prod., 492, 144870, <ext-link xlink:href="https://doi.org/10.1016/j.jclepro.2025.144870" ext-link-type="DOI">10.1016/j.jclepro.2025.144870</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx40"><label>Jones et al.(2014)Jones, Ito, Takano, and Hsu</label><mixed-citation>Jones, D. C., Ito, T., Takano, Y., and Hsu, W.-C.: Spatial and seasonal variability of the air-sea equilibration timescale of carbon dioxide, Global Biogeochem. Cy., 28, 1163–1178, <ext-link xlink:href="https://doi.org/10.1002/2014GB004813" ext-link-type="DOI">10.1002/2014GB004813</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bibx41"><label>Kennedy and Blain(2025)</label><mixed-citation>Kennedy, J. R. and Blain, C. O.: A systematic review of marine macroalgal degradation: Toward a better understanding of macroalgal carbon sequestration potential, J. Phycol., 61, 399–432, <ext-link xlink:href="https://doi.org/10.1111/jpy.70031" ext-link-type="DOI">10.1111/jpy.70031</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx42"><label>Koesling et al.(2021)Koesling, Kvadsheim, Halfdanarson, Emblemsvåg, and Rebours</label><mixed-citation>Koesling, M., Kvadsheim, N. P., Halfdanarson, J., Emblemsvåg, J., and Rebours, C.: Environmental impacts of protein-production from farmed seaweed: Comparison of possible scenarios in Norway, J. Clean. Prod., 307, 127301, <ext-link xlink:href="https://doi.org/10.1016/j.jclepro.2021.127301" ext-link-type="DOI">10.1016/j.jclepro.2021.127301</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx43"><label>Krause-Jensen and Duarte(2016)</label><mixed-citation>Krause-Jensen, D. and Duarte, C. M.: Substantial role of macroalgae in marine carbon sequestration, Nat. Geosci., 9, 737–742, <ext-link xlink:href="https://doi.org/10.1038/ngeo2790" ext-link-type="DOI">10.1038/ngeo2790</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bibx44"><label>Kwiatkowski et al.(2019)Kwiatkowski, Aumont, and Bopp</label><mixed-citation>Kwiatkowski, L., Aumont, O., and Bopp, L.: Consistent trophic amplification of marine biomass declines under climate change, Glob. Change Biol., 25, 218–229, <ext-link xlink:href="https://doi.org/10.1111/gcb.14468" ext-link-type="DOI">10.1111/gcb.14468</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bibx45"><label>Kwon et al.(2024)Kwon, Hawkins, Zaimes, Infante, Kite-Powell, Stekoll, Roberson, Zotter, Augyte, Rocheleau, and Sims</label><mixed-citation>Kwon, H., Hawkins, T. R., Zaimes, G. G., Infante, J., Kite-Powell, H. L., Stekoll, M. S., Roberson, L., Zotter, B., Augyte, S., Rocheleau, G., and Sims, N.: Life-cycle analysis of offshore macroalgae production systems in the United States, Algal Rese., 82, 103654, <ext-link xlink:href="https://doi.org/10.1016/j.algal.2024.103654" ext-link-type="DOI">10.1016/j.algal.2024.103654</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bibx46"><label>Lauvset et al.(2016)Lauvset, Key, Olsen, van Heuven, Velo, Lin, Schirnick, Kozyr, Tanhua, Hoppema, Jutterström, Steinfeldt, Jeansson, Ishii, Perez, Suzuki, and Watelet</label><mixed-citation>Lauvset, S. K., Key, R. M., Olsen, A., van Heuven, S., Velo, A., Lin, X., Schirnick, C., Kozyr, A., Tanhua, T., Hoppema, M., Jutterström, S., Steinfeldt, R., Jeansson, E., Ishii, M., Perez, F. F., Suzuki, T., and Watelet, S.: A new global interior ocean mapped climatology: the 1° <inline-formula><mml:math id="M359" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 1° GLODAP version 2, Earth Syst. Sci. Data, 8, 325–340, <ext-link xlink:href="https://doi.org/10.5194/essd-8-325-2016" ext-link-type="DOI">10.5194/essd-8-325-2016</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bibx47"><label>Levin et al.(2009)Levin, Ekau, Gooday, Jorissen, Middelburg, Naqvi, Neira, Rabalais, and Zhang</label><mixed-citation>Levin, L. A., Ekau, W., Gooday, A. J., Jorissen, F., Middelburg, J. J., Naqvi, S. W. A., Neira, C., Rabalais, N. N., and Zhang, J.: Effects of natural and human-induced hypoxia on coastal benthos, Biogeosciences, 6, 2063–2098, <ext-link xlink:href="https://doi.org/10.5194/bg-6-2063-2009" ext-link-type="DOI">10.5194/bg-6-2063-2009</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bibx48"><label>Levin et al.(2023)Levin, Alfaro-Lucas, Colaço, Cordes, Craik, Danovaro, Hoving, Ingels, Mestre, Seabrook, Thurber, Vivian, and Yasuhara</label><mixed-citation>Levin, L. A., Alfaro-Lucas, J. M., Colaço, A., Cordes, E. E., Craik, N., Danovaro, R., Hoving, H.-J., Ingels, J., Mestre, N. C., Seabrook, S., Thurber, A. R., Vivian, C., and Yasuhara, M.: Deep-sea impacts of climate interventions, Science, 379, 978–981, <ext-link xlink:href="https://doi.org/10.1126/science.ade7521" ext-link-type="DOI">10.1126/science.ade7521</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx49"><label>Macreadie et al.(2021)Macreadie, Costa, Atwood, Friess, Kelleway, Kennedy, Lovelock, Serrano, and Duarte</label><mixed-citation>Macreadie, P. I., Costa, M. D. P., Atwood, T. B., Friess, D. A., Kelleway, J. J., Kennedy, H., Lovelock, C. E., Serrano, O., and Duarte, C. M.: Blue carbon as a natural climate solution, Nat. Rev. Earth Environ., 2, 826–839, <ext-link xlink:href="https://doi.org/10.1038/s43017-021-00224-1" ext-link-type="DOI">10.1038/s43017-021-00224-1</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx50"><label>Madec(2016)</label><mixed-citation> Madec, G.: NEMO ocean engine, Tech. rep., publication Title: Note du Pole de modelisation Issue: 27, Institut Pierre-Simon Laplace No 27, ISSN No 1288-1619, 2016.</mixed-citation></ref>
      <ref id="bib1.bibx51"><label>Magcanta-Mortos et al.(2025)Magcanta-Mortos, Tahiluddin, Mortos, Aspe, Aaron-Amper, Leopardas, and Uy</label><mixed-citation>Magcanta-Mortos, M. L. M., Tahiluddin, A. B., Mortos, J. M. R., Aspe, N. M., Aaron-Amper, J., Leopardas, V. E., and Uy, W. H.: Sargassum hatchery-based cultivation and mariculture techniques in the Philippines: Practices, challenges, and future prospects, Aquat. Bot., 201, 103925, <ext-link xlink:href="https://doi.org/10.1016/j.aquabot.2025.103925" ext-link-type="DOI">10.1016/j.aquabot.2025.103925</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx52"><label>Naylor et al.(2021)Naylor, Hardy, Buschmann, Bush, Cao, Klinger, Little, Lubchenco, Shumway, and Troell</label><mixed-citation>Naylor, R. L., Hardy, R. W., Buschmann, A. H., Bush, S. R., Cao, L., Klinger, D. H., Little, D. C., Lubchenco, J., Shumway, S. E., and Troell, M.: A 20-year retrospective review of global aquaculture, Nature, 591, 551–563, <ext-link xlink:href="https://doi.org/10.1038/s41586-021-03308-6" ext-link-type="DOI">10.1038/s41586-021-03308-6</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx53"><label>N‘Yeurt et al.(2012)N‘Yeurt, Chynoweth, Capron, Stewart, and Hasan</label><mixed-citation>N‘Yeurt, A. D. R., Chynoweth, D. P., Capron, M. E., Stewart, J. R., and Hasan, M. A.: Negative carbon via Ocean Afforestation, Process Saf. Environ., 90, 467–474, <ext-link xlink:href="https://doi.org/10.1016/j.psep.2012.10.008" ext-link-type="DOI">10.1016/j.psep.2012.10.008</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bibx54"><label>Ocean Visions and Monterey Bay Aquarium Research Institute(2022)</label><mixed-citation>Ocean Visions and Monterey Bay Aquarium Research Institute: Answering Critical Questions About Sinking Macroalgae for Carbon Dioxide Removal: A Research Framework to Investigate Sequestration Efficacy and Environmental Impacts, Tech. rep., Ocean Visions, Monterey Bay Aquarium Research Institute, <uri>https://oceanvisions.org/wp-content/uploads/2022/10/Ocean-Visions-Sinking-Seaweed-Report_FINAL.pdf</uri> (last access: 29 October 2025), 2022.</mixed-citation></ref>
      <ref id="bib1.bibx55"><label>Orr and Epitalon(2015)</label><mixed-citation>Orr, J. C. and Epitalon, J.-M.: Improved routines to model the ocean carbonate system: mocsy 2.0, Geosci. Model Dev., 8, 485–499, <ext-link xlink:href="https://doi.org/10.5194/gmd-8-485-2015" ext-link-type="DOI">10.5194/gmd-8-485-2015</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bibx56"><label>Paine et al.(2021)Paine, Schmid, Boyd, Diaz-Pulido, and Hurd</label><mixed-citation>Paine, E. R., Schmid, M., Boyd, P. W., Diaz-Pulido, G., and Hurd, C. L.: Rate and fate of dissolved organic carbon release by seaweeds: A missing link in the coastal ocean carbon cycle, J. Phycol., 57, 1375–1391, <ext-link xlink:href="https://doi.org/10.1111/jpy.13198" ext-link-type="DOI">10.1111/jpy.13198</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx57"><label>Paine et al.(2023)Paine, Boyd, Strzepek, Ellwood, Brewer, Diaz-Pulido, Schmid, and Hurd</label><mixed-citation>Paine, E. R., Boyd, P. W., Strzepek, R. F., Ellwood, M., Brewer, E. A., Diaz-Pulido, G., Schmid, M., and Hurd, C. L.: Iron limitation of kelp growth may prevent ocean afforestation, Commun. Biol., 6, 607, <ext-link xlink:href="https://doi.org/10.1038/s42003-023-04962-4" ext-link-type="DOI">10.1038/s42003-023-04962-4</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx58"><label>Palmiéri and Yool(2024)</label><mixed-citation>Palmiéri, J. and Yool, A.: Global-Scale Evaluation of Coastal Ocean Alkalinity Enhancement in a Fully Coupled Earth System Model, Earths Future, 12, e2023EF004018, <ext-link xlink:href="https://doi.org/10.1029/2023EF004018" ext-link-type="DOI">10.1029/2023EF004018</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bibx59"><label>Pessarrodona et al.(2023)Pessarrodona, Franco-Santos, Wright, Vanderklift, Howard, Pidgeon, Wernberg, and Filbee-Dexter</label><mixed-citation>Pessarrodona, A., Franco-Santos, R. M., Wright, L. S., Vanderklift, M. A., Howard, J., Pidgeon, E., Wernberg, T., and Filbee-Dexter, K.: Carbon sequestration and climate change mitigation using macroalgae: a state of knowledge review, Biol. Rev., 98, 1945–1971, <ext-link xlink:href="https://doi.org/10.1111/brv.12990" ext-link-type="DOI">10.1111/brv.12990</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx60"><label>Peteiro et al.(2014)Peteiro, Sánchez, Dueñas-Liaño, and Martínez</label><mixed-citation>Peteiro, C., Sánchez, N., Dueñas-Liaño, C., and Martínez, B.: Open-sea cultivation by transplanting young fronds of the kelp Saccharina latissima, J. Appl. Phycol., 26, 519–528, <ext-link xlink:href="https://doi.org/10.1007/s10811-013-0096-2" ext-link-type="DOI">10.1007/s10811-013-0096-2</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bibx61"><label>Reagan et al.(2024)Reagan, Boyer, García, Locarnini, Baranova, Bouchard, Cross, Mishonov, Paver, Seidov, Wang, and Dukhovskoy</label><mixed-citation>Reagan, J. R., Boyer, T. P., García, H. E., Locarnini, R. A., Baranova, O. K., Bouchard, C., Cross, S. L., Mishonov, A. V., Paver, C. R., Seidov, D., Wang, Z., and Dukhovskoy, D.: World Ocean Atlas 2023, NCEI [data set], <ext-link xlink:href="https://doi.org/10.25921/va26-hv25" ext-link-type="DOI">10.25921/va26-hv25</ext-link>,  2024.</mixed-citation></ref>
      <ref id="bib1.bibx62"><label>Roberts et al.(2015)Roberts, Paul, Dworjanyn, Bird, and De Nys</label><mixed-citation>Roberts, D. A., Paul, N. A., Dworjanyn, S. A., Bird, M. I., and De Nys, R.: Biochar from commercially cultivated seaweed for soil amelioration, Sci. Rep., 5, 9665, <ext-link xlink:href="https://doi.org/10.1038/srep09665" ext-link-type="DOI">10.1038/srep09665</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bibx63"><label>Roque et al.(2019)Roque, Salwen, Kinley, and Kebreab</label><mixed-citation> Roque, B. M., Salwen, J. K., Kinley, R., and Kebreab, E.: Inclusion of Asparagopsis armata in lactating dairy cows' diet reduces enteric methane emission by over 50 percent, J. Clean. Prod., 234, 132–138, 2019.</mixed-citation></ref>
