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  <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-23-6229-2026</article-id><title-group><article-title>Sinking particle fluxes and biological carbon pump efficiency in the Labrador Sea during a <italic>Phaeocystis</italic> bloom decline</article-title><alt-title>Sinking particle fluxes and biological carbon pump efficiency</alt-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1 aff2">
          <name><surname>Roca-Martí</surname><given-names>Montserrat</given-names></name>
          <email>montserrat.roca.marti@uab.cat</email>
        <ext-link>https://orcid.org/0000-0002-4719-9358</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Healey</surname><given-names>Madeline</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>McBride</surname><given-names>Colleen E.</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3 aff4">
          <name><surname>Sipler</surname><given-names>Rachel</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff5">
          <name><surname>Devred</surname><given-names>Emmanuel</given-names></name>
          
        <ext-link>https://orcid.org/0000-0001-9446-0005</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff6">
          <name><surname>Cisternas-Novoa</surname><given-names>Carolina</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff7">
          <name><surname>Romanelli</surname><given-names>Elisa</given-names></name>
          
        <ext-link>https://orcid.org/0000-0001-7989-9108</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1 aff8">
          <name><surname>Ohashi</surname><given-names>Kyoko</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Kienast</surname><given-names>Stephanie S.</given-names></name>
          
        <ext-link>https://orcid.org/0000-0003-3867-4919</ext-link></contrib>
        <aff id="aff1"><label>1</label><institution>Department of Oceanography, Dalhousie University, Halifax, NS, Canada</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Physics Department, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Department of Ocean Sciences, Memorial University of Newfoundland, St. John's, NL, Canada</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>Bigelow Laboratory for Ocean Sciences, East Boothbay, ME, USA</institution>
        </aff>
        <aff id="aff5"><label>5</label><institution>Fisheries and Oceans Canada, Bedford Institute of Oceanography, Dartmouth, NS, Canada</institution>
        </aff>
        <aff id="aff6"><label>6</label><institution>Ocean Sciences Centre, Memorial University of Newfoundland, St. John's, NL, Canada</institution>
        </aff>
        <aff id="aff7"><label>7</label><institution>Institute of Environmental Engineering, Department of Civil, Environmental and Geomatic Engineering, ETH Zurich, Zurich, Switzerland</institution>
        </aff>
        <aff id="aff8"><label>8</label><institution>Fisheries and Oceans Canada, Institute of Ocean Sciences, Sidney, BC, Canada</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Montserrat Roca-Martí (montserrat.roca.marti@uab.cat)</corresp></author-notes><pub-date><day>11</day><month>September</month><year>2026</year></pub-date>
      
      <volume>23</volume>
      <issue>17</issue>
      <fpage>6229</fpage><lpage>6248</lpage>
      <history>
        <date date-type="received"><day>29</day><month>July</month><year>2025</year></date>
           <date date-type="rev-request"><day>25</day><month>August</month><year>2025</year></date>
           <date date-type="rev-recd"><day>19</day><month>June</month><year>2026</year></date>
           <date date-type="accepted"><day>14</day><month>August</month><year>2026</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2026 Montserrat Roca-Martí 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/23/6229/2026/bg-23-6229-2026.html">This article is available from https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026.pdf</self-uri>
      <abstract><title>Abstract</title>

      <p id="d2e208">The Labrador Sea is a key region for carbon dioxide uptake characterized by deep mixing during winter that supplies nutrients to the upper water column and fuels extensive phytoplankton blooms in spring. Yet, the efficiency by which organic carbon is exported from surface waters during these blooms, as well as their contribution to carbon sequestration, remain poorly constrained. Here, we present an unprecedented number of measurements of sinking export fluxes (particulate organic carbon, POC; and biogenic silica, bSi) collected in the central Labrador Sea during a 2-week-long process study that observed the decline of a historically large <italic>Phaeocystis</italic> bloom in spring 2022. This <italic>Phaeocystis</italic> bloom was unusually large and highly productive, extending over more than half of the Labrador Sea for 6 weeks. During the late stages of the bloom, we found that POC fluxes from the base of the euphotic zone to 500 m were variable but overall moderate to high (average of 8 <inline-formula><mml:math id="M1" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5 mmol C m<sup>−2</sup> d<sup>−1</sup>). Nevertheless, evidence of shallow POC flux remineralization combined with the fact that POC fluxes in the bloom were not higher than in a region sampled outside of the bloom (average of 13 <inline-formula><mml:math id="M4" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3 mmol C m<sup>−2</sup> d<sup>−1</sup>) suggested a limited role of <italic>Phaeocystis</italic> in carbon export. Large (<inline-formula><mml:math id="M7" display="inline"><mml:mo lspace="0mm">&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M8" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m) particles collected using large volume pumps presented relatively low bSi <inline-formula><mml:math id="M9" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC ratios and, therefore, diatoms did not appear to have an important ballasting role of <italic>Phaeocystis</italic>-derived material. Using in situ net primary production (NPP) rates, we determined that 2 weeks after the bloom peak, only 6 % of NPP was exported to 100 m below the euphotic zone. Three weeks after the peak, this value increased to 30 %, reflecting a decline in NPP while POC fluxes remained relatively constant. However, when using satellite-derived NPP integrated from the bloom peak until its end, the overall biological carbon pump (BCP) efficiency was 6 %, indicating that this <italic>Phaeocystis</italic> bloom represented a low-efficiency export system. We stress the importance of multiple observations of both NPP and POC export along the bloom period for estimating meaningful BCP efficiencies. The results presented in this study provide a foundation for comparisons with other datasets collected during this ship-based process study and autonomous platforms present in the area during and beyond this study. These future efforts will provide the opportunity to increase the observational period and further elucidate the mechanisms leading to the low BCP efficiency found during the decline of this <italic>Phaeocystis</italic> bloom in the Labrador Sea.</p>
  </abstract>
    
<funding-group>
<award-group id="gs1">
<funding-source>Ocean Frontier Institute</funding-source>
<award-id>2021-BP-00109</award-id>
<award-id>LCF/BQ/PI24/12040022</award-id>
<award-id>RYC2023-045355-I</award-id>
<award-id>2021 SGR-640</award-id>
</award-group>
<award-group id="gs2">
<funding-source>Natural Sciences and Engineering Research Council of Canada</funding-source>
<award-id>NA</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="d2e324">The biological carbon pump (BCP) encapsulates a set of processes that remove carbon dioxide (CO<sub>2</sub>) from the atmosphere and sequester it in the deep ocean (Passow and Weber, 2025; Volk and Hoffert, 1985). In the surface ocean, phytoplankton produce organic matter via photosynthesis, and a fraction of this organic matter is subsequently transferred to depth through sinking and other processes. Of the material exported below the euphotic zone, most is remineralized in the ocean interior, while only a small fraction reaches depths where CO<sub>2</sub> can be stored for several centuries or longer. A very small fraction of exported carbon is buried in seafloor sediments and thus removed from the atmosphere on geological timescales. While it is well understood that the BCP plays a large role in climate regulation, there is low confidence in the magnitude and even the sign of predicted near-future changes in carbon export fluxes and how these will in turn affect atmospheric CO<sub>2</sub> levels (Henson et al., 2022). Better quantification of particulate matter fluxes in the ocean and of the key parameters controlling them is thus urgently needed.</p>
      <p id="d2e354">Indeed, the various processes transforming and redistributing carbon between the surface and the deep ocean remain poorly quantified. Assessments of carbon budgets below the euphotic zone typically have found that organic carbon sources and sinks are not balanced (Baltar et al., 2009; Boyd et al., 1999; Burd et al., 2010; Reinthaler et al., 2006; Steinberg et al., 2008). This mismatch has been attributed to methodological constraints, the choice of parameters used to estimate budget terms, the exclusion of important midwater processes, and the spatial and temporal scales over which measurements are integrated (Baumas et al., 2023; Giering et al., 2014; Stephens and Roca-Martí et al., 2025).</p>
      <p id="d2e357">Major uncertainties in the magnitude of biologically driven carbon ocean uptake and storage also lie in our fragmented understanding of the different pathways by which the BCP exports particulate and dissolved organic carbon (POC and DOC, respectively) from surface waters to the deep ocean. Advances in remote sensing, in situ imagery, and the development of autonomous platforms (BGC-Argo floats and gliders) over the last 2 decades have revealed that the BCP includes six major pathways for exporting organic carbon from the surface ocean to the deep ocean: gravitational export (sinking), three pumps driven by physical processes (mixed layer, eddy subduction and large-scale subduction), and two pumps driven by vertical migrations ofzooplankton and larger animals (Boyd et al., 2019; Claustre et al., 2021). Process studies and long-term observations of the BCP, combining multiple approaches both ship- and autonomous-based, have been shown to be crucial for better constraining ocean carbon budgets (Stephens and Roca-Martí et al., 2025).</p>
      <p id="d2e360">The North Atlantic is responsible for a significant fraction of the global carbon export (<inline-formula><mml:math id="M13" display="inline"><mml:mo lspace="0mm">∼</mml:mo></mml:math></inline-formula> 15 %), driven by a complex set of BCP processes (Sanders et al., 2014). The subpolar North Atlantic, specifically the Labrador Sea, is an important region for CO<sub>2</sub> uptake (Arruda et al., 2024) characterized by deep vertical mixing during winter that supplies nutrients to the upper water column and fuels extensive phytoplankton blooms in spring after the water column has stratified (Tesdal et al., 2022). Yet, the efficiency by which organic carbon is exported from surface waters during these blooms as well as their contribution to carbon sequestration remain poorly constrained, partly because of the complex physical processes in the region (Baker et al., 2022) and the very limited POC flux observations made so far (e.g., Lemaitre et al., 2018). In addition, impacts of global warming in the physical conditions of the Labrador Sea have already been reported, including freshening and weaker winter convection (Yashayaev, 2024), which add uncertainty to the future state of the BCP and its downstream effects on climate and marine ecosystems. Therefore, our new observations will provide an important baseline for constraining POC export in the Labrador Sea under rapidly changing hydrographic and ecological conditions.</p>
      <p id="d2e380">The phytoplankton assemblages in the Labrador Sea are typically dominated by diatoms, <italic>Phaeocystis</italic> spp. or mixed populations (Devred et al., 2024). Observations of <italic>Phaeocystis</italic> blooms were previously restricted to shelf and slope regimes, but unprecedented large <italic>Phaeocystis</italic> blooms have been reported in the Labrador basin in recent years (in 2015 and 2022; Devred et al., 2025). Here, we present measurements of net primary production (NPP) and sinking export fluxes (POC and biogenic silica, bSi) in the central Labrador Sea during the decline of a historically large <italic>Phaeocystis</italic> bloom in spring 2022. The novelty of our ship-based process study (Biological Carbon Export in the Labrador Sea, BELAS-1) lies in the high temporal resolution of in situ measurements collected over 2 weeks which shed light on the efficiency of the gravitational sinking pump of <italic>Phaeocystis</italic> blooms.</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Methods</title>
      <p id="d2e406">Samples were collected in the Labrador Sea from 19 May to 2 June 2022 during the BELAS-1 expedition (CE22009, RV <italic>Celtic Explorer</italic>). Three regions were targeted (Fig. 1): a grid of nine stations (hereafter “Eastern Grid”, <inline-formula><mml:math id="M15" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 1075 km<sup>2</sup>), where an extensive <italic>Phaeocystis</italic> <italic>pouchetii</italic> bloom was in decline (Devred et al., 2025); two stations located outside of the major <italic>Phaeocystis</italic> bloom to the southwest of the Eastern Grid (hereafter “Central”); and one station located in the bloom area between the Eastern Grid and the central stations which was sampled on two consecutive days (hereafter “Station 28-1” and “Station 28-2”). In the Eastern Grid, six stations were sampled from 20   to 25 May (hereafter “East 1”) before Station 28 and the central stations were occupied, while three stations were sampled after a major storm at the end of the expedition from 30 May to 2 June (hereafter “East 2”). East 1 was characterized by higher <italic>Phaeocystis</italic> biomass levels and primary production rates compared to East 2, which represented post-bloom conditions (Figs. 2 and 3).</p>

      <fig id="F1" specific-use="star"><label>Figure 1</label><caption><p id="d2e443">Map of the stations sampled during the BELAS-1 expedition (CE22009, RV <italic>Celtic Explorer</italic>) in different regions: the Eastern Grid or “East” (East 1: 20  to 25 May; East 2: 30 May to 2 June), Station 28 (St. 28), and “Central”. All stations were sampled for total <sup>234</sup>Th in seawater. Red dots denote stations where in situ large volume pumps (LVP) were deployed. Yellow squares denote stations sampled for net primary production (NPP).</p></caption>
        <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f01.png"/>

      </fig>

      <fig id="F2" specific-use="star"><label>Figure 2</label><caption><p id="d2e466">Satellite-derived chlorophyll <inline-formula><mml:math id="M18" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> (chl <inline-formula><mml:math id="M19" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula>) concentrations (Devred et al., 2025) at the time of sampling of East 1 stations (left panel), Station 28 and central stations (middle panel), and East 2 stations (right panel).</p></caption>
        <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f02.png"/>

      </fig>

      <fig id="F3" specific-use="star"><label>Figure 3</label><caption><p id="d2e492">Profiles of chlorophyll <inline-formula><mml:math id="M20" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> from the CTD (CTD chl <inline-formula><mml:math id="M21" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula>) and net primary production (NPP, absolute uptake) in the upper 200 m of the water column in East 1 (green), Central (dark blue) and East 2 (yellow) stations. The base of the primary production zone (PPZ, Owens et al., 2015) shown with a grey solid line is used to operationally define the base of the euphotic zone. The base of the mixed layer (MLD) is shown with a blue dashed line.</p></caption>
        <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f03.png"/>

      </fig>

      <p id="d2e515">The mixed layer depth (MLD), defined as the depth where potential density exceeds the density at 10 m depth by 0.03 kg m<sup>−3</sup> (de Boyer Montégut et al., 2004), ranged from 22 to 66 m. Stratification, defined as the squared buoyancy frequency, ranged from 5 to 88 m and was on average 20 % deeper than the MLD. The base of the primary production zone (PPZ), defined as the depth at which fluorescence was 10 % of its maximal value (Owens et al., 2015), ranged from 79 to 176 m and was used to operationally define the base of the euphotic zone. Fluorescence was obtained from an optical package mounted on the conductivity-temperature-depth (CTD) rosette system. The CTD was not equipped with a photosynthetic active radiation (PAR) sensor during the BELAS-1 expedition.</p>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Net primary production</title>
      <p id="d2e537">Water was collected from six depths using a CTD-rosette fitted with 10 L Niskin bottles. While collections were dispersed throughout the upper 200 m of the water column, some sampling depths were selected based on key hydrographic features, including the surface and chlorophyll <inline-formula><mml:math id="M23" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> (chl <inline-formula><mml:math id="M24" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula>) maximum (exact depths can be found in Table S1 in the Supplement). Water was transferred from the Niskin to the incubation bottles using acid-rinsed (10 % HCl followed by a minimum of four rinses with type 1 (<inline-formula><mml:math id="M25" display="inline"><mml:mo lspace="0mm">≥</mml:mo></mml:math></inline-formula> 18.2 M<inline-formula><mml:math id="M26" display="inline"><mml:mi mathvariant="normal">Ω</mml:mi></mml:math></inline-formula> cm) reverse osmosis water) silicone tubing fitted with 150 <inline-formula><mml:math id="M27" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m Nitex mesh and covered in electrical tape to exclude light from samples collected at depth. Incubations were performed in triplicate in acid-washed 1 L incubation bottles filled to the bottle neck (approximate volume of 1.2 L). Bottles were amended with H<sup>13</sup>CO<inline-formula><mml:math id="M29" display="inline"><mml:mrow><mml:msubsup><mml:mi/><mml:mn mathvariant="normal">3</mml:mn><mml:mo>-</mml:mo></mml:msubsup></mml:mrow></mml:math></inline-formula> (99 %; Cambridge Isotope Laboratories) at an approximate addition of 330 <inline-formula><mml:math id="M30" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol C L<sup>−1</sup>. For stable isotope tracer studies, we aim for target additions of 10 %, and the average atom percent enrichment for the current study was 13.8 <inline-formula><mml:math id="M32" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5 %.</p>
      <p id="d2e625">Once the isotope was added, bottles were placed in flow-through deck incubators for 24 h with natural daylight. Incubators were covered at night to prevent the unintended impacts of deck lighting. The incubation bottles were covered with screen bags of varying thicknesses to mimic the light availability at the depth where they were collected. For deep samples, bottles were covered with foil and then multiple layers of electrical tape to eliminate light intrusion. Temperature and light in the incubators were monitored using HOBO TidbiT v2 water temperature data loggers (Onset Computer Corporation).</p>
      <p id="d2e628">The experiments were terminated by filtration onto pre-combusted (450 °C for 4 h) GF-75 filters with a nominal pore size of 0.3 <inline-formula><mml:math id="M33" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m. Filters were placed into 2 mL cryovials and frozen at <inline-formula><mml:math id="M34" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>20 °C until analysis at Bigelow Laboratory for Ocean Sciences using a CosTech ECS 4010/Thermo DELTA V Advantage Isotope Ratio Mass Spectrometer. Absolute carbon uptake rates were calculated according to Hama et al. (1983) using the following equation:

            <disp-formula id="Ch1.E1" content-type="numbered"><label>1</label><mml:math id="M35" display="block"><mml:mrow><mml:mi mathvariant="italic">ρ</mml:mi><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mtext>PC at</mml:mtext><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mi mathvariant="italic">%</mml:mi><mml:mi>x</mml:mi><mml:mi>s</mml:mi></mml:mrow><mml:mrow><mml:mtext>DIC at</mml:mtext><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="italic">%</mml:mi><mml:mi>x</mml:mi><mml:mi>s</mml:mi><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mo>×</mml:mo><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mtext>Time</mml:mtext></mml:mrow></mml:mfrac></mml:mstyle><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>×</mml:mo><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>[</mml:mo><mml:mtext>PC</mml:mtext><mml:mo>]</mml:mo><mml:mo>,</mml:mo></mml:mrow></mml:math></disp-formula>

          where <inline-formula><mml:math id="M36" display="inline"><mml:mi mathvariant="italic">ρ</mml:mi></mml:math></inline-formula> is the absolute uptake rate and at %<inline-formula><mml:math id="M37" display="inline"><mml:mrow><mml:mi>x</mml:mi><mml:mi>s</mml:mi></mml:mrow></mml:math></inline-formula> is the atom % excess. For particulate carbon (PC), atom % excess  refers to the percentage of <sup>13</sup>C enrichment at the end of the incubation minus the percentage of <sup>13</sup>C at the beginning of the incubation. For dissolved inorganic carbon (DIC), atom % excess refers to the proportional change in <sup>13</sup>C available in the water when the H<sup>13</sup>CO<inline-formula><mml:math id="M42" display="inline"><mml:mrow><mml:msubsup><mml:mi/><mml:mn mathvariant="normal">3</mml:mn><mml:mo>-</mml:mo></mml:msubsup></mml:mrow></mml:math></inline-formula> label is added. Ambient HCO<inline-formula><mml:math id="M43" display="inline"><mml:mrow><mml:msubsup><mml:mi/><mml:mn mathvariant="normal">3</mml:mn><mml:mo>-</mml:mo></mml:msubsup></mml:mrow></mml:math></inline-formula> concentrations were estimated based on the salinity of each sample (Parsons et al., 1984). Absolute uptake rate represents how much <sup>13</sup>C was taken up by the target community over the incubation period and is presented as units of carbon per volume/area per time (i.e., mmol C m<sup>−3</sup> d<sup>−1</sup>).</p>
      <p id="d2e810">Depth-integrated NPP was calculated using a trapezoidal integration (Knap et al., 1996) up to the base of the PPZ from measured absolute uptake rates. In cases where the base of the PPZ exceeded the deepest sampling depth, the absolute uptake rate from the deepest sampling depth was used for any depths up to and including the base of the PPZ. Due to a limited number of sampling depths, NPP could not be integrated at the central stations.</p>
      <p id="d2e814">Satellite NPP was calculated following the methods described in Devred et al. (2025). Briefly, the satellite product is based on daily 4 km MODIS-Aqua level-3 fields of remote sensing reflectance, sea surface temperature (SST), and PAR. Chlorophyll <inline-formula><mml:math id="M47" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> was derived using a regionally-tuned version of the MODIS OC3M algorithm, and gaps in chl <inline-formula><mml:math id="M48" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> and SST were filled using Data Interpolating Empirical Orthogonal Functions (DINEOF).</p>
</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>Total <sup>234</sup>Th and <sup>238</sup>U in seawater</title>
      <p id="d2e858">Water column samples were taken in the upper 500 m of all stations (<inline-formula><mml:math id="M51" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">12</mml:mn></mml:mrow></mml:math></inline-formula>, Figs. 1, 4) using a CTD-rosette at 15 discrete depths. Unfiltered seawater samples (2 L) were processed on board and analyzed for the activity of total (dissolved <inline-formula><mml:math id="M52" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> particulate) <sup>234</sup>Th following the method described in Clevenger et al. (2021). Samples were immediately acidified after collection and spiked with a yield monitor (<sup>230</sup>Th, 25 disintegrations per minute, dpm, per sample). After a minimum 6 h equilibration time, sample pH was increased to <inline-formula><mml:math id="M55" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 8.5 using ammonium hydroxide, and reagents were added to form a manganese oxide precipitate for scavenging of Th. Samples were allowed to stand for at least 8 h, filtered onto 25 mm diameter quartz microfiber filters (QMA), dried and mounted for beta counting. Samples were counted using three low-level Risø beta multicounters (5 detectors each) until the uncertainty was typically <inline-formula><mml:math id="M56" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 3 %. Beta counters were calibrated using two sets of five deep samples (1500 m) from two stations. At least 5 months after collection (<inline-formula><mml:math id="M57" display="inline"><mml:mo lspace="0mm">&gt;</mml:mo></mml:math></inline-formula> 6 <sup>234</sup>Th half-lives, where the half-life <inline-formula><mml:math id="M59" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 24.1 d), samples were recounted at Dalhousie University to determine the non-<sup>234</sup>Th beta activity stemming from other radionuclides included in the precipitate, which was subtracted from the first count. The net counting rate was corrected for <sup>234</sup>Th decay and ingrowth from <sup>238</sup>U, counting efficiency and chemical recovery. The chemical recovery of <sup>230</sup>Th was determined by using a Thermo Scientific iCAP quadrupole inductively coupled plasma mass spectrometer (ICP-MS) following the procedure detailed in Clevenger et al. (2021). Recoveries averaged 83 <inline-formula><mml:math id="M64" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 10 % (<inline-formula><mml:math id="M65" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">218</mml:mn></mml:mrow></mml:math></inline-formula>). The uranium-238 (<sup>238</sup>U) activity was derived from salinity (Owens et al., 2011).</p>

      <fig id="F4" specific-use="star"><label>Figure 4</label><caption><p id="d2e1003">Profiles of total <sup>234</sup>Th and <sup>238</sup>U in the upper 500 m of the water column in East 1 stations (green), Station 28 (light blue), central stations (dark blue) and East 2 stations (yellow). The base of the primary production zone (PPZ) is shown with a grey solid line and the base of the mixed layer (MLD) is shown with a blue dashed line.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f04.png"/>

        </fig>

</sec>
<sec id="Ch1.S2.SS3">
  <label>2.3</label><title>Particles collected using large volume pumps and marine snow catchers</title>
      <p id="d2e1038">Size-fractionated particles were collected using battery-powered in situ large volume pumps (McLane Research Laboratories, Inc.) equipped with a three-tier 142 mm diameter filter holder (MULVFS style; Bishop et al., 2012; Lam et al., 2015). Five pumps were deployed at depths between 40 and 490 m at six stations (Figs. 1, 5). The pump deployment depths were chosen after examining the fluorescence profile from a CTD cast conducted shortly prior to the pump deployment. One pump was placed close to the base of the PPZ and another pump at 100 m below the PPZ base. Pumps were programmed to sample for 2.5 to 3.0 h at a starting flow rate of 6 L min<sup>−1</sup> and pumped an average of 460 L each. During each pump deployment, pressure sensors (RBR duet 3) were attached to selected pumps to confirm deployment depths.</p>

      <fig id="F5" specific-use="star"><label>Figure 5</label><caption><p id="d2e1055">Profiles of POC <inline-formula><mml:math id="M70" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>234</sup>Th (C <inline-formula><mml:math id="M72" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th) and bSi <inline-formula><mml:math id="M73" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>234</sup>Th (bSi <inline-formula><mml:math id="M75" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th) ratios in <inline-formula><mml:math id="M76" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula>51 <inline-formula><mml:math id="M77" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m particles across regions. Black symbols denote average C <inline-formula><mml:math id="M78" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios (all stations) and average bSi <inline-formula><mml:math id="M79" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios (all stations except East 2) for the following depth horizons: 115–185, 210–240, 260–340, 465–490 m. Empty symbols denote samples collected within the primary production zone. Analytical uncertainties are represented by horizontal bars and uncertainties less than <inline-formula><mml:math id="M80" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 3 <inline-formula><mml:math id="M81" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> are not visible against filled symbols. POC <inline-formula><mml:math id="M83" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> particulate organic carbon, bSi <inline-formula><mml:math id="M84" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> biogenic silica.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f05.png"/>

        </fig>

      <p id="d2e1182">The filter holders contained three filters: two Nitex screens (335 and 51 <inline-formula><mml:math id="M85" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m nominal pore size, acid-leached prior to the cruise) above a pre-combusted QMA filter (<inline-formula><mml:math id="M86" display="inline"><mml:mo lspace="0mm">∼</mml:mo></mml:math></inline-formula> 1 <inline-formula><mml:math id="M87" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m nominal pore size, Graff et al., 2023) for size fractionation (1–51, 51–335 and <inline-formula><mml:math id="M88" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 335 <inline-formula><mml:math id="M89" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m). A total of 28 samples were collected for each size fraction. Size-fractionated particles were analyzed for <sup>234</sup>Th, particulate organic carbon (POC), and biogenic silica (bSi, only from Nitex screens, i.e., 51–335 and <inline-formula><mml:math id="M91" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 335 <inline-formula><mml:math id="M92" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m size fractions). In every cast, an additional filter holder was mounted on the deepest pump to obtain a seawater process blank (“dipped blank” filters, Lam et al., 2015). Immediately after recovering the pumps, residual water from each filter holder was removed by vacuum to avoid particle loss.</p>
      <p id="d2e1249">Zooplankton that were not part of the passive sinking flux (“swimmers”) and were visible to the naked eye were carefully handpicked from the Nitex screens using forceps and discarded. Particles were gently rinsed off the Nitex screens onto 25-mm-diameter silver (Ag) filters (1.2 <inline-formula><mml:math id="M93" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m nominal pore size) using pre-filtered seawater. QMA filters were subsampled for <sup>234</sup>Th and POC using a circular punch tool. Ag and QMA filters were dried, beta counted at sea for <sup>234</sup>Th activities and recounted <inline-formula><mml:math id="M96" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 5 months later at Dalhousie University. After final counting, Ag filters were split into halves by weight and analyzed for POC and bSi. POC was analyzed using high-temperature combustion (Costech Instruments Elemental Combustion System 4010) after acid fumigation. bSi was analyzed following the NaOH digestion method (40 min at 95 °C) using a UV-Visible Spectrophotometer (Genesys 10S, Thermo Scientific) as in Roca-Martí et al. (2021).</p>
      <p id="d2e1285">The average of all dipped blanks was subtracted from total <sup>234</sup>Th, POC, and bSi measurements. Dipped blanks contributed on average <inline-formula><mml:math id="M98" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 3 % to the total <sup>234</sup>Th measured on the filters, <inline-formula><mml:math id="M100" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 7 % to total bSi, and <inline-formula><mml:math id="M101" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 14 % to total POC (Table S2 in the Supplement). All concentrations were above the limit of detection (3 <inline-formula><mml:math id="M102" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> standard deviation of the dipped blanks, Lam et al., 2018). Analysis of triplicate punches from four QMA filters representing depths from 100 to 500 m yielded a relative standard deviation for POC of 1 %–4 %, indicating a relatively homogeneous particle distribution across the filters (Maiti et al., 2012). POC and bSi data have average uncertainties of 3 % and 6 %, respectively, resulting from the dipped blank correction. Particulate <sup>234</sup>Th have average uncertainties of 5 % resulting from counting and the dipped blank correction.</p>
      <p id="d2e1344">At Station 12 in East 1, particles were also collected using marine snow catchers (MSCs) as an independent method to constrain POC <inline-formula><mml:math id="M104" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>234</sup>Th ratios in sinking material. Three MSCs were deployed approximately 3.5 h prior to the large volume pump deployments at similar depths (165, 265, and 485 m; <inline-formula><mml:math id="M106" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>5 m). MSCs are 1.5-m-tall, 100 L water samplers equipped with a removable base section designed to separate particles according to their sinking velocity under low-turbulence conditions (Giering et al., 2016; Riley et al., 2012). Following MSC retrieval, particles were allowed to settle for 2 h, after which they were classified into three operationally defined fractions: suspended, slow-sinking, and fast-sinking (Romanelli et al., 2023, 2026). Suspended particles, with negligible sinking velocity, were collected from a central tap <inline-formula><mml:math id="M107" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 79 cm from the MSC top. Slow-sinking particles, with velocities <inline-formula><mml:math id="M108" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 18 m d<sup>−1</sup>, were siphoned from the base. Fast-sinking particles, with velocities <inline-formula><mml:math id="M110" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 18 m d<sup>−1</sup>, were trapped in the tray at the bottom of the base section. On average, 2.1 L (of 5.0 L) from the base and 1.4 L (of 2.6 L) from the tray were filtered through 25-mm-diameter Ag filters (0.45 <inline-formula><mml:math id="M112" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m nominal pore size) for the determination of <sup>234</sup>Th and POC in slow- and fast-sinking particles. Blank corrections were applied by subtracting the average of two filter blanks from total <sup>234</sup>Th and POC measurements, and the slow- and fast-sinking particle fractions were subsequently combined. Average uncertainties were 1 % for POC and 17 % for particulate <sup>234</sup>Th, resulting from the blank correction (POC and <sup>234</sup>Th) and counting (<sup>234</sup>Th). The sample from 165 m was excluded from analysis due to a low signal-to-noise ratio in the <sup>234</sup>Th measurements.</p>
</sec>
<sec id="Ch1.S2.SS4">
  <label>2.4</label><title><sup>234</sup>Th, POC and bSi export fluxes</title>
      <p id="d2e1496">Export fluxes of <sup>234</sup>Th were calculated for each water column profile by integrating the disequilibrium between <sup>234</sup>Th and <sup>238</sup>U from the surface to each sampled depth over the upper 500 m of the water column. This model implies steady-state (SS) conditions and neglects physical transport (Roca-Martí and Puigcorbé, 2024; Savoye et al., 2006). We discuss the validity of this SS approach in Sect. 2.5.</p>
      <p id="d2e1526">To estimate POC and bSi export fluxes, steady-state <sup>234</sup>Th fluxes were multiplied with POC <inline-formula><mml:math id="M124" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>234</sup>Th (C <inline-formula><mml:math id="M126" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th) and bSi <inline-formula><mml:math id="M127" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>234</sup>Th (bSi <inline-formula><mml:math id="M129" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th) ratios measured in <inline-formula><mml:math id="M130" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M131" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m particles (i.e., combining the 51–335 and <inline-formula><mml:math id="M132" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 335 <inline-formula><mml:math id="M133" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m size fractions) collected using large volume pumps. Using the <inline-formula><mml:math id="M134" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M135" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m fraction for the calculation of fluxes is consistent with the majority of <sup>234</sup>Th studies (Puigcorbé et al., 2020). Furthermore, this choice is directly supported here by comparing C <inline-formula><mml:math id="M137" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios measured at Station 12 in size-fractionated particles with those measured in sinking particles collected using MSCs (Fig. S1 in the Supplement). Sinking particles from MSCs show C <inline-formula><mml:math id="M138" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios similar to all pump size fractions that are greater than 51 <inline-formula><mml:math id="M139" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m (i.e., 51–335, <inline-formula><mml:math id="M140" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 335, and <inline-formula><mml:math id="M141" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M142" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m; Fig. S1 in the Supplement). On the contrary, C <inline-formula><mml:math id="M143" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios from MSCs are clearly higher than C <inline-formula><mml:math id="M144" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios in pump samples that include particles smaller than 51 <inline-formula><mml:math id="M145" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m (i.e., the 1–51 and <inline-formula><mml:math id="M146" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 1 <inline-formula><mml:math id="M147" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m fractions).</p>
      <p id="d2e1723">Based on the samples collected at five depths across six individual stations, we derived average C <inline-formula><mml:math id="M148" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th and bSi <inline-formula><mml:math id="M149" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios (<inline-formula><mml:math id="M150" display="inline"><mml:mo lspace="0mm">±</mml:mo></mml:math></inline-formula> standard deviation) for several depth horizons below the PPZ base (115–185, 210–240, 260–340, 465–490 m; Fig. 5). Ratios at depths not sampled by pump deployments were interpolated linearly between the mid-points of each depth horizon. For POC, all pump casts were combined to determine cruise-average C <inline-formula><mml:math id="M151" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios because no significant differences were found between East 1, Station 28, Central, and East 2 (one-way ANOVA, <inline-formula><mml:math id="M152" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M153" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 0.05). For bSi, pump casts from East 1, Station 28, and Central were combined (one-way ANOVA, <inline-formula><mml:math id="M154" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M155" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 0.05) to determine average bSi <inline-formula><mml:math id="M156" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios and estimate bSi fluxes from 20  to 29 May. bSi <inline-formula><mml:math id="M157" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios in East 2 were used to estimate bSi fluxes from 30 May to 2 June given the higher bSi <inline-formula><mml:math id="M158" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios measured in East 2 (Kruskal-Wallis, <inline-formula><mml:math id="M159" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M160" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 0.01).</p>
</sec>
<sec id="Ch1.S2.SS5">
  <label>2.5</label><title>Testing the assumptions of our export model</title>
<sec id="Ch1.S2.SS5.SSS1">
  <label>2.5.1</label><title>How valid is the steady state assumption?</title>
      <p id="d2e1836">The validity of the steady state (SS) assumption during a phytoplankton bloom depends on when the sampling occurs with respect to the bloom peak and the duration of the bloom (Ceballos-Romero et al., 2018). Following the approach by Ceballos-Romero et al. (2018), satellite-derived chl <inline-formula><mml:math id="M161" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> concentrations suggest that the peak of maximum production in the Eastern Grid area occurred on 11 May, i.e., 9–22 d before our sampling period (20 May–2 June). Chl <inline-formula><mml:math id="M162" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> decreased quite rapidly with time after the peak (Fig. S2 in the Supplement), similar to a hypothetical bloom scenario considered in Ceballos-Romero et al. (2018) (their Fig. 1b). In such conditions, these authors found that the <sup>234</sup>Th SS model provides accurate flux estimates during a <inline-formula><mml:math id="M164" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 1 week-long sampling period, referred to as “window of success”, which commences 10 d after the peak of the bloom. Comparing the sampling dates and bloom timing of the present study with the findings from Ceballos-Romero et al. (2018) suggests that most of our sampling in East 1 happened within the window of success (21–27 May, Fig. S2 in the Supplement), supporting the validity of the SS assumption. Sampling in East 2 occurred 3 to 6 d after the window of success, and therefore, fluxes at the end of the sampling period might represent high end-member estimates (Ceballos-Romero et al., 2018).</p>
      <p id="d2e1869">In addition, the sampling strategy of the BELAS-1 expedition was incompatible with a non-steady state (NSS) model given that: (1) the sampling period was relatively short (6 d in East 1 and 4 d in East 2) relative to the 2–3 weeks recommended by Ceballos-Romero et al. (2018) to capture changes, if any, in <sup>234</sup>Th activities vs. time and apply a NSS approach; and (2) in order to obtain accurate flux estimates with a NSS model, a Lagrangian sampling strategy that tracks the same water mass must be followed (Resplandy et al., 2012; Savoye et al., 2006), which was not possible during this expedition. Indeed, changes in upper water column properties occurred between the sampling of East 1 and East 2 due to a storm (Erin Bertrand, personal communication, 2025), indicating that different water masses might have been sampled. Further, in the Eastern Grid, where we have the highest density of measurements, we did not observe significant changes in <sup>234</sup>Th inventories in the upper 150 m over time (Fig. S3a in the Supplement). Thus, we consider that the SS model gives the best estimate of <sup>234</sup>Th export in this study.</p>
</sec>
<sec id="Ch1.S2.SS5.SSS2">
  <label>2.5.2</label><title>Evaluating the effect of physical transport</title>
      <p id="d2e1907"><italic>Vertical Transport</italic>: The central Labrador Sea in 2022 experienced moderately deep convection reaching 1600 m (Yashayaev, 2024) followed by water column stratification and bloom development in early April. Winter mixing would homogenize vertical <sup>234</sup>Th gradients in the water column. However, sampling during BELAS-1 occurred at least 50 d after convection ended (Yashayaev, 2024), a time interval that exceeds the half-life of <sup>234</sup>Th and its mean life (1 <inline-formula><mml:math id="M170" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> decay constant <inline-formula><mml:math id="M171" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 35 d). Therefore, we are confident that vertical advection associated with winter mixing did not influence the estimates presented in this study. The potential contribution of vertical diffusion was estimated using vertical diffusivity estimates from a hindcast simulation using the model of Ohashi et al. (2024) in the Eastern Grid at the time of the cruise. Vertical diffusivity decreased with depth from 10<sup>−4</sup>–10<sup>−2</sup> m<sup>2</sup> s<sup>−1</sup> at the surface to 10<sup>−6</sup> m<sup>2</sup> s<sup>−1</sup> below 80 m. We estimate that vertical diffusion changed <sup>234</sup>Th fluxes at the base of the PPZ by <inline-formula><mml:math id="M180" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 10 dpm m<sup>−2</sup> d<sup>−1</sup> only, given the small gradients in <sup>234</sup>Th activities observed across the base of the PPZ at each station (Fig. 4). Therefore, the influence of vertical transport on <sup>234</sup>Th export flux estimates must have been negligible.</p>
      <p id="d2e2082"><italic>Horizontal Transport</italic>: Horizontal advection in the Eastern Grid, derived from the hindcast simulation of the cruise period, had mean velocities of 2.8 km d<sup>−1</sup> over the top 150 m in a mostly southeastward direction. The observed <sup>234</sup>Th activities show relatively large variability between profiles in the Eastern Grid (Fig. S4 in the Supplement), however, there are no consistent spatial trends in <sup>234</sup>Th inventories (Fig. S3b in the Supplement). We attribute the variability in <sup>234</sup>Th between profiles to small-scale spatial variations or patchiness rather than horizontal transport.</p>
</sec>
</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
      <p id="d2e2136">Table 1 presents a summary of the integrated NPP rates down to the base of the PPZ, together with the sinking fluxes (POC, bSi) measured at different depths in the upper 500 m across all the stations sampled during BELAS-1.</p>