      <ref id="bib1.bibx64"><label>Ross et al.(2022)Ross, Tarbuck, and Macreadie</label><mixed-citation>Ross, F., Tarbuck, P., and Macreadie, P. I.: Seaweed afforestation at large-scales exclusively for carbon sequestration: Critical assessment of risks, viability and the state of knowledge, Front. Mar. Sci., 9, 1015612, <ext-link xlink:href="https://doi.org/10.3389/fmars.2022.1015612" ext-link-type="DOI">10.3389/fmars.2022.1015612</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx65"><label>Sato et al.(2023)Sato, Fujiwara, and Endo</label><mixed-citation>Sato, Y., Fujiwara, T., and Endo, H.: Density regulation of aquaculture production and its effects on commercial profit and quality as food in the cosmopolitan edible seaweed Undaria pinnatifida, Front. Mar. Sci., 10, <ext-link xlink:href="https://doi.org/10.3389/fmars.2023.1085054" ext-link-type="DOI">10.3389/fmars.2023.1085054</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx66"><label>Sato et al.(2025)Sato, Saito, Inomata, Tanaka, and Nishihara</label><mixed-citation>Sato, Y., Saito, D., Inomata, E., Tanaka, A., and Nishihara, G. N.: Carbon and nitrogen contents depends on macroalgal species, their tissue section, and development stage, Phycol. Res., 74, <ext-link xlink:href="https://doi.org/10.1111/pre.70007" ext-link-type="DOI">10.1111/pre.70007</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx67"><label>Sharma et al.(2018)Sharma, Neves, Funderud, Mydland, Øverland, and Horn</label><mixed-citation>Sharma, S., Neves, L., Funderud, J., Mydland, L. T., Øverland, M., and Horn, S. J.: Seasonal and depth variations in the chemical composition of cultivated <italic>Saccharina latissima</italic>, Algal Rese., 32, 107–112, <ext-link xlink:href="https://doi.org/10.1016/j.algal.2018.03.012" ext-link-type="DOI">10.1016/j.algal.2018.03.012</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bibx68"><label>Sheppard et al.(2023)Sheppard, Hurd, Britton, Reed, and Bach</label><mixed-citation>Sheppard, E. J., Hurd, C. L., Britton, D. D., Reed, D. C., and Bach, L. T.: Seaweed biogeochemistry: Global assessment of <inline-formula><mml:math id="M360" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">N</mml:mi></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M361" display="inline"><mml:mrow class="chem"><mml:mi mathvariant="normal">C</mml:mi><mml:mo>:</mml:mo><mml:mi mathvariant="normal">P</mml:mi></mml:mrow></mml:math></inline-formula> ratios and implications for ocean afforestation, J. Phycol., 59, 879–892, <ext-link xlink:href="https://doi.org/10.1111/jpy.13381" ext-link-type="DOI">10.1111/jpy.13381</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx69"><label>Tagliabue et al.(2016)Tagliabue, Aumont, DeAth, Dunne, Dutkiewicz, Galbraith, Misumi, Moore, Ridgwell, Sherman, Stock, Vichi, Völker, and Yool</label><mixed-citation>Tagliabue, A., Aumont, O., DeAth, R., Dunne, J. P., Dutkiewicz, S., Galbraith, E., Misumi, K., Moore, J. K., Ridgwell, A., Sherman, E., Stock, C., Vichi, M., Völker, C., and Yool, A.: How well do global ocean biogeochemistry models simulate dissolved iron distributions?, Global Biogeochem. Cy., 30, 149–174, <ext-link xlink:href="https://doi.org/10.1002/2015GB005289" ext-link-type="DOI">10.1002/2015GB005289</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bibx70"><label>Tagliabue et al.(2023)Tagliabue, Twining, Barrier, Maury, Berger, and Bopp</label><mixed-citation>Tagliabue, A., Twining, B. S., Barrier, N., Maury, O., Berger, M., and Bopp, L.: Ocean iron fertilization may amplify climate change pressures on marine animal biomass for limited climate benefit, Glob. Change Biol., 29, 5250–5260, <ext-link xlink:href="https://doi.org/10.1111/gcb.16854" ext-link-type="DOI">10.1111/gcb.16854</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx71"><label>Tullberg et al.(2022)Tullberg, Nguyen, and Wang</label><mixed-citation>Tullberg, R. M., Nguyen, H. P., and Wang, C. M.: Review of the Status and Developments in Seaweed Farming Infrastructure, J. Mar. Sci. Eng., 10, <ext-link xlink:href="https://doi.org/10.3390/jmse10101447" ext-link-type="DOI">10.3390/jmse10101447</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bibx72"><label>Valderrama et al.(2013)Valderrama, Cai, Hishamunda, and Ridler</label><mixed-citation>Valderrama, D., Cai, J., Hishamunda, N., and Ridler, N.: Social and economic dimensions of carrageenan seaweed farming, Fisheries and aquaculture technical paper no. 580, Food and Agriculture Organization of the United Nations (FAO), Rome, Italy, <uri>https://www.fao.org/4/i3344e/i3344e.pdf</uri> (last access: 29 October 2025), 2013.</mixed-citation></ref>
      <ref id="bib1.bibx73"><label>Vancoppenolle et al.(2023)</label><mixed-citation>Vancoppenolle, M., Rousset, C., Blockley, E., Aksenov, Y., Feltham, D., Fichefet, T., Garric, G., Guémas, V., Iovino, D., Keeley, S., Madec, G., Massonnet, F., Ridley, J., Schroeder, D., and Tietsche, S.: Sea Ice modelling Integrated Initiative (SI<sup>3</sup>) – The NEMO sea ice engine, Zenodo [software], <ext-link xlink:href="https://doi.org/10.5281/zenodo.7534900" ext-link-type="DOI">10.5281/zenodo.7534900</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx74"><label>Van Der Molen et al.(2018)Van Der Molen, Ruardij, Mooney, Kerrison, O'Connor, Gorman, Timmermans, Wright, Kelly, Hughes, and Capuzzo</label><mixed-citation>van der Molen, J., Ruardij, P., Mooney, K., Kerrison, P., O'Connor, N. E., Gorman, E., Timmermans, K., Wright, S., Kelly, M., Hughes, A. D., and Capuzzo, E.: Modelling potential production of macroalgae farms in UK and Dutch coastal waters, Biogeosciences, 15, 1123–1147, <ext-link xlink:href="https://doi.org/10.5194/bg-15-1123-2018" ext-link-type="DOI">10.5194/bg-15-1123-2018</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bibx75"><label>Veenhof et al.(2024)Veenhof, Burrows, Hughes, Michalek, Ross, Thomson, Fedenko, and Stanley</label><mixed-citation>Veenhof, R. J., Burrows, M. T., Hughes, A. D., Michalek, K., Ross, M. E., Thomson, A. I., Fedenko, J., and Stanley, M. S.: Sustainable seaweed aquaculture and climate change in the North Atlantic: challenges and opportunities, Front. Mar. Sci., 11, 1483330, <ext-link xlink:href="https://doi.org/10.3389/fmars.2024.1483330" ext-link-type="DOI">10.3389/fmars.2024.1483330</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bibx76"><label>Wang et al.(2019)Wang, Hu, Barnes, Mitchum, Lapointe, and Montoya</label><mixed-citation>Wang, M., Hu, C., Barnes, B. B., Mitchum, G., Lapointe, B., and Montoya, J. P.: The great Atlantic <italic>Sargassum</italic> belt, Science, 365, 83–87, <ext-link xlink:href="https://doi.org/10.1126/science.aaw7912" ext-link-type="DOI">10.1126/science.aaw7912</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bibx77"><label>Wang et al.(2020)Wang, He, Ma, Huan, Wang, Xia, and Wang</label><mixed-citation>Wang, X., He, L., Ma, Y., Huan, L., Wang, Y., Xia, B., and Wang, G.: Economically important red algae resources along the Chinese coast: History, status, and prospects for their utilization, Algal Rese., 46, 101817, <ext-link xlink:href="https://doi.org/10.1016/j.algal.2020.101817" ext-link-type="DOI">10.1016/j.algal.2020.101817</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bibx78"><label>Wanninkhof(2014)</label><mixed-citation>Wanninkhof, R.: Relationship between wind speed and gas exchange over the ocean revisited, Limnol. Oceanogr.-Meth., 12, 351–362, <ext-link xlink:href="https://doi.org/10.4319/lom.2014.12.351" ext-link-type="DOI">10.4319/lom.2014.12.351</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bibx79"><label>Wu et al.(2023)Wu, Keller, and Oschlies</label><mixed-citation>Wu, J., Keller, D. P., and Oschlies, A.: Carbon dioxide removal via macroalgae open-ocean mariculture and sinking: an Earth system modeling study, Earth Syst. Dynam., 14, 185–221, <ext-link xlink:href="https://doi.org/10.5194/esd-14-185-2023" ext-link-type="DOI">10.5194/esd-14-185-2023</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bibx80"><label>Wu et al.(2025)Wu, Yao, Keller, and Oschlies</label><mixed-citation>Wu, J., Yao, W., Keller, D. P., and Oschlies, A.: Nearshore Macroalgae Cultivation for Carbon Sequestration by Biomass Harvesting: Evaluating Potential and Impacts With an Earth System Model, Geophys. Res. Lett., 52, e2025GL116774, <ext-link xlink:href="https://doi.org/10.1029/2025GL116774" ext-link-type="DOI">10.1029/2025GL116774</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bibx81"><label>Yamamoto et al.(2017)Yamamoto, Kato, Kanayama, Nakase, and Tsutsumi</label><mixed-citation>Yamamoto, M., Kato, T., Kanayama, S., Nakase, K., and Tsutsumi, N.: Effectiveness of Iron Fertilization for Seaweed Bed Restoration in Coastal Areas, J. Water Environ. Tech., 15, 186–197, <ext-link xlink:href="https://doi.org/10.2965/jwet.16-080" ext-link-type="DOI">10.2965/jwet.16-080</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bibx82"><label>Yool et al.(2013)Yool, Popova, and Anderson</label><mixed-citation>Yool, A., Popova, E. E., and Anderson, T. R.: MEDUSA-2.0: an intermediate complexity biogeochemical model of the marine carbon cycle for climate change and ocean acidification studies, Geosci. Model Dev., 6, 1767–1811, <ext-link xlink:href="https://doi.org/10.5194/gmd-6-1767-2013" ext-link-type="DOI">10.5194/gmd-6-1767-2013</ext-link>, 2013. </mixed-citation></ref>
      <ref id="bib1.bibx83"><label>Yool et al.(2021)Yool, Palmiéri, Jones, de Mora, Kuhlbrodt, Popova, Nurser, Hirschi, Blaker, Coward, Blockley, and Sellar</label><mixed-citation>Yool, A., Palmiéri, J., Jones, C. G., de Mora, L., Kuhlbrodt, T., Popova, E. E., Nurser, A. J. G., Hirschi, J., Blaker, A. T., Coward, A. C., Blockley, E. W., and Sellar, A. A.: Evaluating the physical and biogeochemical state of the global ocean component of UKESM1 in CMIP6 historical simulations, Geosci. Model Dev., 14, 3437–3472, <ext-link xlink:href="https://doi.org/10.5194/gmd-14-3437-2021" ext-link-type="DOI">10.5194/gmd-14-3437-2021</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bibx84"><label>Zhang et al.(2012)Zhang, Fang, Wang et al.</label><mixed-citation>Zhang, J., Fang, J., Wang, W., Du, M., Gao, Y., and Zhang, M.: Growth and loss of mariculture kelp <italic>Saccharina japonica</italic> in Sungo Bay, China, J. Appl. Phycol., 24, 1209–1216, <ext-link xlink:href="https://doi.org/10.1007/s10811-011-9762-4" ext-link-type="DOI">10.1007/s10811-011-9762-4</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bibx85"><label>Zhang et al.(2016)Zhang, Wu, Ren, and Lin</label><mixed-citation>Zhang, J., Wu, W., Ren, J. S., and Lin, F.: A model for the growth of mariculture kelp <italic>Saccharina japonica</italic> in Sanggou Bay, China, Aquacult. Env. Interac., 8, 273–283, <ext-link xlink:href="https://doi.org/10.3354/aei00171" ext-link-type="DOI">10.3354/aei00171</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bibx86"><label>Zollmann et al.(2023)Zollmann, Liberzon, Palatnik et al.</label><mixed-citation>Zollmann, M., Liberzon, A., Palatnik, R. R., Zilberman, D., and Golberg, A.: Effects of season, depth and pre-cultivation fertilizing on Ulva growth dynamics offshore the Eastern Mediterranean Sea, Sci. Rep., 13, 14784, <ext-link xlink:href="https://doi.org/10.1038/s41598-023-41605-4" ext-link-type="DOI">10.1038/s41598-023-41605-4</ext-link>, 2023.</mixed-citation></ref>

  </ref-list></back>
    <!--<article-title-html>The impact of large-scale macroalgae cultivation and harvesting strategies on the marine carbon dioxide removal efficacy and marine biogeochemistry</article-title-html>
<abstract-html/>
<ref-html id="bib1.bib1"><label>Aldridge et al.(2021)Aldridge, Mooney, Dabrowski, and Capuzzo</label><mixed-citation>
      
Aldridge, J., Mooney, K., Dabrowski, T., and Capuzzo, E.:
Modelling effects of seaweed aquaculture on phytoplankton and mussel production. Application to Strangford Lough (Northern Ireland), Aquaculture, 536, 736400, <a href="https://doi.org/10.1016/j.aquaculture.2021.736400" target="_blank">https://doi.org/10.1016/j.aquaculture.2021.736400</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib2"><label>Alevizos and Barillé(2023)</label><mixed-citation>
      
Alevizos, E. and Barillé, L.:
Global ocean spatial suitability for macroalgae offshore cultivation and sinking, Front. Mar. Sci., 10, 1320642, <a href="https://doi.org/10.3389/fmars.2023.1320642" target="_blank">https://doi.org/10.3389/fmars.2023.1320642</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib3"><label>Anugerahanti et al.(2026)Anugerahanti, Palmieri, and Yool</label><mixed-citation>
      