<table-wrap id="T1" orientation="landscape"><label>Table 1</label><caption><p id="d2e2142">Integrated net primary production (NPP) down to the base of the primary production zone (PPZ), and particulate organic carbon (POC) and biogenic silica (bSi) fluxes at different depths across all stations.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="13">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="left"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:colspec colnum="7" colname="col7" align="right"/>
     <oasis:colspec colnum="8" colname="col8" align="right"/>
     <oasis:colspec colnum="9" colname="col9" align="right"/>
     <oasis:colspec colnum="10" colname="col10" align="right"/>
     <oasis:colspec colnum="11" colname="col11" align="right"/>
     <oasis:colspec colnum="12" colname="col12" align="right"/>
     <oasis:colspec colnum="13" colname="col13" align="right"/>
     <oasis:thead>
       <oasis:row>

         <oasis:entry colname="col1">Region</oasis:entry>

         <oasis:entry colname="col2">Station</oasis:entry>

         <oasis:entry colname="col3">Lat</oasis:entry>

         <oasis:entry colname="col4">Long</oasis:entry>

         <oasis:entry colname="col5">Sampling</oasis:entry>

         <oasis:entry colname="col6">Base of</oasis:entry>

         <oasis:entry colname="col7">Integrated</oasis:entry>

         <oasis:entry colname="col8">POC flux at</oasis:entry>

         <oasis:entry colname="col9">bSi flux at</oasis:entry>

         <oasis:entry colname="col10">POC flux at</oasis:entry>

         <oasis:entry colname="col11">bSi flux at</oasis:entry>

         <oasis:entry colname="col12">POC flux at</oasis:entry>

         <oasis:entry colname="col13">bSi flux at</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col1"/>

         <oasis:entry colname="col2">ID</oasis:entry>

         <oasis:entry colname="col3">(° N)</oasis:entry>

         <oasis:entry colname="col4">(° E)</oasis:entry>

         <oasis:entry colname="col5">date (2022)</oasis:entry>

         <oasis:entry colname="col6">PPZ (m)</oasis:entry>

         <oasis:entry colname="col7">NPP</oasis:entry>

         <oasis:entry colname="col8">PPZ base</oasis:entry>

         <oasis:entry colname="col9">PPZ base</oasis:entry>

         <oasis:entry colname="col10">PPZ base <inline-formula><mml:math id="M189" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 100 m</oasis:entry>

         <oasis:entry colname="col11">PPZ base <inline-formula><mml:math id="M190" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> 100 m</oasis:entry>

         <oasis:entry colname="col12">500 m</oasis:entry>

         <oasis:entry colname="col13">500 m</oasis:entry>

       </oasis:row>
       <oasis:row rowsep="1">

         <oasis:entry colname="col1"/>

         <oasis:entry colname="col2"/>

         <oasis:entry colname="col3"/>

         <oasis:entry colname="col4"/>

         <oasis:entry colname="col5"/>

         <oasis:entry colname="col6"/>

         <oasis:entry colname="col7">(mmol C m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

         <oasis:entry colname="col8">(mmol C m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

         <oasis:entry colname="col9">(mmol Si m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

         <oasis:entry colname="col10">(mmol C m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

         <oasis:entry colname="col11">(mmol Si m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

         <oasis:entry colname="col12">(mmol C m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

         <oasis:entry colname="col13">(mmol Si m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>

       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>

         <oasis:entry rowsep="1" colname="col1" morerows="5">East 1</oasis:entry>

         <oasis:entry colname="col2">4</oasis:entry>

         <oasis:entry colname="col3">58.82</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M205" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.11</oasis:entry>

         <oasis:entry colname="col5">20 May</oasis:entry>

         <oasis:entry colname="col6">119</oasis:entry>

         <oasis:entry colname="col7">166 <inline-formula><mml:math id="M206" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 25</oasis:entry>

         <oasis:entry colname="col8">16.3 <inline-formula><mml:math id="M207" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.0</oasis:entry>

         <oasis:entry colname="col9">1.8 <inline-formula><mml:math id="M208" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2</oasis:entry>

         <oasis:entry colname="col10">15.6 <inline-formula><mml:math id="M209" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.5</oasis:entry>

         <oasis:entry colname="col11">2.1 <inline-formula><mml:math id="M210" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col12">13.9 <inline-formula><mml:math id="M211" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 6.1</oasis:entry>

         <oasis:entry colname="col13">2.1 <inline-formula><mml:math id="M212" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">6</oasis:entry>

         <oasis:entry colname="col3">58.82</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M213" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.44</oasis:entry>

         <oasis:entry colname="col5">21 May</oasis:entry>

         <oasis:entry colname="col6">153</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">8.1 <inline-formula><mml:math id="M214" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.1</oasis:entry>

         <oasis:entry colname="col9">0.9 <inline-formula><mml:math id="M215" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col10">8.0 <inline-formula><mml:math id="M216" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.7</oasis:entry>

         <oasis:entry colname="col11">1.1 <inline-formula><mml:math id="M217" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col12">8.9 <inline-formula><mml:math id="M218" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.0</oasis:entry>

         <oasis:entry colname="col13">1.3 <inline-formula><mml:math id="M219" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">9</oasis:entry>

         <oasis:entry colname="col3">58.65</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M220" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.36</oasis:entry>

         <oasis:entry colname="col5">22 May</oasis:entry>

         <oasis:entry colname="col6">163</oasis:entry>

         <oasis:entry colname="col7">263 <inline-formula><mml:math id="M221" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 30</oasis:entry>

         <oasis:entry colname="col8">6.2 <inline-formula><mml:math id="M222" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.9</oasis:entry>

         <oasis:entry colname="col9">0.7 <inline-formula><mml:math id="M223" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col10">5.2 <inline-formula><mml:math id="M224" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.3</oasis:entry>

         <oasis:entry colname="col11">0.8 <inline-formula><mml:math id="M225" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col12">3.0 <inline-formula><mml:math id="M226" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.2</oasis:entry>

         <oasis:entry colname="col13">0.5 <inline-formula><mml:math id="M227" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">8</oasis:entry>

         <oasis:entry colname="col3">58.65</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M228" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.61</oasis:entry>

         <oasis:entry colname="col5">23 May</oasis:entry>

         <oasis:entry colname="col6">153</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">9.3 <inline-formula><mml:math id="M229" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.4</oasis:entry>

         <oasis:entry colname="col9">1.0 <inline-formula><mml:math id="M230" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col10">9.1 <inline-formula><mml:math id="M231" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.1</oasis:entry>

         <oasis:entry colname="col11">1.2 <inline-formula><mml:math id="M232" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col12">11.8 <inline-formula><mml:math id="M233" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.5</oasis:entry>

         <oasis:entry colname="col13">1.8 <inline-formula><mml:math id="M234" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">11</oasis:entry>

         <oasis:entry colname="col3">58.65</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M235" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.11</oasis:entry>

         <oasis:entry colname="col5">24 May</oasis:entry>

         <oasis:entry colname="col6">141</oasis:entry>

         <oasis:entry colname="col7">126 <inline-formula><mml:math id="M236" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 16</oasis:entry>

         <oasis:entry colname="col8">7.4 <inline-formula><mml:math id="M237" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.7</oasis:entry>

         <oasis:entry colname="col9">0.8 <inline-formula><mml:math id="M238" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2</oasis:entry>

         <oasis:entry colname="col10">5.5 <inline-formula><mml:math id="M239" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.3</oasis:entry>

         <oasis:entry colname="col11">0.7 <inline-formula><mml:math id="M240" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col12">–</oasis:entry>

         <oasis:entry colname="col13">–</oasis:entry>

       </oasis:row>
       <oasis:row rowsep="1">

         <oasis:entry colname="col2">12</oasis:entry>

         <oasis:entry colname="col3">58.48</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M241" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.11</oasis:entry>

         <oasis:entry colname="col5">25 May</oasis:entry>

         <oasis:entry colname="col6">176</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">2.9 <inline-formula><mml:math id="M242" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.8</oasis:entry>

         <oasis:entry colname="col9">0.3 <inline-formula><mml:math id="M243" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col10">4.1 <inline-formula><mml:math id="M244" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.1</oasis:entry>

         <oasis:entry colname="col11">0.6 <inline-formula><mml:math id="M245" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col12">7.0 <inline-formula><mml:math id="M246" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.6</oasis:entry>

         <oasis:entry colname="col13">1.1 <inline-formula><mml:math id="M247" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col1">St. 28</oasis:entry>

         <oasis:entry colname="col2">28-1</oasis:entry>

         <oasis:entry colname="col3">57.72</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M248" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>50.07</oasis:entry>

         <oasis:entry colname="col5">26 May</oasis:entry>

         <oasis:entry colname="col6">165</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">4.7 <inline-formula><mml:math id="M249" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.0</oasis:entry>

         <oasis:entry colname="col9">0.5 <inline-formula><mml:math id="M250" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col10">2.1 <inline-formula><mml:math id="M251" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.5</oasis:entry>

         <oasis:entry colname="col11">0.3 <inline-formula><mml:math id="M252" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col12">0.8 <inline-formula><mml:math id="M253" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.7</oasis:entry>

         <oasis:entry colname="col13">0.1 <inline-formula><mml:math id="M254" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6</oasis:entry>

       </oasis:row>
       <oasis:row rowsep="1">

         <oasis:entry colname="col1"/>

         <oasis:entry colname="col2">28-2</oasis:entry>

         <oasis:entry colname="col3">57.72</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M255" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>50.07</oasis:entry>

         <oasis:entry colname="col5">27 May</oasis:entry>

         <oasis:entry colname="col6">135</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">5.1 <inline-formula><mml:math id="M256" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.9</oasis:entry>

         <oasis:entry colname="col9">0.5 <inline-formula><mml:math id="M257" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.3</oasis:entry>

         <oasis:entry colname="col10">10.5 <inline-formula><mml:math id="M258" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.6</oasis:entry>

         <oasis:entry colname="col11">1.4 <inline-formula><mml:math id="M259" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col12">6.7 <inline-formula><mml:math id="M260" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.6</oasis:entry>

         <oasis:entry colname="col13">1.0 <inline-formula><mml:math id="M261" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col1">Central</oasis:entry>

         <oasis:entry colname="col2">24</oasis:entry>

         <oasis:entry colname="col3">56.65</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M262" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>52.47</oasis:entry>

         <oasis:entry colname="col5">28 May</oasis:entry>

         <oasis:entry colname="col6">115</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">12.6 <inline-formula><mml:math id="M263" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.3</oasis:entry>

         <oasis:entry colname="col9">1.4 <inline-formula><mml:math id="M264" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2</oasis:entry>

         <oasis:entry colname="col10">12.0 <inline-formula><mml:math id="M265" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.4</oasis:entry>

         <oasis:entry colname="col11">1.6 <inline-formula><mml:math id="M266" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col12">17.0 <inline-formula><mml:math id="M267" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 7.4</oasis:entry>

         <oasis:entry colname="col13">2.6 <inline-formula><mml:math id="M268" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6</oasis:entry>

       </oasis:row>
       <oasis:row rowsep="1">

         <oasis:entry colname="col1"/>

         <oasis:entry colname="col2">29</oasis:entry>

         <oasis:entry colname="col3">56.82</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M269" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>52.22</oasis:entry>

         <oasis:entry colname="col5">29 May</oasis:entry>

         <oasis:entry colname="col6">170</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">22.1 <inline-formula><mml:math id="M270" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.8</oasis:entry>

         <oasis:entry colname="col9">2.4 <inline-formula><mml:math id="M271" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col10">12.1 <inline-formula><mml:math id="M272" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.7</oasis:entry>

         <oasis:entry colname="col11">1.8 <inline-formula><mml:math id="M273" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5</oasis:entry>

         <oasis:entry colname="col12">9.4 <inline-formula><mml:math id="M274" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.5</oasis:entry>

         <oasis:entry colname="col13">1.4 <inline-formula><mml:math id="M275" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col1" morerows="2">East 2</oasis:entry>

         <oasis:entry colname="col2">15</oasis:entry>

         <oasis:entry colname="col3">58.48</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M276" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.61</oasis:entry>

         <oasis:entry colname="col5">30 May</oasis:entry>

         <oasis:entry colname="col6">155</oasis:entry>

         <oasis:entry colname="col7">–</oasis:entry>

         <oasis:entry colname="col8">7.2 <inline-formula><mml:math id="M277" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.0</oasis:entry>

         <oasis:entry colname="col9">1.3 <inline-formula><mml:math id="M278" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.4</oasis:entry>

         <oasis:entry colname="col10">3.4 <inline-formula><mml:math id="M279" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.7</oasis:entry>

         <oasis:entry colname="col11">0.9 <inline-formula><mml:math id="M280" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.7</oasis:entry>

         <oasis:entry colname="col12">2.7 <inline-formula><mml:math id="M281" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.0</oasis:entry>

         <oasis:entry colname="col13">0.6 <inline-formula><mml:math id="M282" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.8</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">13</oasis:entry>

         <oasis:entry colname="col3">58.48</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M283" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.36</oasis:entry>

         <oasis:entry colname="col5">31 May</oasis:entry>

         <oasis:entry colname="col6">143</oasis:entry>

         <oasis:entry colname="col7">38 <inline-formula><mml:math id="M284" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3</oasis:entry>

         <oasis:entry colname="col8">12.0 <inline-formula><mml:math id="M285" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.6</oasis:entry>

         <oasis:entry colname="col9">2.1 <inline-formula><mml:math id="M286" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5</oasis:entry>

         <oasis:entry colname="col10">11.5 <inline-formula><mml:math id="M287" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.3</oasis:entry>

         <oasis:entry colname="col11">3.0 <inline-formula><mml:math id="M288" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.7</oasis:entry>

         <oasis:entry colname="col12">8.1 <inline-formula><mml:math id="M289" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.3</oasis:entry>

         <oasis:entry colname="col13">1.8 <inline-formula><mml:math id="M290" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1.0</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">7</oasis:entry>

         <oasis:entry colname="col3">58.82</oasis:entry>

         <oasis:entry colname="col4"><inline-formula><mml:math id="M291" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>49.61</oasis:entry>

         <oasis:entry colname="col5">2 Jun</oasis:entry>

         <oasis:entry colname="col6">79</oasis:entry>

         <oasis:entry colname="col7">42 <inline-formula><mml:math id="M292" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 6</oasis:entry>

         <oasis:entry colname="col8">13.9 <inline-formula><mml:math id="M293" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.2</oasis:entry>

         <oasis:entry colname="col9">2.4 <inline-formula><mml:math id="M294" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2</oasis:entry>

         <oasis:entry colname="col10">18.7 <inline-formula><mml:math id="M295" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.4</oasis:entry>

         <oasis:entry colname="col11">3.3 <inline-formula><mml:math id="M296" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.7</oasis:entry>

         <oasis:entry colname="col12">9.9 <inline-formula><mml:math id="M297" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.6</oasis:entry>

         <oasis:entry colname="col13">2.2 <inline-formula><mml:math id="M298" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.9</oasis:entry>

       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

<sec id="Ch1.S3.SS1">
  <label>3.1</label><title>NPP rates</title>
      <p id="d2e3694">The average NPP at the surface and chl <inline-formula><mml:math id="M299" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> maximum depths was higher in East 1 (2.01 and 2.56 mmol C m<sup>−3</sup> d<sup>−1</sup>) compared to Central (1.79 and 1.74 mmol C m<sup>−3</sup> d<sup>−1</sup>) and East 2 (0.92 and 0.75 mmol C m<sup>−3</sup> d<sup>−1</sup>), which correlates with decreasing chl <inline-formula><mml:math id="M306" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> concentrations over the sampling period (Fig. 3). Surface NPP was highest at Station 11 (2.36 <inline-formula><mml:math id="M307" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>  0.47 mmol C m<sup>−3</sup> d<sup>−1</sup>) while NPP at the chl <inline-formula><mml:math id="M310" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> maximum peaked at Station 9 (3.38 <inline-formula><mml:math id="M311" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.83 mmol C m<sup>−3</sup> d<sup>−1</sup>). Station 13 had the lowest surface and chl <inline-formula><mml:math id="M314" display="inline"><mml:mi>a</mml:mi></mml:math></inline-formula> maximum NPP, 0.54 <inline-formula><mml:math id="M315" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.13  and 0.52 <inline-formula><mml:math id="M316" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.10 mmol C m<sup>−3</sup> d<sup>−1</sup>, respectively. Depth-integrated NPP was highest during East 1, peaking at Station 9 with a value of 263 <inline-formula><mml:math id="M319" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 30 mmol C m<sup>−2</sup> d<sup>−1</sup> (Table 1). Between sampling for East 1 and East 2, depth-integrated NPP decreased notably as the bloom declined and transitioned into post-bloom conditions, reaching a minimum of 38 <inline-formula><mml:math id="M322" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3 mmol C m<sup>−2</sup> d<sup>−1</sup> at Station 13 (Table 1). The measured in situ integrated NPP and satellite-derived NPP were in reasonable agreement (Fig. S5 in the Supplement), with a mean measured-satellite ratio of 1.25 <inline-formula><mml:math id="M325" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.69 (<inline-formula><mml:math id="M326" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">5</mml:mn></mml:mrow></mml:math></inline-formula>), supporting the use of satellite-derived NPP for estimating export efficiency (see Sect. 4.3).</p>
</sec>
<sec id="Ch1.S3.SS2">
  <label>3.2</label><title><sup>234</sup>Th <inline-formula><mml:math id="M328" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>238</sup>U profiles and <sup>234</sup>Th fluxes</title>
      <p id="d2e4025">In the upper water column, <sup>234</sup>Th activities were always lower than <sup>238</sup>U activities (<inline-formula><mml:math id="M333" display="inline"><mml:mo lspace="0mm">∼</mml:mo></mml:math></inline-formula> 2.4 dpm L<sup>−1</sup>) across all stations indicating particle export (Fig. 4). However, the magnitude and extent of the <sup>234</sup>Th deficits showed variability. The lowest <sup>234</sup>Th activities were found in surface waters of East 1, with average activities of 1.6 <inline-formula><mml:math id="M337" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2 dpm L<sup>−1</sup>, compared to Station 28, Central and East 2, where average surface activities were 1.9–2.0 dpm L<sup>−1</sup>. <sup>234</sup>Th activities reached equilibrium with <sup>238</sup>U at depths between 55 and 100 m, i.e., between the MLD and the base of the PPZ (Fig. 4). The only exception is Station 7 (East 2), the last station to be occupied, where equilibrium was only reached at <inline-formula><mml:math id="M342" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 150 m, below the PPZ.</p>
      <p id="d2e4141">In general, <sup>234</sup>Th and <sup>238</sup>U activities remained close to each other below the equilibrium depth. However, there were two notable exceptions. At some stations (i.e., East 1 Stations 9 and 12, and Station 28-1), large <sup>234</sup>Th activity excesses relative to <sup>238</sup>U were apparent within the PPZ, which is indicative of remineralization or disaggregation processes. In addition, we found a <sup>234</sup>Th deficit below the PPZ at Station 28-2, which can indicate particle repackaging processes.</p>
      <p id="d2e4189">One-dimensional (1D) steady state <sup>234</sup>Th fluxes (Fig. S6 in the Supplement) reflected the variability observed in <sup>234</sup>Th <inline-formula><mml:math id="M350" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>238</sup>U profiles. At the base of the PPZ, <sup>234</sup>Th fluxes ranged from negligible in East 1 to 1640 dpm m<sup>−2</sup> d<sup>−1</sup> in Central. On average, <sup>234</sup>Th fluxes (dpm m<sup>−2</sup> d<sup>−1</sup>) at the base of the PPZ were 620 <inline-formula><mml:math id="M358" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 330 in East 1, 360 <inline-formula><mml:math id="M359" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 20 at Station 28, 1290 <inline-formula><mml:math id="M360" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 500 in Central and 820 <inline-formula><mml:math id="M361" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 260 in East 2. In general, considering all stations, <sup>234</sup>Th fluxes increased slightly from the base of the PPZ to 100 m below the base (1.4-fold on average), and that increase was most pronounced (2.8-fold) at Station 28-2 due to the <sup>234</sup>Th deficit observed below the PPZ at this station (Fig. 4). At 500 m, <sup>234</sup>Th fluxes were moderately high, with fluxes <inline-formula><mml:math id="M365" display="inline"><mml:mo>≥</mml:mo></mml:math></inline-formula> 1000 dpm m<sup>−2</sup> d<sup>−1</sup> at most stations except three stations located outside the central region (Stations 9, 28-1, 15).</p>
</sec>
<sec id="Ch1.S3.SS3">
  <label>3.3</label><title>C <inline-formula><mml:math id="M368" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th and bSi <inline-formula><mml:math id="M369" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th in particles</title>
      <p id="d2e4404">The C <inline-formula><mml:math id="M370" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios in <inline-formula><mml:math id="M371" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M372" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m particles changed from 41 to 4.0 <inline-formula><mml:math id="M373" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> between near surface (40 m) and 490 m (Fig. 5). The highest C <inline-formula><mml:math id="M375" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios were found within the PPZ in East 1 where <italic>Phaeocystis</italic> dominated the phytoplankton community and phytoplankton biomass, and NPP rates were highest (Fig. 3). Below the PPZ, C <inline-formula><mml:math id="M376" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios decreased with depth from 13.5 <inline-formula><mml:math id="M377" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.8 <inline-formula><mml:math id="M378" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> at 115–185 m to 7.1 <inline-formula><mml:math id="M380" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2.7 <inline-formula><mml:math id="M381" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> at 465–490 m considering all stations.</p>
      <p id="d2e4522">bSi <inline-formula><mml:math id="M383" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios in <inline-formula><mml:math id="M384" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M385" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m particles also showed a general decrease with depth, but not as marked as C <inline-formula><mml:math id="M386" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios (Fig. 5). The highest bSi <inline-formula><mml:math id="M387" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios throughout all depths were found in East 2, with values decreasing from 2.3–2.7 <inline-formula><mml:math id="M388" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> in the upper 220 m to 1.6–1.9 <inline-formula><mml:math id="M390" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup>at deeper depths. In East 1, Station 28 and Central, bSi <inline-formula><mml:math id="M392" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios decreased from 1.5 <inline-formula><mml:math id="M393" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.1 <inline-formula><mml:math id="M394" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> at 115–185 m to 1.1 <inline-formula><mml:math id="M396" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.1 <inline-formula><mml:math id="M397" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> at 465–490 m.</p>
</sec>
<sec id="Ch1.S3.SS4">
  <label>3.4</label><title>POC and bSi fluxes</title>
      <p id="d2e4672">POC flux profiles show large station-to-station variability (Fig. 6a). At the base of the PPZ, POC fluxes ranged by 1 order of magnitude from 2.9 mmol C m<sup>−2</sup> d<sup>−1</sup> in East 1 to 22.1 mmol C m<sup>−2</sup> d<sup>−1</sup> in Central (Table 1). On average, POC fluxes (mmol C m<sup>−2</sup> d<sup>−1</sup>) at the base of the PPZ were 8.4 <inline-formula><mml:math id="M405" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.5 in East 1, 4.9 <inline-formula><mml:math id="M406" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.2 at Station 28, 17.3 <inline-formula><mml:math id="M407" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 6.7 in Central and 11.1 <inline-formula><mml:math id="M408" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.5 in East 2 (Figs. 6a, 7). POC fluxes at 100 m below the PPZ were not significantly different than those at the base of the PPZ (<inline-formula><mml:math id="M409" display="inline"><mml:mi>t</mml:mi></mml:math></inline-formula> test, <inline-formula><mml:math id="M410" display="inline"><mml:mi>p</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M411" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 0.05), indicating negligible attenuation in POC fluxes below the PPZ and a transfer efficiency close to 1 (i.e., flux at 100 m below PPZ base <inline-formula><mml:math id="M412" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> flux at PPZ base; Fig. 7). At 500 m, POC fluxes were, on average, 8.9 <inline-formula><mml:math id="M413" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.2 in East 1, 3.8 <inline-formula><mml:math id="M414" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4.1 at Station 28, 13.2 <inline-formula><mml:math id="M415" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5.3 in Central and 6.9 <inline-formula><mml:math id="M416" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3.7 mmol C m<sup>−2</sup> d<sup>−1</sup> in East 2. Only East 2 showed consistent flux attenuation (30 %–60 %) between the base of the PPZ and 500 m at all stations.</p>

      <fig id="F6" specific-use="star"><label>Figure 6</label><caption><p id="d2e4860">Profiles of particulate organic carbon (POC, panel <bold>a</bold>) and biogenic silica (bSi, panel <bold>b</bold>) flux across regions. Fluxes measured at individual stations are shown in grey, whereas averages are shown in color (green for East 1, light blue for St. 28, dark blue for Central, yellow for East 2). The vertical red lines at 20 mmol C m<sup>−2</sup> d<sup>−1</sup> for POC and 4 mmol Si m<sup>−2</sup> d<sup>−1</sup> for bSi are for visual reference. The base of the primary production zone (PPZ) is shown with green shading (grey solid line indicates the average, black solid lines indicate the minimum and maximum) and the average base of the mixed layer (MLD) is shown with a blue dashed line.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f06.png"/>

        </fig>

      <p id="d2e4924">bSi flux profiles also show large variability across stations (Fig. 6b). At the base of the PPZ, bSi fluxes ranged by 1 order of magnitude from 0.3 mmol Si m<sup>−2</sup> d<sup>−1</sup> in East 1 to 2.4 mmol Si m<sup>−2</sup> d<sup>−1</sup> in Central and East 2 (Table 1). On average, bSi fluxes (mmol Si m<sup>−2</sup> d<sup>−1</sup>) at the base of the PPZ were 0.9 <inline-formula><mml:math id="M429" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.5 in East 1, 0.5 <inline-formula><mml:math id="M430" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.0 at Station 28, 1.9 <inline-formula><mml:math id="M431" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.7 in Central and 1.9 <inline-formula><mml:math id="M432" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.6 in East 2 (Figs. 6b, 7). In deeper waters at most stations, bSi fluxes were similar or even higher than at the base of the PPZ indicating either no flux attenuation or addition of bSi at depth. The exception is East 2, where bSi flux attenuated by 10 %–50 % from the base of the PPZ to 500 m.</p>

      <fig id="F7" specific-use="star"><label>Figure 7</label><caption><p id="d2e5031">Spatial maps of particulate organic carbon (POC) and biogenic silica (bSi) flux at the base of the primary production zone (PPZ, left panels) and transfer efficiency (flux at 100 m below PPZ base <inline-formula><mml:math id="M433" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> flux at PPZ base) of POC and bSi over time in the study area (right panels). The horizontal dashed lines in the right panels indicate a transfer efficiency <inline-formula><mml:math id="M434" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f07.png"/>

        </fig>

</sec>
</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
      <p id="d2e5063">We begin the Discussion by providing an overview of the POC fluxes measured in this study and comparing them with those reported in other studies. We then examine the possible reasons why the massive <italic>Phaeocystis</italic> bloom observed here did not result in enhanced fluxes, with a particular focus on the ballasting hypothesis. Lastly, we present estimates of BCP efficiency during this <italic>Phaeocystis</italic> bloom using both snapshot and longer-term NPP observations. The assumptions underlying the export calculations, including the steady-state framework, are described in Sect. 2.5.</p>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>POC fluxes in the central Labrador Sea during the decline of a <italic>Phaeocystis</italic> bloom and comparison with other studies</title>
      <p id="d2e5084">This study presents an unprecedented number of measurements of sinking fluxes in the central Labrador Sea (<inline-formula><mml:math id="M435" display="inline"><mml:mo lspace="0mm">∼</mml:mo></mml:math></inline-formula> 300 <sup>234</sup>Th measurements) and represents one of the largest assessments of the impact of <italic>Phaeocystis</italic> blooms on POC export. Data collected by Fisheries and Oceans Canada along the Atlantic Repeat Hydrography Line 7 West (AR7W) as part of the Atlantic Zone Off-shelf Monitoring Program (AZOMP) between 2014 and 2022, revealed that the <italic>Phaeocystis</italic> bloom encountered in this study (spring 2022) was unusual given its large spatial extent and high biomass concentration (Devred et al., 2024, 2025). Using satellite remote sensing data combined with an ecological approach, these authors determined that the 2022 <italic>Phaeocystis</italic> bloom extended over more than half of the Labrador Sea, lasted for 6 weeks, and resulted in a remarkably high total primary production that accounted for 60 % of the May production in the Labrador Sea (33.2 Tg C; Devred et al., 2025). Genomic analyses conducted during our expedition, 1 week later than the 2022 AZOMP mission, confirmed that the species of this bloom was <italic>Phaeocystis pouchetii</italic> (Romanelli et al., 2026; Stevens-Green et al., 2024).</p>
      <p id="d2e5116">The sampling of the <italic>Phaeocystis</italic> bloom in this study mostly focused on a <inline-formula><mml:math id="M437" display="inline"><mml:mo>∼</mml:mo></mml:math></inline-formula> 1075 km<sup>2</sup> grid (Eastern Grid) at the southern edge of the bloom about 2 to 3 weeks after its peak. The stations sampled from 20  to 25 May (East 1) showed integrated NPP rates that were, on average, five times higher than those sampled from 30 May to 2 June (East 2) (Table 1) which we consider post-bloom conditions. Two stations located to the southwest of the Eastern Grid outside of the bloom (Central) were also sampled and represented more typical conditions of the central Labrador Sea with a mixed phytoplankton community composition that was more diatom dominated (Devred et al., 2024, 2025). Beyond the Eastern Grid, another station (Station 28) was also sampled in the <italic>Phaeocystis</italic> bloom area, but no in situ NPP data were collected at that station. For this reason, below we focus our discussion on the Eastern Grid and Central (inside vs. outside of the major <italic>Phaeocystis</italic> bloom).</p>
      <p id="d2e5144">POC fluxes at the base of the PPZ were, overall, moderate to high, with average fluxes of 8 and 11 mmol C m<sup>−2</sup> d<sup>−1</sup> in East 1 and East 2, respectively, and 17 mmol C m<sup>−2</sup> d<sup>−1</sup> in Central. This indicates that POC export was lower in the <italic>Phaeocystis</italic> bloom region relative to the stations sampled outside of the bloom. Our observations of large excesses of <sup>234</sup>Th relative to <sup>238</sup>U at around 70–100 m in the Eastern Grid (Stations 9 and 12) and Station 28-1 (Fig. 4) indicate remineralization of sinking particles within the euphotic zone in the <italic>Phaeocystis</italic> bloom region, and limited POC export to the mesopelagic. In general, POC fluxes did not decrease from the base of the PPZ to 500 m, except for East 2 where all stations showed a consistent decrease (30 %–60 %). This lack of flux attenuation down to 500 m suggests that the material that was not subject to remineralization in the euphotic zone was more refractory in nature or sank at relatively fast rates. POC fluxes at 500 m were on average 9 and 7 mmol C m<sup>−2</sup> d<sup>−1</sup> in East 1 and East 2, respectively, and 13 mmol C m<sup>−2</sup> d<sup>−1</sup> in Central. Therefore, our results also show lower fluxes at depth in the <italic>Phaeocystis</italic> bloom region than at the stations outside of the bloom. Despite the limited flux attenuation found in the upper mesopelagic, taken together, these findings suggest a limited role of <italic>Phaeocystis</italic> in export.</p>
      <p id="d2e5276">Only three prior studies have used <sup>234</sup>Th to estimate POC fluxes in the Labrador Sea: Moran et al. (2003) in July 1999 (three stations), Puigcorbé et al. (2017) in early May 2010 (one station), and Lemaitre et al. (2018) in late June 2014 (three stations), the last one during the decline of a diatom bloom. The POC flux results from our study are 1 order of magnitude higher than those measured at the PPZ base (or at 100 m, Moran et al., 2003) in May (0.6 mmol C m<sup>−2</sup> d<sup>−1</sup>, Puigcorbé et al., 2017), but encompass the range of POC fluxes measured in June (6.1–10 mmol C m<sup>−2</sup> d<sup>−1</sup>, Lemaitre et al., 2018) and July (5.7–21 mmol C m<sup>−2</sup> d<sup>−1</sup>, Moran et al., 2003). Compared to other <italic>Phaeocystis</italic> blooms (Table 2), our POC fluxes in the Eastern Grid compare well with those found in the Barents Sea using sediment traps (Lalande et al., 2008), but are lower than those measured in that same study using <sup>234</sup>Th and large volume sampling. Our POC fluxes in the Eastern Grid are also lower (2 to 8-fold) than those reported in a number of other Arctic and sub-Arctic studies during <italic>Phaeocystis</italic> blooms (Coppola et al., 2002; Dybwad et al., 2021; Le Moigne et al., 2015; Reigstad and Wassmann, 2007). Compared to studies that reported blooms dominated by both <italic>Phaeocystis</italic> and diatoms (Table 2), our POC fluxes fall within the range of those measured in the Crozet Plateau (Morris et al., 2007; Salter et al., 2007), but are lower (4-fold) than those reported in the Barents Sea (Andreassen and Wassmann, 1998) and the Ross Sea (Asper and Smith, 1999).</p>