Anugerahanti, P., Palmieri, J., and Yool, A.:
The Impact of Large-Scale Macroalgae Cultivation and Harvesting Strategies on the Marine Carbon Dioxide Removal Efficacy and Marine Biogeochemistry – Supplementary materials, Zenodo [code and data set], <a href="https://doi.org/10.5281/zenodo.20154696" target="_blank">https://doi.org/10.5281/zenodo.20154696</a>, 2026.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib4"><label>Arzeno-Soltero et al.(2023)Arzeno-Soltero, Saenz, Frieder, Long, DeAngelo, Davis, and Davis</label><mixed-citation>
      
Arzeno-Soltero, I. B., Saenz, B. T., Frieder, C. A., Long, M. C., DeAngelo, J., Davis, S. J., and Davis, K. A.:
Large global variations in the carbon dioxide removal potential of seaweed farming due to biophysical constraints, Commun. Earth Environ., 4, 185, <a href="https://doi.org/10.1038/s43247-023-00833-2" target="_blank">https://doi.org/10.1038/s43247-023-00833-2</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib5"><label>Bach et al.(2021)Bach, Tamsitt, Gower, Hurd, Raven, and Boyd</label><mixed-citation>
      
Bach, L. T., Tamsitt, V., Gower, J., Hurd, C. L., Raven, J. A., and Boyd, P. W.:
Testing the climate intervention potential of ocean afforestation using the Great Atlantic Sargassum Belt, Nat. Commun., 12, 2556, <a href="https://doi.org/10.1038/s41467-021-22837-2" target="_blank">https://doi.org/10.1038/s41467-021-22837-2</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib6"><label>Bach et al.(2024)Bach, Vaughan, Law, and Williamson</label><mixed-citation>
      
Bach, L. T., Vaughan, N. E., Law, C. S., and Williamson, P.:
Implementation of marine CO2 removal for climate mitigation: The challenges of additionality, predictability, and governability, Elem. Sci. Anth., 12, 00034, <a href="https://doi.org/10.1525/elementa.2023.00034" target="_blank">https://doi.org/10.1525/elementa.2023.00034</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib7"><label>Bak et al.(2018)Bak, Mols-Mortensen, and Gregersen</label><mixed-citation>
      
Bak, U. G., Mols-Mortensen, A., and Gregersen, O.:
Production method and cost of commercial-scale offshore cultivation of kelp in the Faroe Islands using multiple partial harvesting, Algal Res., 33, 36–47, <a href="https://doi.org/10.1016/j.algal.2018.05.001" target="_blank">https://doi.org/10.1016/j.algal.2018.05.001</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib8"><label>Bak et al.(2020)Bak, Gregersen, and Infante</label><mixed-citation>
      
Bak, U. G., Gregersen, O., and Infante, J.:
Technical challenges for offshore cultivation of kelp species: lessons learned and future directions, Bot. Mar., 63, 341–353, <a href="https://doi.org/10.1515/bot-2019-0005" target="_blank">https://doi.org/10.1515/bot-2019-0005</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib9"><label>Barrett et al.(2022)Barrett, Theuerkauf, Rose, Alleway, Bricker, Parker, Petrolia, and Jones</label><mixed-citation>
      
Barrett, L. T., Theuerkauf, S. J., Rose, J. M., Alleway, H. K., Bricker, S. B., Parker, M., Petrolia, D. R., and Jones, R. C.:
Sustainable growth of non-fed aquaculture can generate valuable ecosystem benefits, Ecosyst. Serv., 53, 101396, <a href="https://doi.org/10.1016/j.ecoser.2021.101396" target="_blank">https://doi.org/10.1016/j.ecoser.2021.101396</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib10"><label>Behera et al.(2022)Behera, Vadodariya, Veeragurunathan, Sigamani, Moovendhan, Srinivasan, Kolandhasamy, and Ingle</label><mixed-citation>
      
Behera, D. P., Vadodariya, V., Veeragurunathan, V., Sigamani, S., Moovendhan, M., Srinivasan, R., Kolandhasamy, P., and Ingle, K. N.:
Seaweeds cultivation methods and their role in climate mitigation and environmental cleanup, Total Environment Research Themes, 3–4, 100016, <a href="https://doi.org/10.1016/j.totert.2022.100016" target="_blank">https://doi.org/10.1016/j.totert.2022.100016</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib11"><label>Berger et al.(2023)Berger, Kwiatkowski, Ho, and Bopp</label><mixed-citation>
      
Berger, M., Kwiatkowski, L., Ho, D. T., and Bopp, L.:
Ocean dynamics and biological feedbacks limit the potential of macroalgae carbon dioxide removal, Environ. Res. Lett., 18, 024039, <a href="https://doi.org/10.1088/1748-9326/acb06e" target="_blank">https://doi.org/10.1088/1748-9326/acb06e</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib12"><label>Berger et al.(2025)Berger, Kwiatkowski, Bopp, and Ho</label><mixed-citation>
      
Berger, M., Kwiatkowski, L., Bopp, L., and Ho, D. T.:
Efficacy of seaweed-based carbon dioxide removal reduced by iron limitation and nutrient competition with phytoplankton, CDRxiv [Preprints], <a href="https://doi.org/10.70212/cdrxiv.2025385.v1" target="_blank">https://doi.org/10.70212/cdrxiv.2025385.v1</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib13"><label>Boderskov et al.(2023)Boderskov, Rasmussen, and Bruhn</label><mixed-citation>
      
Boderskov, T., Rasmussen, M. B., and Bruhn, A.:
Upscaling cultivation of <i>Saccharina latissima</i> on net or line systems; comparing biomass yields and nutrient extraction potentials, Front. Mar. Sci., 10, 992179, <a href="https://doi.org/10.3389/fmars.2023.992179" target="_blank">https://doi.org/10.3389/fmars.2023.992179</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib14"><label>Boyd and Vivian(2019)</label><mixed-citation>
      
Boyd, P. and Vivian, C. M. G., e.:
GESAMP “High level review of a wide range of proposed marine geoengineering techniques”, Tech. rep., Joint Group of Experts on the Scientific Aspects of Marine Environmental Protection, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib15"><label>Boyd et al.(2022)Boyd, Bach, Hurd, Paine, Raven, and Tamsitt</label><mixed-citation>
      
Boyd, P. W., Bach, L. T., Hurd, C. L., Paine, E., Raven, J. A., and Tamsitt, V.:
Potential negative effects of ocean afforestation on offshore ecosystems, Nat. Ecol. Evol., 6, 675–683, <a href="https://doi.org/10.1038/s41559-022-01722-1" target="_blank">https://doi.org/10.1038/s41559-022-01722-1</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib16"><label>Bullen et al.(2024)Bullen, Driscoll, Burt, Stephens, Hessing-Lewis, and Gregr</label><mixed-citation>
      
Bullen, C. D., Driscoll, J., Burt, J., Stephens, T., Hessing-Lewis, M., and Gregr, E. J.:
The potential climate benefits of seaweed farming in temperate waters, Sci. Rep., 14, 15021, <a href="https://doi.org/10.1038/s41598-024-65408-3" target="_blank">https://doi.org/10.1038/s41598-024-65408-3</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib17"><label>Chen et al.(2020)Chen, Li, Zhang, He, Shi, Jiao, and Zhang</label><mixed-citation>
      
Chen, J., Li, H., Zhang, Z., He, C., Shi, Q., Jiao, N., and Zhang, Y.:
DOC dynamics and bacterial community succession during long-term degradation of <i>Ulva prolifera</i> and their implications for the legacy effect of green tides on refractory DOC pool in seawater, Water Res., 185, 116268, <a href="https://doi.org/10.1016/j.watres.2020.116268" target="_blank">https://doi.org/10.1016/j.watres.2020.116268</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib18"><label>Choi et al.(2025)Choi, Lee, Jung, Lee, Han, Hyeon, and Choi</label><mixed-citation>
      
Choi, D., Lee, H.-G., Jung, Y.-H., Lee, D.-W., Han, J., Hyeon, J.-Y., and Choi, Y.-U.:
Pilot-Scale Cultivation of Seaweed (<i>Undaria pinnatifida</i>) Along an Offshore Wind Farm in Southwestern Korea, J. Mar. Sci. Eng., 13, <a href="https://doi.org/10.3390/jmse13050882" target="_blank">https://doi.org/10.3390/jmse13050882</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib19"><label>Chopin et al.(2024)Chopin, Costa-Pierce, Troell, Hurd, Costello, Backman, Buschmann, Cuhel, Duarte, Gröndahl, Heasman, Haroun, Johansen, Jueterbock, Lench, Lindell, Pavia, Ricart, Sundell, and Yarish</label><mixed-citation>
      
Chopin, T., Costa-Pierce, B. A., Troell, M., Hurd, C. L., Costello, M. J., Backman, S., Buschmann, A. H., Cuhel, R., Duarte, C. M., Gröndahl, F., Heasman, K., Haroun, R. J., Johansen, J., Jueterbock, A., Lench, M., Lindell, S., Pavia, H., Ricart, A. M., Sundell, K. S., and Yarish, C.:
Deep-ocean seaweed dumping for carbon sequestration: Questionable, risky, and not the best use of valuable biomass, One Earth, 7, S2590332224000356, <a href="https://doi.org/10.1016/j.oneear.2024.01.013" target="_blank">https://doi.org/10.1016/j.oneear.2024.01.013</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib20"><label>Chung et al.(2011)Chung, Beardall, Mehta, Sahoo, and Stojkovic</label><mixed-citation>
      