<table-wrap id="T2" specific-use="star" orientation="landscape"><label>Table 2</label><caption><p id="d2e5382">Compilation of sinking fluxes and biological carbon pump efficiency metrics in <italic>Phaeocystis</italic> and diatom blooms. Values given in parentheses next to ranges indicate averages. PPZ <inline-formula><mml:math id="M457" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> primary production zone, PAR <inline-formula><mml:math id="M458" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> photosynthetic active radiation, Eq depth <inline-formula><mml:math id="M459" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> <sup>234</sup>Th and <sup>238</sup>U equilibrium depth, NPP <inline-formula><mml:math id="M462" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> net primary production, POC <inline-formula><mml:math id="M463" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> particulate organic carbon, bSi <inline-formula><mml:math id="M464" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> biogenic silica, ISP <inline-formula><mml:math id="M465" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> in situ pump, ST <inline-formula><mml:math id="M466" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> sediment trap.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="13">
     <oasis:colspec colnum="1" colname="col1" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="2" colname="col2" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="3" colname="col3" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="4" colname="col4" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="5" colname="col5" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="6" colname="col6" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="7" colname="col7" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="8" colname="col8" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="9" colname="col9" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="10" colname="col10" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="11" colname="col11" align="justify" colwidth="2cm"/>
     <oasis:colspec colnum="12" colname="col12" align="justify" colwidth="2.5cm"/>
     <oasis:colspec colnum="13" colname="col13" align="justify" colwidth="3cm"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">Study area</oasis:entry>
         <oasis:entry colname="col2" align="right">Reference depth <inline-formula><mml:math id="M467" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula> (m)</oasis:entry>
         <oasis:entry colname="col3" align="right">Integrated NPP                (mmol C m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>
         <oasis:entry colname="col4" align="right">POC flux at <inline-formula><mml:math id="M470" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula> (mmol C m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>
         <oasis:entry colname="col5" align="right">Export efficiency (%)</oasis:entry>
         <oasis:entry colname="col6" align="right">Transfer efficiency <inline-formula><mml:math id="M473" display="inline"><mml:mrow><mml:mi>z</mml:mi><mml:mo>+</mml:mo><mml:mn mathvariant="normal">100</mml:mn></mml:mrow></mml:math></inline-formula> m (%)</oasis:entry>
         <oasis:entry colname="col7" align="right">bSi flux at <inline-formula><mml:math id="M474" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula> (mmol Si m<sup>−2</sup> d<sup>−1</sup>)</oasis:entry>
         <oasis:entry colname="col8" align="right">Molar bSi <inline-formula><mml:math id="M477" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC at <inline-formula><mml:math id="M478" display="inline"><mml:mi>z</mml:mi></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom stage</oasis:entry>
         <oasis:entry colname="col10" align="left">Dominant species</oasis:entry>
         <oasis:entry colname="col11" align="left">Flux method</oasis:entry>
         <oasis:entry colname="col12" align="left">Number of stations considered</oasis:entry>
         <oasis:entry colname="col13" align="left">References</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1" align="left">Labrador Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">119–176 (PPZ)</oasis:entry>
         <oasis:entry colname="col3" align="right">126–263 (185)</oasis:entry>
         <oasis:entry colname="col4" align="right">2.9–16 (8.4)</oasis:entry>
         <oasis:entry colname="col5" align="right">2–10 (6)</oasis:entry>
         <oasis:entry colname="col6" align="right">74–139 (98)</oasis:entry>
         <oasis:entry colname="col7" align="right">0.3–1.8 (0.9)</oasis:entry>
         <oasis:entry colname="col8" align="right">0.1</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom decline</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic></oasis:entry>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th + ISPs</oasis:entry>
         <oasis:entry colname="col12" align="left">n = 6, East 1</oasis:entry>
         <oasis:entry colname="col13" align="left">This study</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left"/>
         <oasis:entry colname="col2" align="right">79–155 (PPZ)</oasis:entry>
         <oasis:entry colname="col3" align="right">38–42 (40)</oasis:entry>
         <oasis:entry colname="col4" align="right">7.2–14 (11)</oasis:entry>
         <oasis:entry colname="col5" align="right">32–34 (33)</oasis:entry>
         <oasis:entry colname="col6" align="right">48–134 (92)</oasis:entry>
         <oasis:entry colname="col7" align="right">1.3–2.4 (1.9)</oasis:entry>
         <oasis:entry colname="col8" align="right">0.2</oasis:entry>
         <oasis:entry colname="col9" align="left">Post-bloom</oasis:entry>
         <oasis:entry colname="col10" align="left"/>
         <oasis:entry colname="col11" align="left"/>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M480" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:math></inline-formula>, East 2</oasis:entry>
         <oasis:entry colname="col13" align="left"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1" align="left">Fram Strait</oasis:entry>
         <oasis:entry colname="col2" align="right">100</oasis:entry>
         <oasis:entry colname="col3" align="right">59 <inline-formula><mml:math id="M481" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 2</oasis:entry>
         <oasis:entry colname="col4" align="right">78 <inline-formula><mml:math id="M482" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 9</oasis:entry>
         <oasis:entry colname="col5" align="right">130 <inline-formula><mml:math id="M483" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 30</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">After bloom peak</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic></oasis:entry>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th <inline-formula><mml:math id="M485" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> ISP</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M486" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Le Moigne et al. (2015)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left"/>
         <oasis:entry colname="col2" align="right">100</oasis:entry>
         <oasis:entry colname="col3" align="right">48–64 (56)</oasis:entry>
         <oasis:entry colname="col4" align="right">6.8–13 (9.8)</oasis:entry>
         <oasis:entry colname="col5" align="right">11–30 (21)</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">During/after bloom peak</oasis:entry>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left"/>
         <oasis:entry colname="col12" align="left">n = 2</oasis:entry>
         <oasis:entry colname="col13" align="left"/>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1" align="left">Barents Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">60</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">14–62 (33)</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Early bloom/bloom/late bloom</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic></oasis:entry>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th <inline-formula><mml:math id="M488" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> large volume sampling</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M489" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">5</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Lalande et al. (2008)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left"/>
         <oasis:entry colname="col2" align="right">60</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">5.6–19 (12)</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left"/>
         <oasis:entry colname="col10" align="left"/>
         <oasis:entry colname="col11" align="left">Drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left"/>
         <oasis:entry colname="col13" align="left"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">Barents Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">90</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">17</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic></oasis:entry>
         <oasis:entry colname="col11" align="left">Drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M490" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Coppola et al. (2002)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1" align="left">Barents Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">40 (0.1 %PAR)</oasis:entry>
         <oasis:entry colname="col3" align="right">67–76 (71)</oasis:entry>
         <oasis:entry colname="col4" align="right">35–48 (42)</oasis:entry>
         <oasis:entry colname="col5" align="right">47–72 (59)</oasis:entry>
         <oasis:entry colname="col6" align="right">56–78 (67)</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Early bloom</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis </italic>and diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left">Drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M491" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Andreassen and Wassmann (1998), Buesseler et al. (2020)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left"/>
         <oasis:entry colname="col2" align="right">35 (0.1 %PAR)</oasis:entry>
         <oasis:entry colname="col3" align="right">68</oasis:entry>
         <oasis:entry colname="col4" align="right">54</oasis:entry>
         <oasis:entry colname="col5" align="right">79</oasis:entry>
         <oasis:entry colname="col6" align="right">41</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Well-developed bloom</oasis:entry>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left"/>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M492" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">North Norwegian fjords &amp; Barents Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">90–100</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">15–64 (40)</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom and after bloom</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic></oasis:entry>
         <oasis:entry colname="col11" align="left">Drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left">Average of several stations and cruises</oasis:entry>
         <oasis:entry colname="col13" align="left">Reigstad and Wassmann (2007)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1" align="left">North of Svalbard</oasis:entry>
         <oasis:entry colname="col2" align="right">50</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">13–33 (23)</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic></oasis:entry>
         <oasis:entry colname="col11" align="left">Ice-tethered STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M493" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">6</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Dybwad et al. (2021)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left"/>
         <oasis:entry colname="col2" align="right">60</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">15–47 (34)</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left"/>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left"/>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M494" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">4</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left"/>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">Ross Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">50</oasis:entry>
         <oasis:entry colname="col3" align="right">49–206 (105)</oasis:entry>
         <oasis:entry colname="col4" align="right">21–66 (34)</oasis:entry>
         <oasis:entry colname="col5" align="right">21–44 (34)</oasis:entry>
         <oasis:entry colname="col6" align="right">15–50 (35)</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">After bloom peak</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic> and diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left">Drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M495" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">5</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Asper and Smith (1999)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1" align="left">Crozet Plateau, Southern Ocean</oasis:entry>
         <oasis:entry colname="col2" align="right">87–248</oasis:entry>
         <oasis:entry colname="col3" align="right">–</oasis:entry>
         <oasis:entry colname="col4" align="right">1.0–17 (4.8)</oasis:entry>
         <oasis:entry colname="col5" align="right">–</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">0.3–29 (7.9)</oasis:entry>
         <oasis:entry colname="col8" align="right">0.2–2.8 (1.4)</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom decline</oasis:entry>
         <oasis:entry colname="col10" align="left"><italic>Phaeocystis</italic> and diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left">Neutrally buoyant STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M496" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M497" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> reoccupations</oasis:entry>
         <oasis:entry colname="col13" align="left">Salter et al. (2007)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left"/>
         <oasis:entry colname="col2" align="right">101–204</oasis:entry>
         <oasis:entry colname="col3" align="right">10–250 (70)</oasis:entry>
         <oasis:entry colname="col4" align="right">4.9–30 (17)</oasis:entry>
         <oasis:entry colname="col5" align="right">7–305 (54)</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left"/>
         <oasis:entry colname="col10" align="left"/>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th <inline-formula><mml:math id="M499" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> ISPs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M500" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">8</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M501" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> reoccupations</oasis:entry>
         <oasis:entry colname="col13" align="left">Morris et al. (2007)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">Labrador Sea</oasis:entry>
         <oasis:entry colname="col2" align="right">40–80 (Eq depth &amp; PPZ; <inline-formula><mml:math id="M502" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula>5)</oasis:entry>
         <oasis:entry colname="col3" align="right">27–80 (54)</oasis:entry>
         <oasis:entry colname="col4" align="right">6.1–10 (8.0)</oasis:entry>
         <oasis:entry colname="col5" align="right">8–38 (20)</oasis:entry>
         <oasis:entry colname="col6" align="right">30–63 (48)</oasis:entry>
         <oasis:entry colname="col7" align="right">2.7–6.6 (4.8)</oasis:entry>
         <oasis:entry colname="col8" align="right">0.4–0.8 (0.6)</oasis:entry>
         <oasis:entry colname="col9" align="left">After bloom peak</oasis:entry>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th <inline-formula><mml:math id="M504" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> ISPs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M505" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Lemaitre et al. (2018), Lemaitre (2017)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">Porcupine Abyssal Plain</oasis:entry>
         <oasis:entry colname="col2" align="right">67–133 (PPZ)</oasis:entry>
         <oasis:entry colname="col3" align="right">54–90 (75)</oasis:entry>
         <oasis:entry colname="col4" align="right">9.1–14 (11)</oasis:entry>
         <oasis:entry colname="col5" align="right">8–26 (16)</oasis:entry>
         <oasis:entry colname="col6" align="right">107–155 (139)</oasis:entry>
         <oasis:entry colname="col7" align="right">3.4–6.1 (4.4)</oasis:entry>
         <oasis:entry colname="col8" align="right">0.3–0.4 (0.4)</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom decline</oasis:entry>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th <inline-formula><mml:math id="M507" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> ISPs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M508" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">23</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Clevenger et al. (2024)</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1" align="left">Atlantic sector Southern Ocean</oasis:entry>
         <oasis:entry colname="col2" align="right">100–120</oasis:entry>
         <oasis:entry colname="col3" align="right">66–234 (153)</oasis:entry>
         <oasis:entry colname="col4" align="right">11–44 (19)</oasis:entry>
         <oasis:entry colname="col5" align="right">7–34 (12)</oasis:entry>
         <oasis:entry colname="col6" align="right">–</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom decline</oasis:entry>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left">Drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M509" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">11</mml:mn></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col13" align="left">Roca-Martí et al. (2017)</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1" align="left">Kerguelen Plateau, Southern Ocean</oasis:entry>
         <oasis:entry colname="col2" align="right">100</oasis:entry>
         <oasis:entry colname="col3" align="right">82</oasis:entry>
         <oasis:entry colname="col4" align="right">23 <inline-formula><mml:math id="M510" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 4</oasis:entry>
         <oasis:entry colname="col5" align="right">28</oasis:entry>
         <oasis:entry colname="col6" align="right">107</oasis:entry>
         <oasis:entry colname="col7" align="right">–</oasis:entry>
         <oasis:entry colname="col8" align="right">–</oasis:entry>
         <oasis:entry colname="col9" align="left">Bloom decline</oasis:entry>
         <oasis:entry colname="col10" align="left">Diatoms</oasis:entry>
         <oasis:entry colname="col11" align="left"><sup>234</sup>Th <inline-formula><mml:math id="M512" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> hose pump &amp; drifting STs</oasis:entry>
         <oasis:entry colname="col12" align="left"><inline-formula><mml:math id="M513" display="inline"><mml:mrow><mml:mi>n</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M514" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> reoccupations</oasis:entry>
         <oasis:entry colname="col13" align="left">Savoye et al. (2008)</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>Why did this <italic>Phaeocystis</italic> bloom not lead to enhanced fluxes?</title>
      <p id="d2e6813">Our observations during the decline of a massive <italic>Phaeocystis</italic> bloom in the Labrador Sea do not show indications of enhanced sinking fluxes when compared to an area located outside of the major bloom. These observations cover a period of 2 weeks during which NPP in the bloom area strongly declined, but POC fluxes remained relatively constant. As described in Sect. 2.5, the assumptions behind the <sup>234</sup>Th export model used in this study have been assessed and validated to the best of our possibilities based on information of the dynamics specific to that bloom and a hindcast simulation of physical transport at the time of the cruise. Furthermore, the conversion from <sup>234</sup>Th to POC fluxes using <inline-formula><mml:math id="M517" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M518" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m particles from large volume filtration in this study is supported with independent data from marine snow catchers (see Sect. 2.4). We also note that the C <inline-formula><mml:math id="M519" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> Th ratios used to estimate POC fluxes in this study (<inline-formula><mml:math id="M520" display="inline"><mml:mo lspace="0mm">≤</mml:mo></mml:math></inline-formula> 16 <inline-formula><mml:math id="M521" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup>) are far below the ratios measured by Lalande et al. (2008) of up to <inline-formula><mml:math id="M523" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 100 <inline-formula><mml:math id="M524" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>mol dpm<sup>−1</sup> which were considered to be biased towards high values and in turn led to overestimated POC-derived fluxes. Hence, the comparison of our data with context information and independent methods gives confidence in the export estimates presented here.</p>
      <p id="d2e6915"><italic>Phaeocystis</italic> spp. play an exceptional role in marine ecosystems due to its high carbon content when blooming and because of its unique polymorphic life cycle, including free-living cells of <inline-formula><mml:math id="M526" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 10 <inline-formula><mml:math id="M527" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m and gelatinous colonies that usually reach sizes of several mm (Schoemann et al., 2005). However, despite the large sizes of <italic>Phaeocystis</italic> colonies, most of the literature available on the topic shows that <italic>Phaeocystis</italic>-derived material is largely recycled in the upper ocean (Smith and Trimborn, 2024 and references therein). For instance, a compilation of sediment trap studies, including data ranging from polar to sub-Arctic and boreal regions, revealed that <italic>Phaeocystis</italic> POC fluxes strongly decline throughout the upper 100 m of the water column (Reigstad and Wassmann, 2007). In our study, the large excesses of <sup>234</sup>Th observed within the euphotic zone in the bloom region (Fig. 4), also support remineralization of sinking particles in the upper water column during <italic>Phaeocystis</italic> blooms. Yet, <italic>Phaeocystis</italic>-derived material can be efficiently exported to depth under certain circumstances, including: the formation of aggregates by <italic>Phaeocystis</italic> colonies facilitated by the release of transparent exopolymer particles (TEP) which can scavenge other particles like ballasting minerals (Andreassen and Wassmann, 1998; Passow and Wassmann, 1994; Wollenburg et al., 2018); and through physical mixing processes, such as downwelling associated with eddy activity (Lalande et al., 2011). The incorporation of <italic>Phaeocystis</italic> into fast-sinking zooplankton pellets has also been identified as an important export pathway (Dybwad et al., 2021; Wiedmann et al., 2020).</p>
      <p id="d2e6966">One hypothesis for the lack of enhanced sinking fluxes is that <italic>Phaeocystis</italic> material resulting from the decline of the bloom was not sufficiently ballasted. <italic>Phaeocystis</italic> colonies can release large amounts of TEP during their growth and their senescence, which, thanks to the high stickiness of TEP, can aggregate suspended particles and facilitate the formation of marine snow (Passow, 2002). However, the density of TEP is lower than that of seawater, which may cause the aggregates to remain in surface waters if not ballasted by other particles (Mari et al., 2017). In line with this, other studies have shown that below the mixed layer, marine biogels such as TEP can adhere to and accumulate on particles, reducing their sinking velocity by increasing both their buoyancy and hydrodynamic resistance, which may enhance carbon flux attenuation (Alcolombri et al., 2025; Romanelli et al., 2023). Sinking rates of <italic>Phaeocystis</italic> colonies and aggregates range from negligible to up to 200 m d<sup>−1</sup> (Schoemann et al., 2005), indicating that their capacity to sink and get transferred from the upper ocean to depth depends on their characteristics including the amount and type of particles that might have been scavenged. For instance, in the ice-covered Arctic Ocean, sinking of <italic>Phaeocystis</italic> aggregates throughout the water column down to the seafloor has been associated with ballasting by cryogenic gypsum which would have increased the density of such aggregates, facilitating their export to abyssal depths (Wollenburg et al., 2018). More common ballasting minerals that are known to increase the density of phytoplankton aggregates and enhance their export include continental dust, calcium carbonate, and biogenic silica (Armstrong et al., 2002). Yet, to our knowledge, the role of those minerals in controlling the sinking of <italic>Phaeocystis</italic> aggregates have not been explored.</p>
      <p id="d2e6998">If insufficient ballasting limited export, then we would expect a low contribution of dense mineral phases such as bSi to sinking particles. In line with this, molar bSi to POC ratios in particles <inline-formula><mml:math id="M530" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M531" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m within the PPZ were lower in East 1 (0.02–0.08) relative to those in Central (0.13) and East 2 (0.15–0.20), which likely suggests a change in phytoplankton community composition towards more diatoms in East 2. This observation is consistent with genomic samples taken during the cruise showing a lower relative contribution of diatoms to biomass in East 1 (Romanelli et al., 2026; Stevens-Green et al., 2024). Likewise, bSi fluxes at the base of the PPZ were 2-fold lower in East 1 than in Central and East 2 (Table 1), indicating a smaller contribution of siliceous plankton to export flux in East 1. However, below the PPZ, bSi fluxes in East 1 and East 2 became more similar, with fluxes within the same range at 500 m. Overall, bSi <inline-formula><mml:math id="M532" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC ratios were relatively low in all areas, with averages from the base of the PPZ down to 500 m of 0.14 <inline-formula><mml:math id="M533" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.03 (East 1), 0.18 <inline-formula><mml:math id="M534" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.06 (Central), and 0.22 <inline-formula><mml:math id="M535" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.04 (East 2), supporting a limited contribution of siliceous ballast in this study.</p>
      <p id="d2e7045">To our knowledge, bSi <inline-formula><mml:math id="M536" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC ratios during the decline of other <italic>Phaeocystis</italic> blooms have not been reported. Our bSi <inline-formula><mml:math id="M537" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC ratios fall in the lower range of those measured in the Labrador Sea (Lemaitre et al., 2018), the Porcupine Abyssal Plain (Clevenger et al., 2024), and the Crozet Plateau (Salter et al., 2007) during the decline of either diatom or mixed diatom and <italic>Phaeocystis</italic> blooms (Table 2). The bSi <inline-formula><mml:math id="M538" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC ratios measured in this study are also lower than those found in the upper mesopelagic of Ocean Station Papa during low flux conditions (0.63 <inline-formula><mml:math id="M539" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 0.28; Roca-Martí et al., 2021). Taken together, these results suggest that sinking particles during the BELAS-1 expedition were not heavily ballasted with bSi or, in other words, that diatoms did not appear to play an important ballasting role, which could at least partly explain the limited export of <italic>Phaeocystis</italic> in this study.</p>
      <p id="d2e7086">Other factors that may help explain why this <italic>Phaeocystis</italic> bloom did not lead to enhanced fluxes will be assessed elsewhere, including the analysis of the biochemical composition and morphology of suspended and sinking particles (Cisternas-Novoa et al., 2026) or their susceptibility to be degraded by heterotrophic bacteria (Romanelli et al., 2026).</p>
</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>BCP efficiency</title>
      <p id="d2e7101">The export efficiency (i.e., flux at PPZ base <inline-formula><mml:math id="M540" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> integrated NPP within PPZ) and transfer efficiency (i.e., flux at 100 m below PPZ base <inline-formula><mml:math id="M541" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> flux at PPZ base) are two common metrics for assessing the efficiency of the biological carbon pump and allowing comparisons between studies across different regions and seasons (Buesseler et al., 2020; Buesseler and Boyd, 2009). Here, we obtain export efficiencies ranging from 2 %–10 % in East 1 and 32 %–34 % in East 2 using in situ estimates of integrated NPP throughout the PPZ in combination with POC flux estimates measured at the base of that layer (Tables 1, 2). Transfer efficiencies ranged from 48 to 139 % and were on average 98 % in East 1 and 92 % in East 2 suggesting that flux attenuation throughout the upper 100 m of the mesopelagic zone was low. Combining both metrics, we obtain an overall BCP efficiency, defined as the amount of NPP reaching 100 m below the PPZ base, of 6 % in East 1 and 30 % in East 2. That would place this <italic>Phaeocystis</italic> bloom in the Labrador Sea as either a low BCP efficiency system (when looking at East 1 results) like the oligotrophic site ALOHA near Hawaii (Buesseler and Boyd, 2009), or a high BCP efficiency system (when looking at East 2 results) similar to that found in the Kerguelen Plateau (Savoye et al., 2008) and in the Barents Sea (Andreassen and Wassmann, 1998; Buesseler et al., 2020) during diatom and mixed diatom and <italic>Phaeocystis</italic> blooms (Fig. 8, Table 2).</p>