Chung, I. K., Beardall, J., Mehta, S., Sahoo, D., and Stojkovic, S.:
Using marine macroalgae for carbon sequestration: a critical appraisal, J. Appl. Phycol., 23, 877–886, <a href="https://doi.org/10.1007/s10811-010-9604-9" target="_blank">https://doi.org/10.1007/s10811-010-9604-9</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib21"><label>Coleman et al.(2022)Coleman, Dewhurst, Fredriksson, St. Gelais, Cole, MacNicoll, Laufer, and Brady</label><mixed-citation>
      
Coleman, S., Dewhurst, T., Fredriksson, D. W., St. Gelais, A. T., Cole, K. L., MacNicoll, M., Laufer, E., and Brady, D. C.:
Quantifying baseline costs and cataloging potential optimization strategies for kelp aquaculture carbon dioxide removal, Front. Marine Sci., 9, <a href="https://doi.org/10.3389/fmars.2022.966304" target="_blank">https://doi.org/10.3389/fmars.2022.966304</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib22"><label>Corrigan et al.(2022)Corrigan, Brown, Ashton, Smale, and Tyler</label><mixed-citation>
      
Corrigan, S., Brown, A. R., Ashton, I. G. C., Smale, D. A., and Tyler, C. R.:
Quantifying habitat provisioning at macroalgal cultivation sites, Rev. Aquacult., 14, 1671–1694, <a href="https://doi.org/10.1111/raq.12669" target="_blank">https://doi.org/10.1111/raq.12669</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib23"><label>DeAngelo et al.(2021)DeAngelo, Azevedo, Bistline, Clarke, Luderer, Byers, and Davis</label><mixed-citation>
      
DeAngelo, J., Azevedo, I., Bistline, J., Clarke, L., Luderer, G., Byers, E., and Davis, S. J.:
Energy systems in scenarios at net-zero CO2 emissions, Nat. Commun., 12, 6096, <a href="https://doi.org/10.1038/s41467-021-26356-y" target="_blank">https://doi.org/10.1038/s41467-021-26356-y</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib24"><label>DeAngelo et al.(2022)DeAngelo, Saenz, Arzeno-Soltero, Frieder, Long, Hamman, Davis, and Davis</label><mixed-citation>
      
DeAngelo, J., Saenz, B. T., Arzeno-Soltero, I. B., Frieder, C. A., Long, M. C., Hamman, J., Davis, K. A., and Davis, S. J.:
Economic and biophysical limits to seaweed farming for climate change mitigation, Nat. Plants, 9, 45–57, <a href="https://doi.org/10.1038/s41477-022-01305-9" target="_blank">https://doi.org/10.1038/s41477-022-01305-9</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib25"><label>Duarte et al.(2017)Duarte, Wu, Xiao, Bruhn, and Krause-Jensen</label><mixed-citation>
      
Duarte, C. M., Wu, J., Xiao, X., Bruhn, A., and Krause-Jensen, D.:
Can Seaweed Farming Play a Role in Climate Change Mitigation and Adaptation?, Front. Mar. Sci., 4, <a href="https://doi.org/10.3389/fmars.2017.00100" target="_blank">https://doi.org/10.3389/fmars.2017.00100</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib26"><label>Duarte et al.(2021)Duarte, Bruhn, and Krause-Jensen</label><mixed-citation>
      
Duarte, C. M., Bruhn, A., and Krause-Jensen, D.:
A seaweed aquaculture imperative to meet global sustainability targets, Nat. Sustain., 5, 185–193, <a href="https://doi.org/10.1038/s41893-021-00773-9" target="_blank">https://doi.org/10.1038/s41893-021-00773-9</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib27"><label>Enríquez et al.(1996)Enríquez, Duarte, Sand-Jensen, and Nielsen</label><mixed-citation>
      
Enríquez, S., Duarte, C. M., Sand-Jensen, K., and Nielsen, S. L.:
Broad-scale comparison of photosynthetic rates across phototrophic organisms, Oecologia, 108, 197–206, <a href="https://doi.org/10.1007/BF00334642" target="_blank">https://doi.org/10.1007/BF00334642</a>, 1996.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib28"><label>Froehlich et al.(2019)Froehlich, Afflerbach, Frazier, and Halpern</label><mixed-citation>
      
Froehlich, H. E., Afflerbach, J. C., Frazier, M., and Halpern, B. S.:
Blue Growth Potential to Mitigate Climate Change through Seaweed Offsetting, Curr. Biol., 29, 3087–3093.e3, <a href="https://doi.org/10.1016/j.cub.2019.07.041" target="_blank">https://doi.org/10.1016/j.cub.2019.07.041</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib29"><label>Gallagher et al.(2022)Gallagher, Shelamoff, and Layton</label><mixed-citation>
      
Gallagher, J. B., Shelamoff, V., and Layton, C.:
Seaweed ecosystems may not mitigate CO2 emissions, ICES J. Mar. Sci., 79, 585–592, <a href="https://doi.org/10.1093/icesjms/fsac011" target="_blank">https://doi.org/10.1093/icesjms/fsac011</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib30"><label>Guibourd de Luzinais et al.(2023)Guibourd de Luzinais, du Pontavice, Reygondeau, Barrier, Blanchard, Bornarel, Büchner, Cheung, Eddy, Everett, Guiet, Harrison, Maury, Novaglio, Petrik, Steenbeek, Tittensor, and Gascuel</label><mixed-citation>
      
Guibourd de Luzinais, V., du Pontavice, H., Reygondeau, G., Barrier, N., Blanchard, J. L., Bornarel, V., Büchner, M., Cheung, W. W. L., Eddy, T. D., Everett, J. D., Guiet, J., Harrison, C. S., Maury, O., Novaglio, C., Petrik, C. M., Steenbeek, J., Tittensor, D. P., and Gascuel, D.:
Trophic amplification: A model intercomparison of climate driven changes in marine food webs, PLOS ONE, 18, 1–23, <a href="https://doi.org/10.1371/journal.pone.0287570" target="_blank">https://doi.org/10.1371/journal.pone.0287570</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib31"><label>Hayashi et al.(2017)Hayashi, Reis, Dos Santos, Castelar, Robledo, De Vega, Msuya, Eswaran, Yasir, Ali, and Hurtado</label><mixed-citation>
      
Hayashi, L., Reis, R. P., Dos Santos, A. A., Castelar, B., Robledo, D., De Vega, G. B., Msuya, F. E., Eswaran, K., Yasir, S. M., Ali, M. K. M., and Hurtado, A. Q.:
The Cultivation of Kappaphycus and Eucheuma in Tropical and Sub-Tropical Waters, in: Tropical Seaweed Farming Trends, Problems and Opportunities, edited by: Hurtado, A. Q., Critchley, A. T., and Neish, I. C., Springer International Publishing, Cham, pp. 55–90, <a href="https://doi.org/10.1007/978-3-319-63498-2_4" target="_blank">https://doi.org/10.1007/978-3-319-63498-2_4</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib32"><label>Hersbach et al.(2020)Hersbach, Bell, Berrisford, Hirahara, Horányi, Muñoz-Sabater, Nicolas, Peubey, Radu, Schepers, Simmons, Soci, Abdalla, Abellan, Balsamo, Bechtold, Biavati, Bidlot, Bonavita, De Chiara, Dahlgren, Dee, Diamantakis, Dragani, Flemming, Forbes, Fuentes, Geer, Haimberger, Healy, Hogan, Hólm, Janisková, Keeley, Laloyaux, Lopez, Lupu, Radnoti, de Rosnay, Rozum, Vamborg, Villaume, and Thépaut</label><mixed-citation>
      
Hersbach, H., Bell, B., Berrisford, P., Hirahara, S., Horányi, A., Muñoz-Sabater, J., Nicolas, J., Peubey, C., Radu, R., Schepers, D., Simmons, A., Soci, C., Abdalla, S., Abellan, X., Balsamo, G., Bechtold, P., Biavati, G., Bidlot, J., Bonavita, M., De Chiara, G., Dahlgren, P., Dee, D., Diamantakis, M., Dragani, R., Flemming, J., Forbes, R., Fuentes, M., Geer, A., Haimberger, L., Healy, S., Hogan, R. J., Hólm, E., Janisková, M., Keeley, S., Laloyaux, P., Lopez, P., Lupu, C., Radnoti, G., de Rosnay, P., Rozum, I., Vamborg, F., Villaume, S., and Thépaut, J.-N.:
The ERA5 global reanalysis, Q. J. Roy. Meteor. Soc., 146, 1999–2049, <a href="https://doi.org/10.1002/qj.3803" target="_blank">https://doi.org/10.1002/qj.3803</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib33"><label>Hurd et al.(2014)Hurd, Harrison, Bischof, and Lobban</label><mixed-citation>
      
Hurd, C. L., Harrison, P. J., Bischof, K., and Lobban, C. S.:
Seaweed Ecology and Physiology, Cambridge University Press, Cambridge, ISBN
9781139192637, <a href="https://doi.org/10.1017/CBO9781139192637" target="_blank">https://doi.org/10.1017/CBO9781139192637</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib34"><label>Hurd et al.(2022)Hurd, Law, Bach, Britton, Hovenden, Paine, Raven, Tamsitt, and Boyd</label><mixed-citation>
      
Hurd, C. L., Law, C. S., Bach, L. T., Britton, D., Hovenden, M., Paine, E. R., Raven, J. A., Tamsitt, V., and Boyd, P. W.:
Forensic carbon accounting: Assessing the role of seaweeds for carbon sequestration, J. Phycol., 58, 347–363, <a href="https://doi.org/10.1111/jpy.13249" target="_blank">https://doi.org/10.1111/jpy.13249</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib35"><label>Hurd et al.(2023)Hurd, Gattuso, and Boyd</label><mixed-citation>
      
Hurd, C. L., Gattuso, J., and Boyd, P. W.:
Air-sea carbon dioxide equilibrium: Will it be possible to use seaweeds for carbon removal offsets?, J. Phycol., 60, jpy.13405, <a href="https://doi.org/10.1111/jpy.13405" target="_blank">https://doi.org/10.1111/jpy.13405</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib36"><label>Hwang et al.(2018)Hwang, Liu, Lee, Ha, and Park</label><mixed-citation>
      
Hwang, E. K., Liu, F., Lee, K. H., Ha, D. S., and Park, C. S.:
Comparison of the cultivation performance between Korean (Sugwawon No. 301) and Chinese strains (Huangguan No. 1) of kelp <i>Saccharina japonica</i> in an aquaculture farm in Korea, Algae, 33, 101–108, <a href="https://doi.org/10.4490/algae.2018.33.2.4" target="_blank">https://doi.org/10.4490/algae.2018.33.2.4</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib37"><label>IPCC(2022)</label><mixed-citation>
      
IPCC:
Summary for Policymakers, Cambridge University Press, 3–48, <a href="https://doi.org/10.1017/9781009157896.001" target="_blank">https://doi.org/10.1017/9781009157896.001</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib38"><label>Jiang et al.(2013)Jiang, Fang, Mao, Han, and Wang</label><mixed-citation>
      
Jiang, Z., Fang, J., Mao, Y., Han, T., and Wang, G.:
Influence of Seaweed Aquaculture on Marine Inorganic Carbon Dynamics and Sea-air CO<sub>2</sub> Flux, J. World Aquacult. Soc., 44, 133–140, <a href="https://doi.org/10.1111/jwas.12000" target="_blank">https://doi.org/10.1111/jwas.12000</a>, 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib39"><label>Jiao et al.(2025)Jiao, Feng, Li, and Tian</label><mixed-citation>
      
Jiao, T., Feng, E. Y., Li, Y., and Tian, Y.:
Carbon dioxide removal dilemma of macroalgae products: Evidence from carbon footprint and profitability, J. Clean. Prod., 492, 144870, <a href="https://doi.org/10.1016/j.jclepro.2025.144870" target="_blank">https://doi.org/10.1016/j.jclepro.2025.144870</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib40"><label>Jones et al.(2014)Jones, Ito, Takano, and Hsu</label><mixed-citation>
      
Jones, D. C., Ito, T., Takano, Y., and Hsu, W.-C.:
Spatial and seasonal variability of the air-sea equilibration timescale of carbon dioxide, Global Biogeochem. Cy., 28, 1163–1178, <a href="https://doi.org/10.1002/2014GB004813" target="_blank">https://doi.org/10.1002/2014GB004813</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib41"><label>Kennedy and Blain(2025)</label><mixed-citation>
      
Kennedy, J. R. and Blain, C. O.:
A systematic review of marine macroalgal degradation: Toward a better understanding of macroalgal carbon sequestration potential, J. Phycol., 61, 399–432, <a href="https://doi.org/10.1111/jpy.70031" target="_blank">https://doi.org/10.1111/jpy.70031</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib42"><label>Koesling et al.(2021)Koesling, Kvadsheim, Halfdanarson, Emblemsvåg, and Rebours</label><mixed-citation>
      
Koesling, M., Kvadsheim, N. P., Halfdanarson, J., Emblemsvåg, J., and Rebours, C.:
Environmental impacts of protein-production from farmed seaweed: Comparison of possible scenarios in Norway, J. Clean. Prod., 307, 127301, <a href="https://doi.org/10.1016/j.jclepro.2021.127301" target="_blank">https://doi.org/10.1016/j.jclepro.2021.127301</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib43"><label>Krause-Jensen and Duarte(2016)</label><mixed-citation>
      
Krause-Jensen, D. and Duarte, C. M.:
Substantial role of macroalgae in marine carbon sequestration, Nat. Geosci., 9, 737–742, <a href="https://doi.org/10.1038/ngeo2790" target="_blank">https://doi.org/10.1038/ngeo2790</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib44"><label>Kwiatkowski et al.(2019)Kwiatkowski, Aumont, and Bopp</label><mixed-citation>
      
Kwiatkowski, L., Aumont, O., and Bopp, L.:
Consistent trophic amplification of marine biomass declines under climate change, Glob. Change Biol., 25, 218–229, <a href="https://doi.org/10.1111/gcb.14468" target="_blank">https://doi.org/10.1111/gcb.14468</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib45"><label>Kwon et al.(2024)Kwon, Hawkins, Zaimes, Infante, Kite-Powell, Stekoll, Roberson, Zotter, Augyte, Rocheleau, and Sims</label><mixed-citation>
      
Kwon, H., Hawkins, T. R., Zaimes, G. G., Infante, J., Kite-Powell, H. L., Stekoll, M. S., Roberson, L., Zotter, B., Augyte, S., Rocheleau, G., and Sims, N.:
Life-cycle analysis of offshore macroalgae production systems in the United States, Algal Rese., 82, 103654, <a href="https://doi.org/10.1016/j.algal.2024.103654" target="_blank">https://doi.org/10.1016/j.algal.2024.103654</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib46"><label>Lauvset et al.(2016)Lauvset, Key, Olsen, van Heuven, Velo, Lin, Schirnick, Kozyr, Tanhua, Hoppema, Jutterström, Steinfeldt, Jeansson, Ishii, Perez, Suzuki, and Watelet</label><mixed-citation>
      
Lauvset, S. K., Key, R. M., Olsen, A., van Heuven, S., Velo, A., Lin, X., Schirnick, C., Kozyr, A., Tanhua, T., Hoppema, M., Jutterström, S., Steinfeldt, R., Jeansson, E., Ishii, M., Perez, F. F., Suzuki, T., and Watelet, S.:
A new global interior ocean mapped climatology: the 1°&thinsp; × &thinsp;1° GLODAP version 2, Earth Syst. Sci. Data, 8, 325–340, <a href="https://doi.org/10.5194/essd-8-325-2016" target="_blank">https://doi.org/10.5194/essd-8-325-2016</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib47"><label>Levin et al.(2009)Levin, Ekau, Gooday, Jorissen, Middelburg, Naqvi, Neira, Rabalais, and Zhang</label><mixed-citation>
      
Levin, L. A., Ekau, W., Gooday, A. J., Jorissen, F., Middelburg, J. J., Naqvi, S. W. A., Neira, C., Rabalais, N. N., and Zhang, J.:
Effects of natural and human-induced hypoxia on coastal benthos, Biogeosciences, 6, 2063–2098, <a href="https://doi.org/10.5194/bg-6-2063-2009" target="_blank">https://doi.org/10.5194/bg-6-2063-2009</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib48"><label>Levin et al.(2023)Levin, Alfaro-Lucas, Colaço, Cordes, Craik, Danovaro, Hoving, Ingels, Mestre, Seabrook, Thurber, Vivian, and Yasuhara</label><mixed-citation>
      
Levin, L. A., Alfaro-Lucas, J. M., Colaço, A., Cordes, E. E., Craik, N., Danovaro, R., Hoving, H.-J., Ingels, J., Mestre, N. C., Seabrook, S., Thurber, A. R., Vivian, C., and Yasuhara, M.:
Deep-sea impacts of climate interventions, Science, 379, 978–981, <a href="https://doi.org/10.1126/science.ade7521" target="_blank">https://doi.org/10.1126/science.ade7521</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib49"><label>Macreadie et al.(2021)Macreadie, Costa, Atwood, Friess, Kelleway, Kennedy, Lovelock, Serrano, and Duarte</label><mixed-citation>
      
Macreadie, P. I., Costa, M. D. P., Atwood, T. B., Friess, D. A., Kelleway, J. J., Kennedy, H., Lovelock, C. E., Serrano, O., and Duarte, C. M.:
Blue carbon as a natural climate solution, Nat. Rev. Earth Environ., 2, 826–839, <a href="https://doi.org/10.1038/s43017-021-00224-1" target="_blank">https://doi.org/10.1038/s43017-021-00224-1</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib50"><label>Madec(2016)</label><mixed-citation>
      
Madec, G.:
NEMO ocean engine, Tech. rep., publication Title: Note du Pole de modelisation Issue: 27, Institut Pierre-Simon Laplace No 27, ISSN No 1288-1619, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib51"><label>Magcanta-Mortos et al.(2025)Magcanta-Mortos, Tahiluddin, Mortos, Aspe, Aaron-Amper, Leopardas, and Uy</label><mixed-citation>
      
Magcanta-Mortos, M. L. M., Tahiluddin, A. B., Mortos, J. M. R., Aspe, N. M., Aaron-Amper, J., Leopardas, V. E., and Uy, W. H.:
Sargassum hatchery-based cultivation and mariculture techniques in the Philippines: Practices, challenges, and future prospects, Aquat. Bot., 201, 103925, <a href="https://doi.org/10.1016/j.aquabot.2025.103925" target="_blank">https://doi.org/10.1016/j.aquabot.2025.103925</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib52"><label>Naylor et al.(2021)Naylor, Hardy, Buschmann, Bush, Cao, Klinger, Little, Lubchenco, Shumway, and Troell</label><mixed-citation>
      