      <fig id="F8"><label>Figure 8</label><caption><p id="d2e7126">Biological carbon pump efficiency (BCP) comparing bloom studies across different regions (see Table 2). The <inline-formula><mml:math id="M542" display="inline"><mml:mi>x</mml:mi></mml:math></inline-formula>-axis depicts transfer efficiency (flux at 100 m below PPZ base <inline-formula><mml:math id="M543" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> flux at PPZ base) and the <inline-formula><mml:math id="M544" display="inline"><mml:mi>y</mml:mi></mml:math></inline-formula>-axis export efficiency (flux at PPZ base <inline-formula><mml:math id="M545" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> integrated NPP within PPZ). Contour lines represent overall BCP efficiency (percentages), and symbol area is proportional to NPP. The “East 1” and “East 2” symbols and the diamond correspond to this study. The diamond represents the overall BCP efficiency for the Labrador Sea (LS) 2022 bloom, calculated using satellite-derived NPP integrated from the bloom peak to the final day of sampling. Studies are colour-coded according to the dominant phytoplankton species. The ALOHA site (Buesseler and Boyd, 2009) is also shown as a low-end member. PPZ <inline-formula><mml:math id="M546" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> primary production zone, NPP <inline-formula><mml:math id="M547" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> net primary production.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/6229/2026/bg-23-6229-2026-f08.png"/>

        </fig>

      <p id="d2e7178">Yet, these BCP metrics are based on NPP and export estimates measured during the same time window. Given the similar POC fluxes measured in East 1 and East 2, the increase in export efficiencies observed with time in the Eastern Grid, from East 1 to East 2, is due to a decrease in NPP with time. This increase in export efficiencies may reflect either (1) a more efficient BCP during post-bloom conditions, or more likely (2) a temporal decoupling between primary production and export (Henson et al., 2015; Kiørboe et al., 1996; Smetacek et al., 1984) which have been shown to explain the inverse relationship between NPP and export efficiency found in different oceanic regions (Laws and Maiti, 2019; Roca-Martí et al., 2017). In other words, in this study, the export measured during post-bloom conditions in East 2, when NPP was low, could have been fueled with higher NPP rates that occurred at an earlier phase of the bloom. It is important to keep in mind that the NPP rates presented in this study were measured in the late stages of the bloom during its decline (East 1) and post-bloom phase (East 2) and therefore do not reflect the higher rates that occurred before the field campaign.</p>
      <p id="d2e7182">In order to estimate an overall BCP efficiency for the 2022 <italic>Phaeocystis</italic> bloom in the Labrador Sea (see diamond in Fig. 8), where NPP and export estimates are compared over longer timescales (Laws and Maiti, 2019), we have used satellite-derived NPP (Devred et al., 2025) in the Eastern Grid area from the peak of the <italic>Phaeocystis</italic> bloom until the last sampling day of the BELAS-1 expedition (9 May–2 June, Fig. S5 in the Supplement). The generally good agreement between in situ and satellite-derived NPP over the course of the expedition (Fig. S5 in the Supplement) provides confidence in the use of satellite NPP for calculating the overall efficiency. We have obtained an integrated NPP during that period of 3920 mmol C m<sup>−2</sup> which, combined with the POC fluxes measured in the Eastern Grid (average of 9.3 mmol C m<sup>−2</sup> d<sup>−1</sup> in East 1 and 2) multiplied by 25 d (i.e., 232 mmol C m<sup>−2</sup>), results in an overall export efficiency for the <italic>Phaeocystis</italic> bloom decline of 6 %. This estimate assumes that the POC export measured in East 1 and 2 using <sup>234</sup>Th (half-life <inline-formula><mml:math id="M553" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 24.1 d) is representative of the export that occurred over the 25 d from the peak of the bloom until the end of the bloom. Using that export efficiency combined with an average transfer efficiency in the Eastern Grid of 96 % (East 1 and 2), we would obtain an overall BCP efficiency of 6 %, which is the same as that obtained in East 1 (Fig. 8). This analysis suggests that East 1 results represent better the overall BCP efficiency of the extensive <italic>Phaeocystis</italic> bloom occurred in the Labrador Sea in spring 2022 and stresses the importance of long-term observations of the BCP. The overall BCP efficiency obtained in this study is lower than that reported during other blooms in the North Atlantic and Southern Ocean, dominated by either diatoms or <italic>Phaeocystis</italic> and diatoms (Fig. 8, Table 2). Insufficient mineral ballasting may have contributed to the low BCP efficiency observed during this massive <italic>Phaeocystis</italic> bloom.</p>
      <p id="d2e7269">Altogether, our findings highlight the complexity of quantifying the contribution of phytoplankton blooms to carbon export and underscore the importance of sustained observations to better capture their variability and broader implications in a changing ocean. This study provides a foundation for future comparisons with other datasets from the BELAS-1 expedition. Integrating these datasets will help elucidate the mechanisms underlying low BCP efficiency events, such as the one observed during this study, thereby improving our ability to predict the consequences of shifting bloom dynamics for global carbon export. Moreover, future efforts will offer the opportunity to extend the observational period by incorporating data from autonomous platforms, thereby shedding light on the evolution of sinking fluxes beyond the limited timeframe of ship-based campaigns.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusions</title>
      <p id="d2e7282">This study represents one of the largest assessments ever of the impact of <italic>Phaeocystis</italic> blooms on sinking fluxes using <sup>234</sup>Th as a tracer during a 2-week-long process study in spring 2022. The <italic>Phaeocystis</italic> bloom encountered was unusually large and highly productive extending over more than half of the Labrador Sea for 6 weeks. The main conclusions of this work are summarized below: <list list-type="bullet"><list-item>
      <p id="d2e7302">During the late stages of the bloom, POC fluxes in the upper mesopelagic down to 500 m were variable but overall moderate to high (average of 8 <inline-formula><mml:math id="M555" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 5 mmol C m<sup>−2</sup> d<sup>−1</sup>). Yet, evidence of shallow POC flux remineralization combined with the fact that POC fluxes in the bloom were not higher than in a region sampled outside of the bloom (average of 13 <inline-formula><mml:math id="M558" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 3 mmol C m<sup>−2</sup> d<sup>−1</sup>) suggests a limited role of <italic>Phaeocystis</italic> in carbon export.</p></list-item><list-item>
      <p id="d2e7372">Large (<inline-formula><mml:math id="M561" display="inline"><mml:mo lspace="0mm">&gt;</mml:mo></mml:math></inline-formula> 51 <inline-formula><mml:math id="M562" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">µ</mml:mi></mml:mrow></mml:math></inline-formula>m) particles collected using large volume pumps presented relatively low bSi <inline-formula><mml:math id="M563" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC ratios and, therefore, diatoms did not appear to have an important ballasting role of <italic>Phaeocystis</italic>-derived material during the observation period.</p></list-item><list-item>
      <p id="d2e7401">2 weeks after the peak of the bloom, the total amount of in situ NPP that reached 100 m below the euphotic zone was only 6 %. However, 3 weeks after the peak of the bloom, the value had increased to 30 %. This apparent change was driven primarily by a decline in NPP over the sampling period, highlighting how sensitive the BCP efficiency determination is to temporal changes in NPP.</p></list-item><list-item>
      <p id="d2e7405">Using satellite-derived NPP from the peak of the bloom until its end, we obtain an overall BCP efficiency of 6 % supporting a low BCP efficiency system. We stress the importance of long-term observations of both NPP and POC export for estimating meaningful BCP efficiencies. Future research including data from autonomous platforms will elucidate how sinking fluxes might have changed after this ship-based observation study.</p></list-item><list-item>
      <p id="d2e7409">The BCP efficiency of this <italic>Phaeocystis</italic> bloom is clearly lower than that found during the decline of either diatom blooms or mixed diatom and <italic>Phaeocystis</italic> blooms (albeit with no bSi <inline-formula><mml:math id="M564" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> POC information) in the North Atlantic and Southern Ocean.</p></list-item><list-item>
      <p id="d2e7426">To elucidate under which conditions <italic>Phaeocystis</italic> can be a good exporter, future research should include the analysis of bSi and other ballasting minerals, if possible, in all particle size fractions. These measurements will be key to shedding light on the role of the BCP in sequestering carbon in a future ocean where small cells, such as <italic>Phaeocysti</italic>s, will increasingly dominate phytoplankton communities (Finkel et al., 2010; Passow and Carlson, 2012).</p></list-item></list></p>
</sec>

      
      </body>
    <back><notes notes-type="dataavailability"><title>Data availability</title>

      <p id="d2e7439">Thorium-234 (<sup>234</sup>Th) and size-fractionated particulate data were published open access (Roca‐Martí et al., 2025a, b).</p>
  </notes><app-group>
        <supplementary-material position="anchor"><p id="d2e7451">The supplement related to this article is available online at <inline-supplementary-material xlink:href="https://doi.org/10.5194/bg-23-6229-2026-supplement" xlink:title="pdf">https://doi.org/10.5194/bg-23-6229-2026-supplement</inline-supplementary-material>.</p></supplementary-material>
        </app-group><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d2e7460">SK contributed to funding acquisition. MRM, MH, SK contributed to the experimental conceptual design. All authors contributed to data generation and analysis. MRM, MH, CM, SK contributed to the initial manuscript draft (MH data visualization), and all co-authors contributed to the revision of the paper.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d2e7466">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="d2e7472">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="d2e7478">This work is dedicated to Markus Kienast, a dear colleague whose approach to both science and life deeply inspires us all. We will always remember his passion, enthusiasm, positivity, and sense of humor. This study is part of the Ocean Frontier Institute's research project “The Northwest Atlantic Biological Carbon Pump” (NWA-BCP). We are sincerely grateful to the crew and scientific party aboard the RV <italic>Celtic Explorer</italic> during the BELAS-1 expedition and the whole NWA-BCP team. We would like to acknowledge Maria Armstrong (<sup>234</sup>Th team), and Britton Dempsey and Brianna Stanley (NPP team) for their invaluable help with the expedition's preparation, sample collection and/or processing. We also thank Stephanie Clay for providing satellite data, Gretchen Swarr and Claire Normandeau for measuring <sup>230</sup>Th <inline-formula><mml:math id="M568" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>229</sup>Th and POC, and Erin Bertrand, Ken Buesseler and Uta Passow for providing insightful discussions about the data. We thank OBPG-NASA for making satellite ocean colour data available and the support from the DFO Atlantic Zone Off-shelf Monitoring Program to collect in situ data used to develop satellite-based data products.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d2e7520">MRM acknowledges support from the Ocean Frontier Institute (OFI) International Postdoctoral Fellowship Program, the Beatriu de Pinós Fellowship (grant no. 2021-BP-00109), the “la Caixa” Foundation (ID 100010434, fellowship code LCF/BQ/PI24/12040022), the Ramón y Cajal Program (grant no. RYC2023-045355-I), and ICTA-UAB MERS (grant no. 2021 SGR-640) of the Generalitat de Catalunya. MH was supported by the Nova Scotia Graduate Scholarship (NSGS), and SK by the Natural Sciences and Engineering Research Council of Canada (NSERC) and OFI.</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d2e7526">This paper was edited by Andrew Thurber and reviewed by two anonymous referees.</p>
  </notes><ref-list>
    <title>References</title>

      <ref id="bib1.bib1"><label>1</label><mixed-citation>Alcolombri, U., Nissan, A., Słomka, J., Charlton, S., Secchi, E., Short, I., Lee, K. S., Peaudecerf, F. J., Baumgartner, D. A., Sichert, A., Sauer, U., Sengupta, A., and Stocker, R.: Biogel scavenging slows the sinking of organic particles to the ocean depths, Nat. Commun., 161,  1–10, <ext-link xlink:href="https://doi.org/10.1038/s41467-025-57982-5" ext-link-type="DOI">10.1038/s41467-025-57982-5</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bib2"><label>2</label><mixed-citation>Andreassen, I. J. and Wassmann, P.: Vertical flux of phytoplankton and particulate biogenic matter in the marginal ice zone of the Barents Sea in May 1993, Mar. Ecol. Prog. Ser., 170, 1–14, <ext-link xlink:href="https://doi.org/10.3354/MEPS170001" ext-link-type="DOI">10.3354/MEPS170001</ext-link>, 1998.</mixed-citation></ref>
      <ref id="bib1.bib3"><label>3</label><mixed-citation>Armstrong, R. A., Lee, C., Hedges, J. I., Honjo, S., and Wakeham, S. G.: A new, mechanistic model for organic carbon fluxes in the ocean based on the quantitative association of POC with ballast minerals, Deep-Sea Res. Pt. II, 49, 219–236, <ext-link xlink:href="https://doi.org/10.1016/S0967-0645(01)00101-1" ext-link-type="DOI">10.1016/S0967-0645(01)00101-1</ext-link>, 2002.</mixed-citation></ref>
      <ref id="bib1.bib4"><label>4</label><mixed-citation>Arruda, R., Atamanchuk, D., Boteler, C., and Wallace, D. W. R.: Seasonality of pCO<sub>2</sub> and air-sea CO<sub>2</sub> fluxes in the Central Labrador Sea, Front. Mar. Sci., 11, 1472697, <ext-link xlink:href="https://doi.org/10.3389/fmars.2024.1472697" ext-link-type="DOI">10.3389/fmars.2024.1472697</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib5"><label>5</label><mixed-citation>Asper, V. L. and Smith, W. O.: Particle fluxes during austral spring and summer in the southern Ross Sea, Antarctica, J. Geophys. Res.-Ocean., 104, 5345–5359, <ext-link xlink:href="https://doi.org/10.1029/1998JC900067" ext-link-type="DOI">10.1029/1998JC900067</ext-link>, 1999.</mixed-citation></ref>
      <ref id="bib1.bib6"><label>6</label><mixed-citation>Baker, C. A., Martin, A. P., Yool, A., and Popova, E.: Biological Carbon Pump Sequestration Efficiency in the North Atlantic: A Leaky or a Long-Term Sink?, Global Biogeochem. Cy., 36, e2021GB007286, <ext-link xlink:href="https://doi.org/10.1029/2021GB007286" ext-link-type="DOI">10.1029/2021GB007286</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib7"><label>7</label><mixed-citation>Baltar, F., Arístegui, J., Gasol, J. M., Sintes, E., and Herndl, G. J.: Evidence of prokaryotic metabolism on suspended particulate organic matter in the dark waters of the subtropical North Atlantic, Limnol. Oceanogr., 54, 182–193, <ext-link xlink:href="https://doi.org/10.4319/LO.2009.54.1.0182" ext-link-type="DOI">10.4319/LO.2009.54.1.0182</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib8"><label>8</label><mixed-citation>Baumas, C., Fuchs, R., Garel, M., Poggiale, J.-C., Memery, L., Le Moigne, F. A. C., and Tamburini, C.: Reconstructing the ocean's mesopelagic zone carbon budget: sensitivity and estimation of parameters associated with prokaryotic remineralization, Biogeosciences, 20, 4165–4182, <ext-link xlink:href="https://doi.org/10.5194/BG-20-4165-2023" ext-link-type="DOI">10.5194/BG-20-4165-2023</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib9"><label>9</label><mixed-citation>Bishop, J. K. B., Lam, P. J., and Wood, T. J.: Getting good particles: Accurate sampling of particles by large volume in-situ filtration, Limnol. Oceanogr. Method., 10, 681–710, <ext-link xlink:href="https://doi.org/10.4319/lom.2012.10.681" ext-link-type="DOI">10.4319/lom.2012.10.681</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib10"><label>10</label><mixed-citation>Boyd, P. W., Sherry, N. D., Berges, J. A., Bishop, J. K. B., Calvert, S. E., Charette, M. A., Giovannoni, S. J., Goldblatt, R., Harrison, P. J., Moran, S. B., Roy, S., Soon, M., Strom, S., Thibault, D., Vergin, K. L., Whitney, F. A., and Wong, C. S.: Transformations of biogenic particulates from the pelagic to the deep ocean realm, Deep-Sea Res. Pt. II, 46, 2761–2792, <ext-link xlink:href="https://doi.org/10.1016/S0967-0645(99)00083-1" ext-link-type="DOI">10.1016/S0967-0645(99)00083-1</ext-link>, 1999.</mixed-citation></ref>
      <ref id="bib1.bib11"><label>11</label><mixed-citation>Boyd, P. W., Claustre, H., Levy, M., Siegel, D. A., and Weber, T.: Multi-faceted particle pumps drive carbon sequestration in the ocean, Nature, 568, 327–335, <ext-link xlink:href="https://doi.org/10.1038/s41586-019-1098-2" ext-link-type="DOI">10.1038/s41586-019-1098-2</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib12"><label>12</label><mixed-citation>Buesseler, K. O. and Boyd, P. W.: Shedding light on processes that control particle export and flux attenuation in the twilight zone of the open ocean, Limnol. Oceanogr., 54, 1210–1232, <ext-link xlink:href="https://doi.org/10.4319/lo.2009.54.4.1210" ext-link-type="DOI">10.4319/lo.2009.54.4.1210</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib13"><label>13</label><mixed-citation>Buesseler, K. O., Boyd, P. W., Black, E. E., and Siegel, D. A.: Metrics that matter for assessing the ocean biological carbon pump, P. Natl. Acad. Sci. USA,  117, 9679–9687, <ext-link xlink:href="https://doi.org/10.1073/PNAS.1918114117" ext-link-type="DOI">10.1073/PNAS.1918114117</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib14"><label>14</label><mixed-citation>Burd, A. B., Hansell, D. A., Steinberg, D. K., Anderson, T. R., Arístegui, J., Baltar, F., Beaupré, S. R., Buesseler, K. O., DeHairs, F., Jackson, G. A., Kadko, D. C., Koppelmann, R., Lampitt, R. S., Nagata, T., Reinthaler, T., Robinson, C., Robison, B. H., Tamburini, C., and Tanaka, T.: Assessing the apparent imbalance between geochemical and biochemical indicators of meso- and bathypelagic biological activity: What the @$♯! is wrong with present calculations of carbon budgets?, Deep-Sea Res. Pt. II, 57, 1557–1571, <ext-link xlink:href="https://doi.org/10.1016/j.dsr2.2010.02.022" ext-link-type="DOI">10.1016/j.dsr2.2010.02.022</ext-link>, 2010.</mixed-citation></ref>
      <ref id="bib1.bib15"><label>15</label><mixed-citation>Ceballos-Romero, E., De Soto, F., Le Moigne, F. A. C., García-Tenorio, R., and Villa-Alfageme, M.: <sup>234</sup>Th-derived particle fluxes and seasonal variability: when is the SS assumption reliable? Insights from a novel approach for carbon flux simulation, Geophys. Res. Lett., 45, 13414–13426, <ext-link xlink:href="https://doi.org/10.1029/2018GL079968" ext-link-type="DOI">10.1029/2018GL079968</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib16"><label>16</label><mixed-citation>Cisternas-Novoa, C., Romanelli, E., and Passow, U.: Differences between suspended and sinking particles regulate carbon flux in the upper mesopelagic during a <italic>Phaeocystis</italic> Bloom, preprint, <ext-link xlink:href="https://doi.org/10.64898/2026.06.09.731151" ext-link-type="DOI">10.64898/2026.06.09.731151</ext-link>, 2026.</mixed-citation></ref>
      <ref id="bib1.bib17"><label>17</label><mixed-citation>Claustre, H., Legendre, L., Boyd, P. W., and Levy, M.: The Oceans' Biological Carbon Pumps: Framework for a Research Observational Community Approach, Front. Mar. Sci., 8, <ext-link xlink:href="https://doi.org/10.3389/fmars.2021.780052" ext-link-type="DOI">10.3389/fmars.2021.780052</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib18"><label>18</label><mixed-citation>Clevenger, S. J., Benitez-Nelson, C. R., Drysdale, J., Pike, S., Puigcorbé, V., and Buesseler, K. O.: Review of the analysis of <sup>234</sup>Th in small volume (2–4 L) seawater samples: improvements and recommendations, J. Radioanal. Nucl. Chem., 329, 1–13, <ext-link xlink:href="https://doi.org/10.1007/s10967-021-07772-2" ext-link-type="DOI">10.1007/s10967-021-07772-2</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib19"><label>19</label><mixed-citation>Clevenger, S. J., Benitez-Nelson, C. R., Roca-Martí, M., Bam, W., Estapa, M., Kenyon, J. A., Pike, S., Resplandy, L., Wyatt, A., and Buesseler, K. O.: Carbon and silica fluxes during a declining North Atlantic spring bloom as part of the EXPORTS program, Mar. Chem., 258, 104346, <ext-link xlink:href="https://doi.org/10.1016/J.MARCHEM.2023.104346" ext-link-type="DOI">10.1016/J.MARCHEM.2023.104346</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib20"><label>20</label><mixed-citation>Coppola, L., Roy-Barman, M., Wassmann, P., Mulsow, S., and Jeandel, C.: Calibration of sediment traps and particulate organic carbon export using <sup>234</sup>Th in the Barents Sea, Mar. Chem., 80, 11–26, <ext-link xlink:href="https://doi.org/10.1016/S0304-4203(02)00071-3" ext-link-type="DOI">10.1016/S0304-4203(02)00071-3</ext-link>, 2002.</mixed-citation></ref>
      <ref id="bib1.bib21"><label>21</label><mixed-citation>de Boyer Montégut, C., Madec, G., Fischer, A. S., Lazar, A., and Iudicone, D.: Mixed layer depth over the global ocean: An examination of profile data and a profile-based climatology, J. Geophys. Res., 109, C12003, <ext-link xlink:href="https://doi.org/10.1029/2004JC002378" ext-link-type="DOI">10.1029/2004JC002378</ext-link>, 2004.</mixed-citation></ref>
      <ref id="bib1.bib22"><label>22</label><mixed-citation> Devred, E., Wilson, K. L., Perry, T., Hardy, M., Brosnahan, M., and Ringuette, M.: Identification and validation of phytoplankton taxonomic assemblages derived from pigment signatures using samples collected in the Labrador Sea from 2014 to 2022, Dartmouth, Nova Scotia, Canada, 37 pp., ISBN: 978-0-660-70848-5, 2024.</mixed-citation></ref>
      <ref id="bib1.bib23"><label>23</label><mixed-citation>Devred, E., Clay, S., Ringuette, M., Perry, T., Amirian, M., Irwin, A., and Finkel, Z.: Net primary production in the Labrador Sea between 2014 and 2022 derived from ocean colour remote sensing based on ecological regimes, Remote Sens. Environ., 323, 114713, <ext-link xlink:href="https://doi.org/10.1016/J.RSE.2025.114713" ext-link-type="DOI">10.1016/J.RSE.2025.114713</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bib24"><label>24</label><mixed-citation>Dybwad, C., Assmy, P., Olsen, L. M., Peeken, I., Nikolopoulos, A., Krumpen, T., Randelhoff, A., Tatarek, A., Wiktor, J. M., and Reigstad, M.: Carbon Export in the Seasonal Sea Ice Zone North of Svalbard From Winter to Late Summer, Front. Mar. Sci., 7, 525800, <ext-link xlink:href="https://doi.org/10.3389/fmars.2020.525800" ext-link-type="DOI">10.3389/fmars.2020.525800</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib25"><label>25</label><mixed-citation>Finkel, Z. V., Beardall, J., Flynn, K. J., Quigg, A., Rees, T. A. V., and Raven, J. A.: Phytoplankton in a changing world: cell size and elemental stoichiometry, J. Plankton Res., 32, 119–137, <ext-link xlink:href="https://doi.org/10.1093/PLANKT/FBP098" ext-link-type="DOI">10.1093/PLANKT/FBP098</ext-link>, 2010.</mixed-citation></ref>
      <ref id="bib1.bib26"><label>26</label><mixed-citation>Giering, S. L. C., Sanders, R., Lampitt, R. S., Anderson, T. R., Tamburini, C., Boutrif, M., Zubkov, M. V, Marsay, C. M., Henson, S. A., Saw, K., Cook, K., and Mayor, D. J.: Reconciliation of the carbon budget in the ocean's twilight zone, Nature, 507, 480–483, <ext-link xlink:href="https://doi.org/10.1038/nature13123" ext-link-type="DOI">10.1038/nature13123</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib27"><label>27</label><mixed-citation>Giering, S. L. C., Sanders, R., Martin, A. P., Lindemann, C., Möller, K. O., Daniels, C. J., Mayor, D. J., and St. John, M. A.: High export via small particles before the onset of the North Atlantic spring bloom, J. Geophys. Res.-Ocean., 121, 6929–6945, <ext-link xlink:href="https://doi.org/10.1002/2016JC012048" ext-link-type="DOI">10.1002/2016JC012048</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib28"><label>28</label><mixed-citation>Graff, J. R., Nelson, N. B., Roca-Martí, M., Romanelli, E., Kramer, S. J., Erickson, Z., Cetinic, I., Buesseler, K. O., Passow, U., Zhang, X., Benitez-Nelson, C., Bisson, K., Close, H. G., Crockford, T., Fox, J., Halewood, S., Lam, P., Roesler, C., Sweet, J., VerWey, B., Xiong, Y., and Siegel, D. A.: Reconciliation of total particulate organic carbon and nitrogen measurements determined using contrasting methods in the North Pacific Ocean as part of the NASA EXPORTS field campaign, Elem. Sci. Anthr., 11, <ext-link xlink:href="https://doi.org/10.1525/elementa.2022.00112" ext-link-type="DOI">10.1525/elementa.2022.00112</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib29"><label>29</label><mixed-citation>Hama, T., Miyazaki, T., Ogawa, Y., Iwakuma, T., Takahashi, M., Otsuki, A., and Ichimura, S.: Measurement of photosynthetic production of a marine phytoplankton population using a stable <sup>13</sup>C isotope, Mar. Biol., 73, 31–36, <ext-link xlink:href="https://doi.org/10.1007/BF00396282" ext-link-type="DOI">10.1007/BF00396282</ext-link>, 1983.</mixed-citation></ref>
      <ref id="bib1.bib30"><label>30</label><mixed-citation>Henson, S. A., Yool, A., and Sanders, R.: Variability in efficiency of particulate organic carbon export: A model study, Global Biogeochem. Cy., 29, 33–45, <ext-link xlink:href="https://doi.org/10.1002/2014GB004965" ext-link-type="DOI">10.1002/2014GB004965</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib31"><label>31</label><mixed-citation>Henson, S. A., Laufkötter, C., Leung, S., Giering, S. L. C., Palevsky, H. I., and Cavan, E. L.: Uncertain response of ocean biological carbon export in a changing world, Nat. Geosci. 2022 154, 15, 248–254, <ext-link xlink:href="https://doi.org/10.1038/s41561-022-00927-0" ext-link-type="DOI">10.1038/s41561-022-00927-0</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib32"><label>32</label><mixed-citation> Kiørboe, T., Hansen, J. L. S., Alldredge, A. L., Jackson, G. A., Passow, U., Dam, H. G., Drapeau, D. T., Waite, A., and Garcia, C. M.: Sedimentation of phytoplankton during a diatom bloom: Rates and mechanisms, J. Mar. Res., 54, 1123–148, 1996.</mixed-citation></ref>
      <ref id="bib1.bib33"><label>33</label><mixed-citation> Knap, A., Michaels, A., Close, A., Ducklow, H., and Dickson, A.: Protocols for the Joint Global Ocean Flux Study (JGOFS) Core Measurements, JGOFS Report Nr. 19, 1–210,  1996.</mixed-citation></ref>
      <ref id="bib1.bib34"><label>34</label><mixed-citation>Lalande, C., Moran, S. B., Wassmann, P., Grebmeier, J. M., and Cooper, L. W.: <sup>234</sup>Th-derived particulate organic carbon fluxes in the northern Barents Sea with comparison to drifting sediment trap fluxes, J. Mar. Syst., 73, 103–113, <ext-link xlink:href="https://doi.org/10.1016/j.jmarsys.2007.09.004" ext-link-type="DOI">10.1016/j.jmarsys.2007.09.004</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib35"><label>35</label><mixed-citation>Lalande, C., Bauerfeind, E., and Nöthig, E. M.: Downward particulate organic carbon export at high temporal resolution in the eastern Fram Strait: influence of Atlantic Water on flux composition, Mar. Ecol. Prog. Ser., 440, 127–136, <ext-link xlink:href="https://doi.org/10.3354/MEPS09385" ext-link-type="DOI">10.3354/MEPS09385</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bib36"><label>36</label><mixed-citation>Lam, P. J., Ohnemus, D. C., and Auro, M. E.: Size-fractionated major particle composition and concentrations from the US GEOTRACES North Atlantic Zonal Transect, Deep-Sea Res. Pt. II, 116, 303–320, <ext-link xlink:href="https://doi.org/10.1016/J.DSR2.2014.11.020" ext-link-type="DOI">10.1016/J.DSR2.2014.11.020</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib37"><label>37</label><mixed-citation>Lam, P. J., Lee, J.-M., Heller, M. I., Mehic, S., Xiang, Y., and Bates, N. R.: Size-fractionated distributions of suspended particle concentration and major phase composition from the U.S. GEOTRACES Eastern Pacific Zonal Transect (GP16), Mar. Chem., 201, 90–107, <ext-link xlink:href="https://doi.org/10.1016/j.marchem.2017.08.013" ext-link-type="DOI">10.1016/j.marchem.2017.08.013</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib38"><label>38</label><mixed-citation>Laws, E. A. and Maiti, K.: The relationship between primary production and export production in the ocean: Effects of time lags and temporal variability, Deep-Sea Res. Pt. I, 148, 100–107, <ext-link xlink:href="https://doi.org/10.1016/J.DSR.2019.05.006" ext-link-type="DOI">10.1016/J.DSR.2019.05.006</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib39"><label>39</label><mixed-citation>Lemaitre, N.: Multi-proxy approach (Thorium-234, excess Barium) of export and remineralization fluxes of carbon and biogenic elements associated with the oceanic biological pump, Université de Bretagne Occidentale, 319 pp., <ext-link xlink:href="https://doi.org/10.70675/7b3eb896z1f81z4818zb4d4za9cac3dc2592" ext-link-type="DOI">10.70675/7b3eb896z1f81z4818zb4d4za9cac3dc2592</ext-link>,  2017.</mixed-citation></ref>
      <ref id="bib1.bib40"><label>40</label><mixed-citation>Lemaitre, N., Planchon, F., Planquette, H., Dehairs, F., Fonseca-Batista, D., Roukaerts, A., Deman, F., Tang, Y., Mariez, C., and Sarthou, G.: High variability of particulate organic carbon export along the North Atlantic GEOTRACES section GA01 as deduced from <sup>234</sup>Th fluxes, Biogeosciences, 15, 6417–6437, <ext-link xlink:href="https://doi.org/10.5194/bg-15-6417-2018" ext-link-type="DOI">10.5194/bg-15-6417-2018</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib41"><label>41</label><mixed-citation>Maiti, K., Buesseler, K. O., Pike, S. M., Benitez-Nelson, C., Cai, P., Chen, W., Cochran, K., Dai, M., Dehairs, F., Gasser, B., Kelly, R. P., Masque, P., Miller, L. A., Miquel, J. C., Moran, S. B., Morris, P. J., Peine, F., Planchon, F., Renfro, A. A., van der Loeff, M. R., Santschi, P. H., Turnewitsch, R., Waples, J. T., and Xu, C.: Intercalibration studies of short-lived thorium-234 in the water column and marine particles, Limnol. Oceanogr. Methods, 10, 631–644, <ext-link xlink:href="https://doi.org/10.4319/lom.2012.10.631" ext-link-type="DOI">10.4319/lom.2012.10.631</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib42"><label>42</label><mixed-citation>Mari, X., Passow, U., Migon, C., Burd, A. B., and Legendre, L.: Transparent exopolymer particles: Effects on carbon cycling in the ocean, Prog. Oceanogr., 151, 13–37, <ext-link xlink:href="https://doi.org/10.1016/J.POCEAN.2016.11.002" ext-link-type="DOI">10.1016/J.POCEAN.2016.11.002</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib43"><label>43</label><mixed-citation>Le Moigne, F. A. C., Poulton, A. J., Henson, S. A., Daniels, C. J., Fragoso, G. M., Mitchell, E., Richier, S., Russell, B. C., Smith, H. E. K., Tarling, G. A., Young, J. R., and Zubkov, M.: Carbon export efficiency and phytoplankton community composition in the Atlantic sector of the Arctic Ocean, J. Geophys. Res.-Ocean., 120, 3896–3912, <ext-link xlink:href="https://doi.org/10.1002/2015JC010700" ext-link-type="DOI">10.1002/2015JC010700</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib44"><label>44</label><mixed-citation>Moran, S. B., Weinstein, S. E., Edmonds, H. N., Smith, J. N., Kelly, R. P., Pilson, M. E. Q., and Harrison, W. G.: Does <sup>234</sup>Th <inline-formula><mml:math id="M579" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <sup>238</sup>U disequilibrium provide an accurate record of the export flux of particulate organic carbon from the upper ocean?, Limnol. Oceanogr., 48, 1018–1029, <ext-link xlink:href="https://doi.org/10.4319/lo.2003.48.3.1018" ext-link-type="DOI">10.4319/lo.2003.48.3.1018</ext-link>, 2003.</mixed-citation></ref>
      <ref id="bib1.bib45"><label>45</label><mixed-citation>Morris, P. J., Sanders, R., Turnewitsch, R., and Thomalla, S.: <sup>234</sup>Th-derived particulate organic carbon export from an island-induced phytoplankton bloom in the Southern Ocean, Deep-Sea Res. Pt. II, 54, 2208–2232, <ext-link xlink:href="https://doi.org/10.1016/j.dsr2.2007.06.002" ext-link-type="DOI">10.1016/j.dsr2.2007.06.002</ext-link>, 2007.</mixed-citation></ref>
      <ref id="bib1.bib46"><label>46</label><mixed-citation>Ohashi, K., Laurent, A., Renkl, C., Sheng, J., Fennel, K., and Oliver, E.: DalROMS-NWA12 v1.0, a coupled circulation-ice-biogeochemistry modelling system for the northwest Atlantic Ocean: Development and validation, Geosci. Model Dev., 17, 8697–8733, <ext-link xlink:href="https://doi.org/10.5194/GMD-17-8697-2024" ext-link-type="DOI">10.5194/GMD-17-8697-2024</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib47"><label>47</label><mixed-citation>Owens, S. A., Buesseler, K. O., and Sims, K. W. W.: Re-evaluating the <sup>238</sup>U-salinity relationship in seawater: Implications for the <sup>238</sup>U–<sup>234</sup>Th disequilibrium method, Mar. Chem., 127, 31–39, <ext-link xlink:href="https://doi.org/10.1016/j.marchem.2011.07.005" ext-link-type="DOI">10.1016/j.marchem.2011.07.005</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bib48"><label>48</label><mixed-citation>Owens, S. A., Pike, S., and Buesseler, K. O.: Thorium-234 as a tracer of particle dynamics and upper ocean export in the Atlantic Ocean, Deep-Sea Res. Pt. II, 116, 42–59, <ext-link xlink:href="https://doi.org/10.1016/j.dsr2.2014.11.010" ext-link-type="DOI">10.1016/j.dsr2.2014.11.010</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib49"><label>49</label><mixed-citation>Parsons, T. R., Maita, Y., and Lalli, C. M.: A manual of chemical and biological methods for seawater analysis, Pergamon Press, Oxford, UK, 173 pp., <ext-link xlink:href="https://doi.org/10.25607/OBP-1830" ext-link-type="DOI">10.25607/OBP-1830</ext-link>, 1984.</mixed-citation></ref>
      <ref id="bib1.bib50"><label>50</label><mixed-citation>Passow, U.: Transparent exopolymer particles (TEP) in aquatic environments, Prog. Oceanogr., 55, 287–333, <ext-link xlink:href="https://doi.org/10.1016/S0079-6611(02)00138-6" ext-link-type="DOI">10.1016/S0079-6611(02)00138-6</ext-link>, 2002.</mixed-citation></ref>
      <ref id="bib1.bib51"><label>51</label><mixed-citation>Passow, U. and Carlson, C.: The biological pump in a high CO<sub>2</sub> world, Mar. Ecol. Prog. Ser., 470, 249–271, <ext-link xlink:href="https://doi.org/10.3354/meps09985" ext-link-type="DOI">10.3354/meps09985</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib52"><label>52</label><mixed-citation>Passow, U. and Wassmann, P.: On the trophic fate of <italic>Phaeocystis pouchetii</italic> (Hariot): IV. The formation of marine snow by <italic>P. pouchetii</italic> on JSTOR, Mar. Ecol. Prog. Ser., 104, 153–161, 1994.</mixed-citation></ref>
      <ref id="bib1.bib53"><label>53</label><mixed-citation>Passow, U. and Weber, T.: The biological carbon pump, in: Treatise on Geochemistry, 3rd edn., vol. 4, Elsevier, 333–369, <ext-link xlink:href="https://doi.org/10.1016/B978-0-323-99762-1.00031-0" ext-link-type="DOI">10.1016/B978-0-323-99762-1.00031-0</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bib54"><label>54</label><mixed-citation>Puigcorbé, V., Roca-Martí, M., Masqué, P., Benitez-Nelson, C., Rutgers van der Loeff, M., Bracher, A., and Moreau, S.: Latitudinal distributions of particulate carbon export across the North Western Atlantic Ocean, Deep-Sea Res. Pt. I, 129, 116–130, <ext-link xlink:href="https://doi.org/10.1016/J.DSR.2017.08.016" ext-link-type="DOI">10.1016/J.DSR.2017.08.016</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib55"><label>55</label><mixed-citation>Puigcorbé, V., Masqué, P., and Le Moigne, F. A. C.: Global database of ratios of particulate organic carbon to thorium-234 in the ocean: improving estimates of the biological carbon pump, Earth Syst. Sci. Data, 12, 1267–1285, <ext-link xlink:href="https://doi.org/10.5194/essd-12-1267-2020" ext-link-type="DOI">10.5194/essd-12-1267-2020</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib56"><label>56</label><mixed-citation>Reigstad, M. and Wassmann, P.: Does <italic>Phaeocystis</italic> spp. contribute significantly to vertical export of organic carbon?, Biogeochemistry, 83, 217–234, <ext-link xlink:href="https://doi.org/10.1007/s10533-007-9093-3" ext-link-type="DOI">10.1007/s10533-007-9093-3</ext-link>, 2007.</mixed-citation></ref>
      <ref id="bib1.bib57"><label>57</label><mixed-citation>Reinthaler, T., Van Aken, H., Veth, C., Arístegui, J., Robinson, C., Williams, P. J. L. B., Lebaron, P., and Herndl, G. J.: Prokaryotic respiration and production in the meso- and bathypelagic realm of the eastern and western North Atlantic basin, Limnol. Oceanogr., 51, 1262–1273, <ext-link xlink:href="https://doi.org/10.4319/LO.2006.51.3.1262" ext-link-type="DOI">10.4319/LO.2006.51.3.1262</ext-link>, 2006.</mixed-citation></ref>
      <ref id="bib1.bib58"><label>58</label><mixed-citation>Resplandy, L., Martin, A. P., Le Moigne, F., Martin, P., Aquilina, A., Mémery, L., Lévy, M., and Sanders, R.: How does dynamical spatial variability impact <sup>234</sup>Th-derived estimates of organic export?, Deep-Sea Res. Pt. I, 68, 24–45, <ext-link xlink:href="https://doi.org/10.1016/j.dsr.2012.05.015" ext-link-type="DOI">10.1016/j.dsr.2012.05.015</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib59"><label>59</label><mixed-citation>Riley, J. S., Sanders, R., Marsay, C., Le Moigne, F. A. C., Achterberg, E. P., and Poulton, A. J.: The relative contribution of fast and slow sinking particles to ocean carbon export, Global Biogeochem. Cy., 26, <ext-link xlink:href="https://doi.org/10.1029/2011GB004085" ext-link-type="DOI">10.1029/2011GB004085</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib60"><label>60</label><mixed-citation>Roca-Martí, M. and Puigcorbé, V.: Combined Use of Short-Lived Radionuclides (<sup>234</sup>Th and <sup>210</sup>Po) as Tracers of Sinking Particles in the Ocean, Ann. Rev. Mar. Sci., 16, 551–575, <ext-link xlink:href="https://doi.org/10.1146/annurev-marine-041923-013807" ext-link-type="DOI">10.1146/annurev-marine-041923-013807</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib61"><label>61</label><mixed-citation>Roca-Martí, M., Puigcorbé, V., Iversen, M. H., Rutgers van der Loeff, M. M., Klaas, C., Cheah, W., Bracher, A., and Masqué, P.: High particulate organic carbon export during the decline of a vast diatom bloom in the Atlantic sector of the Southern Ocean, Deep-Sea Res. Pt. II, 138, 102–115, <ext-link xlink:href="https://doi.org/10.1016/j.dsr2.2015.12.007" ext-link-type="DOI">10.1016/j.dsr2.2015.12.007</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib62"><label>62</label><mixed-citation>Roca-Martí, M., Benitez-Nelson, C. R., Umhau, B. P., Wyatt, A. M., Clevenger, S. J., Pike, S., Horner, T. J., Estapa, M. L., Resplandy, L., and Buesseler, K. O.: Concentrations, ratios, and sinking fluxes of major bioelements at Ocean Station Papa, Elem. Sci. Anthr., 9, <ext-link xlink:href="https://doi.org/10.1525/elementa.2020.00166" ext-link-type="DOI">10.1525/elementa.2020.00166</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib63"><label>63</label><mixed-citation>Roca‐Martí, M., Healey, M., and Kienast, S. S.: Particulate Th-234, organic carbon, organic nitrogen and biogenic silica concentrations in size-fractionated particles in the Labrador Sea in spring 2022, PANGAEA [data set], <ext-link xlink:href="https://doi.org/10.1594/PANGAEA.983961" ext-link-type="DOI">10.1594/PANGAEA.983961</ext-link>, 2025a.</mixed-citation></ref>
      <ref id="bib1.bib64"><label>64</label><mixed-citation>Roca‐Martí, M., Healey, M., and Kienast, S. S.: Total (dissolved <inline-formula><mml:math id="M589" display="inline"><mml:mo>+</mml:mo></mml:math></inline-formula> particulate) Th-234 and U-238 activities in seawater in the Labrador Sea in spring 2022, PANGAEA [data set], <ext-link xlink:href="https://doi.org/10.1594/PANGAEA.983957" ext-link-type="DOI">10.1594/PANGAEA.983957</ext-link>, 2025b.</mixed-citation></ref>
      <ref id="bib1.bib65"><label>65</label><mixed-citation>Romanelli, E., Sweet, J., Giering, S. L. C., Siegel, D. A., and Passow, U.: The importance of transparent exopolymer particles over ballast in determining both sinking and suspension of small particles during late summer in the Northeast Pacific Ocean, Elementa, 11, <ext-link xlink:href="https://doi.org/10.1525/elementa.2022.00122" ext-link-type="DOI">10.1525/elementa.2022.00122</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib66"><label>66</label><mixed-citation>Romanelli, E., Stevens-Green, R., Cisternas-Novoa, C., LaRoche, J., Siegel, D. A., Carlson, C. A., and Passow, U.: Particle lability drives degradation dynamics and bacterial community assembly during a <italic>Phaeocystis</italic> bloom decline, preprint, <ext-link xlink:href="https://doi.org/10.64898/2026.04.19.716305" ext-link-type="DOI">10.64898/2026.04.19.716305</ext-link>, 2026.</mixed-citation></ref>
      <ref id="bib1.bib67"><label>67</label><mixed-citation>Salter, I., Lampitt, R. S., Sanders, R., Poulton, A., Kemp, A. E. S., Boorman, B., Saw, K., and Pearce, R.: Estimating carbon, silica and diatom export from a naturally fertilised phytoplankton bloom in the Southern Ocean using PELAGRA: A novel drifting sediment trap, Deep-Sea Res. Pt. II, 54, 2233–2259, <ext-link xlink:href="https://doi.org/10.1016/j.dsr2.2007.06.008" ext-link-type="DOI">10.1016/j.dsr2.2007.06.008</ext-link>, 2007.</mixed-citation></ref>
      <ref id="bib1.bib68"><label>68</label><mixed-citation>Sanders, R., Henson, S. A., Koski, M., De La Rocha, C. L., Painter, S. C., Poulton, A. J., Riley, J., Salihoglu, B., Visser, A., Yool, A., Bellerby, R., and Martin, A. P.: The Biological Carbon Pump in the North Atlantic, Prog. Oceanogr., 129, 200–218, <ext-link xlink:href="https://doi.org/10.1016/j.pocean.2014.05.005" ext-link-type="DOI">10.1016/j.pocean.2014.05.005</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib69"><label>69</label><mixed-citation>Savoye, N., Benitez-Nelson, C., Burd, A. B., Cochran, J. K., Charette, M., Buesseler, K. O., Jackson, G. A., Roy-Barman, M., Schmidt, S., and Elskens, M.: <sup>234</sup>Th sorption and export models in the water column: A review, Mar. Chem., 100, 234–249, <ext-link xlink:href="https://doi.org/10.1016/j.marchem.2005.10.014" ext-link-type="DOI">10.1016/j.marchem.2005.10.014</ext-link>, 2006.</mixed-citation></ref>
      <ref id="bib1.bib70"><label>70</label><mixed-citation>Savoye, N., Trull, T. W., Jacquet, S. H. M., Navez, J., and Dehairs, F.: <sup>234</sup>Th-based export fluxes during a natural iron fertilization experiment in the Southern Ocean (KEOPS), Deep-Sea Res. Pt. II, 55, 841–855, <ext-link xlink:href="https://doi.org/10.1016/j.dsr2.2007.12.036" ext-link-type="DOI">10.1016/j.dsr2.2007.12.036</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib71"><label>71</label><mixed-citation>Schoemann, V., Becquevort, S., Stefels, J., Rousseau, V., and Lancelot, C.: <italic>Phaeocystis</italic> blooms in the global ocean and their controlling mechanisms: a review, J. Sea Res., 53, 43–66, <ext-link xlink:href="https://doi.org/10.1016/J.SEARES.2004.01.008" ext-link-type="DOI">10.1016/J.SEARES.2004.01.008</ext-link>, 2005.</mixed-citation></ref>
      <ref id="bib1.bib72"><label>72</label><mixed-citation> Smetacek, V., von Bodungen, B., Knoppers, B., Peinert, R., Pollehne, F., Stegmann, P., and Zeitzschel, B.: Seasonal stages characterizing the annual cycle of an inshore pelagic system, Rapp. Proces-Verbaux des Reun. Cons. Int. pour l'Exploration la Mer, 183, 126–135, 1984.</mixed-citation></ref>
      <ref id="bib1.bib73"><label>73</label><mixed-citation>Smith, W. O. and Trimborn, S.: <italic>Phaeocystis</italic>: A Global Enigma, Ann. Rev. Mar. Sci., 16, 417–441, <ext-link xlink:href="https://doi.org/10.1146/annurev-marine-022223-025031" ext-link-type="DOI">10.1146/annurev-marine-022223-025031</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib74"><label>74</label><mixed-citation>Steinberg, D. K., Van Mooy, B. A. S., Buesseler, K. O., Boyd, P. W., Kobari, T., and Karl, D. M.: Bacterial vs. zooplankton control of sinking particle flux in the ocean's twilight zone, Limnol. Oceanogr., 53, 1327–1338, <ext-link xlink:href="https://doi.org/10.4319/lo.2008.53.4.1327" ext-link-type="DOI">10.4319/lo.2008.53.4.1327</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib75"><label>75</label><mixed-citation>Stephens, B. M., Roca-Martí, M., Maas, A. E., Amaral, V. J., Clevenger, S., Traylor, S., Benitez-Nelson, C. R., Boyd, P. W., Buesseler, K. O., Carlson, C. A., Cassar, N., Estapa, M., Fassbender, A. J., Huang, Y., Lam, P. J., Marchal, O., Menden-Deuer, S., Paul, N. L., Santoro, A. E., Siegel, D. A., and Nicholson, D. P.: An upper-mesopelagic-zone carbon budget for the subarctic North Pacific, Biogeosciences, 22, 3301–3328, <ext-link xlink:href="https://doi.org/10.5194/bg-22-3301-2025" ext-link-type="DOI">10.5194/bg-22-3301-2025</ext-link>, 2025. </mixed-citation></ref>
      <ref id="bib1.bib76"><label>76</label><mixed-citation>Stevens-Green, R., Romanelli, E., Cisternas-Novoa, C., Pollara, S., Jabre, L., Passow, U., Bertrand, E. M., LaRoche, J., Stevens-Green, R., Romanelli, E., Cisternas-Novoa, C., Pollara, S., Jabre, L., Passow, U., Bertrand, E. M., and LaRoche, J.: Exploring the Microbial Communities Involved in the Biological Carbon Pump Throughout the Decline of a Large <italic>Phaeocystis pouchetii</italic> Bloom in the Labrador Sea, in: Ocean Sciences Meeting (OSM) 2024,  2024.</mixed-citation></ref>
      <ref id="bib1.bib77"><label>77</label><mixed-citation>Tesdal, J. E., Ducklow, H. W., Goes, J. I., and Yashayaev, I.: Recent nutrient enrichment and high biological productivity in the Labrador Sea is tied to enhanced winter convection, Prog. Oceanogr., 206, 102848, <ext-link xlink:href="https://doi.org/10.1016/J.POCEAN.2022.102848" ext-link-type="DOI">10.1016/J.POCEAN.2022.102848</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib78"><label>78</label><mixed-citation>Volk, T. and Hoffert, M. I.: Ocean carbon pumps: analysis of relative strengths and efficiencies in ocean-driven atmospheric CO<sub>2</sub> changes, in: The Carbon Cycle and Atmospheric CO<sub>2</sub>: Natural Variations Archean to Present, Vol. 32, edited by: Sundquist, E. T. and Broecker, W. S., American Geophysical Union, Washington, DC, 99–110, <ext-link xlink:href="https://doi.org/10.1029/GM032" ext-link-type="DOI">10.1029/GM032</ext-link>, 1985.</mixed-citation></ref>
      <ref id="bib1.bib79"><label>79</label><mixed-citation>Wiedmann, I., Ceballos‐Romero, E., Villa‐Alfageme, M., Renner, A. H. H., Dybwad, C., Jagt, H., Svensen, C., Assmy, P., Wiktor, J. M., Tatarek, A., Różańska‐Pluta, M., and Iversen, M. H.: Arctic Observations Identify Phytoplankton Community Composition as Driver of Carbon Flux Attenuation, Geophys. Res. Lett., 47, e2020GL087465, <ext-link xlink:href="https://doi.org/10.1029/2020GL087465" ext-link-type="DOI">10.1029/2020GL087465</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib80"><label>80</label><mixed-citation>Wollenburg, J. E., Katlein, C., Nehrke, G., Nöthig, E.-M., Matthiessen, J., Wolf- Gladrow, D. A., Nikolopoulos, A., Gázquez-Sanchez, F., Rossmann, L., Assmy, P., Babin, M., Bruyant, F., Beaulieu, M., Dybwad, C., and Peeken, I.: Ballasting by cryogenic gypsum enhances carbon export in a <italic>Phaeocystis</italic> under-ice bloom, Sci. Rep., 8, 7703, <ext-link xlink:href="https://doi.org/10.1038/s41598-018-26016-0" ext-link-type="DOI">10.1038/s41598-018-26016-0</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib81"><label>81</label><mixed-citation>Yashayaev, I.: Intensification and shutdown of deep convection in the Labrador Sea were caused by changes in atmospheric and freshwater dynamics, Commun. Earth Environ., 5, <ext-link xlink:href="https://doi.org/10.1038/S43247-024-01296-9" ext-link-type="DOI">10.1038/S43247-024-01296-9</ext-link>, 2024.</mixed-citation></ref>