Naylor, R. L., Hardy, R. W., Buschmann, A. H., Bush, S. R., Cao, L., Klinger, D. H., Little, D. C., Lubchenco, J., Shumway, S. E., and Troell, M.:
A 20-year retrospective review of global aquaculture, Nature, 591, 551–563, <a href="https://doi.org/10.1038/s41586-021-03308-6" target="_blank">https://doi.org/10.1038/s41586-021-03308-6</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib53"><label>N‘Yeurt et al.(2012)N‘Yeurt, Chynoweth, Capron, Stewart, and Hasan</label><mixed-citation>
      
N‘Yeurt, A. D. R., Chynoweth, D. P., Capron, M. E., Stewart, J. R., and Hasan, M. A.:
Negative carbon via Ocean Afforestation, Process Saf. Environ., 90, 467–474, <a href="https://doi.org/10.1016/j.psep.2012.10.008" target="_blank">https://doi.org/10.1016/j.psep.2012.10.008</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib54"><label>Ocean Visions and Monterey Bay Aquarium Research Institute(2022)</label><mixed-citation>
      
Ocean Visions and Monterey Bay Aquarium Research Institute:
Answering Critical Questions About Sinking Macroalgae for Carbon Dioxide Removal: A Research Framework to Investigate Sequestration Efficacy and Environmental Impacts, Tech. rep., Ocean Visions, Monterey Bay Aquarium Research Institute, <a href="https://oceanvisions.org/wp-content/uploads/2022/10/Ocean-Visions-Sinking-Seaweed-Report_FINAL.pdf" target="_blank"/> (last access: 29 October 2025), 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib55"><label>Orr and Epitalon(2015)</label><mixed-citation>
      
Orr, J. C. and Epitalon, J.-M.:
Improved routines to model the ocean carbonate system: mocsy 2.0, Geosci. Model Dev., 8, 485–499, <a href="https://doi.org/10.5194/gmd-8-485-2015" target="_blank">https://doi.org/10.5194/gmd-8-485-2015</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib56"><label>Paine et al.(2021)Paine, Schmid, Boyd, Diaz-Pulido, and Hurd</label><mixed-citation>
      
Paine, E. R., Schmid, M., Boyd, P. W., Diaz-Pulido, G., and Hurd, C. L.:
Rate and fate of dissolved organic carbon release by seaweeds: A missing link in the coastal ocean carbon cycle, J. Phycol., 57, 1375–1391, <a href="https://doi.org/10.1111/jpy.13198" target="_blank">https://doi.org/10.1111/jpy.13198</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib57"><label>Paine et al.(2023)Paine, Boyd, Strzepek, Ellwood, Brewer, Diaz-Pulido, Schmid, and Hurd</label><mixed-citation>
      
Paine, E. R., Boyd, P. W., Strzepek, R. F., Ellwood, M., Brewer, E. A., Diaz-Pulido, G., Schmid, M., and Hurd, C. L.:
Iron limitation of kelp growth may prevent ocean afforestation, Commun. Biol., 6, 607, <a href="https://doi.org/10.1038/s42003-023-04962-4" target="_blank">https://doi.org/10.1038/s42003-023-04962-4</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib58"><label>Palmiéri and Yool(2024)</label><mixed-citation>
      
Palmiéri, J. and Yool, A.:
Global-Scale Evaluation of Coastal Ocean Alkalinity Enhancement in a Fully Coupled Earth System Model, Earths Future, 12, e2023EF004018, <a href="https://doi.org/10.1029/2023EF004018" target="_blank">https://doi.org/10.1029/2023EF004018</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib59"><label>Pessarrodona et al.(2023)Pessarrodona, Franco-Santos, Wright, Vanderklift, Howard, Pidgeon, Wernberg, and Filbee-Dexter</label><mixed-citation>
      
Pessarrodona, A., Franco-Santos, R. M., Wright, L. S., Vanderklift, M. A., Howard, J., Pidgeon, E., Wernberg, T., and Filbee-Dexter, K.:
Carbon sequestration and climate change mitigation using macroalgae: a state of knowledge review, Biol. Rev., 98, 1945–1971, <a href="https://doi.org/10.1111/brv.12990" target="_blank">https://doi.org/10.1111/brv.12990</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib60"><label>Peteiro et al.(2014)Peteiro, Sánchez, Dueñas-Liaño, and Martínez</label><mixed-citation>
      
Peteiro, C., Sánchez, N., Dueñas-Liaño, C., and Martínez, B.:
Open-sea cultivation by transplanting young fronds of the kelp Saccharina latissima, J. Appl. Phycol., 26, 519–528, <a href="https://doi.org/10.1007/s10811-013-0096-2" target="_blank">https://doi.org/10.1007/s10811-013-0096-2</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib61"><label>Reagan et al.(2024)Reagan, Boyer, García, Locarnini, Baranova, Bouchard, Cross, Mishonov, Paver, Seidov, Wang, and Dukhovskoy</label><mixed-citation>
      
Reagan, J. R., Boyer, T. P., García, H. E., Locarnini, R. A., Baranova, O. K., Bouchard, C., Cross, S. L., Mishonov, A. V., Paver, C. R., Seidov, D., Wang, Z., and Dukhovskoy, D.:
World Ocean Atlas 2023, NCEI [data set], <a href="https://doi.org/10.25921/va26-hv25" target="_blank">https://doi.org/10.25921/va26-hv25</a>,  2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib62"><label>Roberts et al.(2015)Roberts, Paul, Dworjanyn, Bird, and De Nys</label><mixed-citation>
      
Roberts, D. A., Paul, N. A., Dworjanyn, S. A., Bird, M. I., and De Nys, R.:
Biochar from commercially cultivated seaweed for soil amelioration, Sci. Rep., 5, 9665, <a href="https://doi.org/10.1038/srep09665" target="_blank">https://doi.org/10.1038/srep09665</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib63"><label>Roque et al.(2019)Roque, Salwen, Kinley, and Kebreab</label><mixed-citation>
      
Roque, B. M., Salwen, J. K., Kinley, R., and Kebreab, E.:
Inclusion of Asparagopsis armata in lactating dairy cows' diet reduces enteric methane emission by over 50 percent, J. Clean. Prod., 234, 132–138, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib64"><label>Ross et al.(2022)Ross, Tarbuck, and Macreadie</label><mixed-citation>
      
Ross, F., Tarbuck, P., and Macreadie, P. I.:
Seaweed afforestation at large-scales exclusively for carbon sequestration: Critical assessment of risks, viability and the state of knowledge, Front. Mar. Sci., 9, 1015612, <a href="https://doi.org/10.3389/fmars.2022.1015612" target="_blank">https://doi.org/10.3389/fmars.2022.1015612</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib65"><label>Sato et al.(2023)Sato, Fujiwara, and Endo</label><mixed-citation>
      
Sato, Y., Fujiwara, T., and Endo, H.:
Density regulation of aquaculture production and its effects on commercial profit and quality as food in the cosmopolitan edible seaweed Undaria pinnatifida, Front. Mar. Sci., 10, <a href="https://doi.org/10.3389/fmars.2023.1085054" target="_blank">https://doi.org/10.3389/fmars.2023.1085054</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib66"><label>Sato et al.(2025)Sato, Saito, Inomata, Tanaka, and Nishihara</label><mixed-citation>
      
Sato, Y., Saito, D., Inomata, E., Tanaka, A., and Nishihara, G. N.:
Carbon and nitrogen contents depends on macroalgal species, their tissue section, and development stage, Phycol. Res., 74, <a href="https://doi.org/10.1111/pre.70007" target="_blank">https://doi.org/10.1111/pre.70007</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib67"><label>Sharma et al.(2018)Sharma, Neves, Funderud, Mydland, Øverland, and Horn</label><mixed-citation>
      
Sharma, S., Neves, L., Funderud, J., Mydland, L. T., Øverland, M., and Horn, S. J.:
Seasonal and depth variations in the chemical composition of cultivated <i>Saccharina latissima</i>, Algal Rese., 32, 107–112, <a href="https://doi.org/10.1016/j.algal.2018.03.012" target="_blank">https://doi.org/10.1016/j.algal.2018.03.012</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib68"><label>Sheppard et al.(2023)Sheppard, Hurd, Britton, Reed, and Bach</label><mixed-citation>
      
Sheppard, E. J., Hurd, C. L., Britton, D. D., Reed, D. C., and Bach, L. T.:
Seaweed biogeochemistry: Global assessment of C : N and C : P ratios and implications for ocean afforestation, J. Phycol., 59, 879–892, <a href="https://doi.org/10.1111/jpy.13381" target="_blank">https://doi.org/10.1111/jpy.13381</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib69"><label>Tagliabue et al.(2016)Tagliabue, Aumont, DeAth, Dunne, Dutkiewicz, Galbraith, Misumi, Moore, Ridgwell, Sherman, Stock, Vichi, Völker, and Yool</label><mixed-citation>
      
Tagliabue, A., Aumont, O., DeAth, R., Dunne, J. P., Dutkiewicz, S., Galbraith, E., Misumi, K., Moore, J. K., Ridgwell, A., Sherman, E., Stock, C., Vichi, M., Völker, C., and Yool, A.:
How well do global ocean biogeochemistry models simulate dissolved iron distributions?, Global Biogeochem. Cy., 30, 149–174, <a href="https://doi.org/10.1002/2015GB005289" target="_blank">https://doi.org/10.1002/2015GB005289</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib70"><label>Tagliabue et al.(2023)Tagliabue, Twining, Barrier, Maury, Berger, and Bopp</label><mixed-citation>
      
Tagliabue, A., Twining, B. S., Barrier, N., Maury, O., Berger, M., and Bopp, L.:
Ocean iron fertilization may amplify climate change pressures on marine animal biomass for limited climate benefit, Glob. Change Biol., 29, 5250–5260, <a href="https://doi.org/10.1111/gcb.16854" target="_blank">https://doi.org/10.1111/gcb.16854</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib71"><label>Tullberg et al.(2022)Tullberg, Nguyen, and Wang</label><mixed-citation>
      
Tullberg, R. M., Nguyen, H. P., and Wang, C. M.:
Review of the Status and Developments in Seaweed Farming Infrastructure, J. Mar. Sci. Eng., 10, <a href="https://doi.org/10.3390/jmse10101447" target="_blank">https://doi.org/10.3390/jmse10101447</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib72"><label>Valderrama et al.(2013)Valderrama, Cai, Hishamunda, and Ridler</label><mixed-citation>
      
Valderrama, D., Cai, J., Hishamunda, N., and Ridler, N.:
Social and economic dimensions of carrageenan seaweed farming, Fisheries and aquaculture technical paper no. 580, Food and Agriculture Organization of the United Nations (FAO), Rome, Italy, <a href="https://www.fao.org/4/i3344e/i3344e.pdf" target="_blank"/> (last access: 29 October 2025), 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib73"><label>Vancoppenolle et al.(2023)</label><mixed-citation>
      
Vancoppenolle, M., Rousset, C., Blockley, E., Aksenov, Y., Feltham, D., Fichefet, T., Garric, G., Guémas, V., Iovino, D., Keeley, S., Madec, G., Massonnet, F., Ridley, J., Schroeder, D., and Tietsche, S.:
Sea Ice modelling Integrated Initiative (SI<sup>3</sup>) – The NEMO sea ice engine, Zenodo [software], <a href="https://doi.org/10.5281/zenodo.7534900" target="_blank">https://doi.org/10.5281/zenodo.7534900</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib74"><label>Van Der Molen et al.(2018)Van Der Molen, Ruardij, Mooney, Kerrison, O'Connor, Gorman, Timmermans, Wright, Kelly, Hughes, and Capuzzo</label><mixed-citation>
      
van der Molen, J., Ruardij, P., Mooney, K., Kerrison, P., O'Connor, N. E., Gorman, E., Timmermans, K., Wright, S., Kelly, M., Hughes, A. D., and Capuzzo, E.:
Modelling potential production of macroalgae farms in UK and Dutch coastal waters, Biogeosciences, 15, 1123–1147, <a href="https://doi.org/10.5194/bg-15-1123-2018" target="_blank">https://doi.org/10.5194/bg-15-1123-2018</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib75"><label>Veenhof et al.(2024)Veenhof, Burrows, Hughes, Michalek, Ross, Thomson, Fedenko, and Stanley</label><mixed-citation>
      
Veenhof, R. J., Burrows, M. T., Hughes, A. D., Michalek, K., Ross, M. E., Thomson, A. I., Fedenko, J., and Stanley, M. S.:
Sustainable seaweed aquaculture and climate change in the North Atlantic: challenges and opportunities, Front. Mar. Sci., 11, 1483330, <a href="https://doi.org/10.3389/fmars.2024.1483330" target="_blank">https://doi.org/10.3389/fmars.2024.1483330</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib76"><label>Wang et al.(2019)Wang, Hu, Barnes, Mitchum, Lapointe, and Montoya</label><mixed-citation>
      
Wang, M., Hu, C., Barnes, B. B., Mitchum, G., Lapointe, B., and Montoya, J. P.:
The great Atlantic <i>Sargassum</i> belt, Science, 365, 83–87, <a href="https://doi.org/10.1126/science.aaw7912" target="_blank">https://doi.org/10.1126/science.aaw7912</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib77"><label>Wang et al.(2020)Wang, He, Ma, Huan, Wang, Xia, and Wang</label><mixed-citation>
      
Wang, X., He, L., Ma, Y., Huan, L., Wang, Y., Xia, B., and Wang, G.:
Economically important red algae resources along the Chinese coast: History, status, and prospects for their utilization, Algal Rese., 46, 101817, <a href="https://doi.org/10.1016/j.algal.2020.101817" target="_blank">https://doi.org/10.1016/j.algal.2020.101817</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib78"><label>Wanninkhof(2014)</label><mixed-citation>
      
Wanninkhof, R.:
Relationship between wind speed and gas exchange over the ocean revisited, Limnol. Oceanogr.-Meth., 12, 351–362, <a href="https://doi.org/10.4319/lom.2014.12.351" target="_blank">https://doi.org/10.4319/lom.2014.12.351</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib79"><label>Wu et al.(2023)Wu, Keller, and Oschlies</label><mixed-citation>
      
Wu, J., Keller, D. P., and Oschlies, A.:
Carbon dioxide removal via macroalgae open-ocean mariculture and sinking: an Earth system modeling study, Earth Syst. Dynam., 14, 185–221, <a href="https://doi.org/10.5194/esd-14-185-2023" target="_blank">https://doi.org/10.5194/esd-14-185-2023</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib80"><label>Wu et al.(2025)Wu, Yao, Keller, and Oschlies</label><mixed-citation>
      
Wu, J., Yao, W., Keller, D. P., and Oschlies, A.:
Nearshore Macroalgae Cultivation for Carbon Sequestration by Biomass Harvesting: Evaluating Potential and Impacts With an Earth System Model, Geophys. Res. Lett., 52, e2025GL116774, <a href="https://doi.org/10.1029/2025GL116774" target="_blank">https://doi.org/10.1029/2025GL116774</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib81"><label>Yamamoto et al.(2017)Yamamoto, Kato, Kanayama, Nakase, and Tsutsumi</label><mixed-citation>
      
Yamamoto, M., Kato, T., Kanayama, S., Nakase, K., and Tsutsumi, N.:
Effectiveness of Iron Fertilization for Seaweed Bed Restoration in Coastal Areas, J. Water Environ. Tech., 15, 186–197, <a href="https://doi.org/10.2965/jwet.16-080" target="_blank">https://doi.org/10.2965/jwet.16-080</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib82"><label>Yool et al.(2013)Yool, Popova, and Anderson</label><mixed-citation>
      
Yool, A., Popova, E. E., and Anderson, T. R.:
MEDUSA-2.0: an intermediate complexity biogeochemical model of the marine carbon cycle for climate change and ocean acidification studies, Geosci. Model Dev., 6, 1767–1811, <a href="https://doi.org/10.5194/gmd-6-1767-2013" target="_blank">https://doi.org/10.5194/gmd-6-1767-2013</a>, 2013.


    </mixed-citation></ref-html>
<ref-html id="bib1.bib83"><label>Yool et al.(2021)Yool, Palmiéri, Jones, de Mora, Kuhlbrodt, Popova, Nurser, Hirschi, Blaker, Coward, Blockley, and Sellar</label><mixed-citation>
      
Yool, A., Palmiéri, J., Jones, C. G., de Mora, L., Kuhlbrodt, T., Popova, E. E., Nurser, A. J. G., Hirschi, J., Blaker, A. T., Coward, A. C., Blockley, E. W., and Sellar, A. A.:
Evaluating the physical and biogeochemical state of the global ocean component of UKESM1 in CMIP6 historical simulations, Geosci. Model Dev., 14, 3437–3472, <a href="https://doi.org/10.5194/gmd-14-3437-2021" target="_blank">https://doi.org/10.5194/gmd-14-3437-2021</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib84"><label>Zhang et al.(2012)Zhang, Fang, Wang et al.</label><mixed-citation>
      
Zhang, J., Fang, J., Wang, W., Du, M., Gao, Y., and Zhang, M.: Growth and loss of mariculture kelp <i>Saccharina japonica</i> in Sungo Bay, China, J. Appl. Phycol., 24, 1209–1216, <a href="https://doi.org/10.1007/s10811-011-9762-4" target="_blank">https://doi.org/10.1007/s10811-011-9762-4</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib85"><label>Zhang et al.(2016)Zhang, Wu, Ren, and Lin</label><mixed-citation>
      
Zhang, J., Wu, W., Ren, J. S., and Lin, F.:
A model for the growth of mariculture kelp <i>Saccharina japonica</i> in Sanggou Bay, China, Aquacult. Env. Interac., 8, 273–283, <a href="https://doi.org/10.3354/aei00171" target="_blank">https://doi.org/10.3354/aei00171</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib86"><label>Zollmann et al.(2023)Zollmann, Liberzon, Palatnik et al.</label><mixed-citation>
      
Zollmann, M., Liberzon, A., Palatnik, R. R., Zilberman, D., and Golberg, A.: Effects of season, depth and pre-cultivation fertilizing on Ulva growth dynamics offshore the Eastern Mediterranean Sea, Sci. Rep., 13, 14784, <a href="https://doi.org/10.1038/s41598-023-41605-4" target="_blank">https://doi.org/10.1038/s41598-023-41605-4</a>, 2023.

    </mixed-citation></ref-html>--></article>