  </ref-list></back>
    <!--<article-title-html>Sinking particle fluxes and biological carbon pump efficiency in the Labrador Sea during a <i>Phaeocystis</i> bloom decline</article-title-html>
<abstract-html/>
<ref-html id="bib1.bib1"><label>1</label><mixed-citation>
      
Alcolombri, U., Nissan, A., Słomka, J., Charlton, S., Secchi, E., Short, I., Lee, K. S., Peaudecerf, F. J., Baumgartner, D. A., Sichert, A., Sauer, U., Sengupta, A., and Stocker, R.: Biogel scavenging slows the sinking of organic particles to the ocean depths, Nat. Commun., 161,  1–10, <a href="https://doi.org/10.1038/s41467-025-57982-5" target="_blank">https://doi.org/10.1038/s41467-025-57982-5</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib2"><label>2</label><mixed-citation>
      
Andreassen, I. J. and Wassmann, P.: Vertical flux of phytoplankton and particulate biogenic matter in the marginal ice zone of the Barents Sea in May 1993, Mar. Ecol. Prog. Ser., 170, 1–14, <a href="https://doi.org/10.3354/MEPS170001" target="_blank">https://doi.org/10.3354/MEPS170001</a>, 1998.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib3"><label>3</label><mixed-citation>
      
Armstrong, R. A., Lee, C., Hedges, J. I., Honjo, S., and Wakeham, S. G.: A new, mechanistic model for organic carbon fluxes in the ocean based on the quantitative association of POC with ballast minerals, Deep-Sea Res. Pt. II, 49, 219–236, <a href="https://doi.org/10.1016/S0967-0645(01)00101-1" target="_blank">https://doi.org/10.1016/S0967-0645(01)00101-1</a>, 2002.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib4"><label>4</label><mixed-citation>
      
Arruda, R., Atamanchuk, D., Boteler, C., and Wallace, D. W. R.: Seasonality of pCO<sub>2</sub> and air-sea CO<sub>2</sub> fluxes in the Central Labrador Sea, Front. Mar. Sci., 11, 1472697, <a href="https://doi.org/10.3389/fmars.2024.1472697" target="_blank">https://doi.org/10.3389/fmars.2024.1472697</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib5"><label>5</label><mixed-citation>
      
Asper, V. L. and Smith, W. O.: Particle fluxes during austral spring and summer in the southern Ross Sea, Antarctica, J. Geophys. Res.-Ocean., 104, 5345–5359, <a href="https://doi.org/10.1029/1998JC900067" target="_blank">https://doi.org/10.1029/1998JC900067</a>, 1999.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib6"><label>6</label><mixed-citation>
      
Baker, C. A., Martin, A. P., Yool, A., and Popova, E.: Biological Carbon Pump Sequestration Efficiency in the North Atlantic: A Leaky or a Long-Term Sink?, Global Biogeochem. Cy., 36, e2021GB007286, <a href="https://doi.org/10.1029/2021GB007286" target="_blank">https://doi.org/10.1029/2021GB007286</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib7"><label>7</label><mixed-citation>
      
Baltar, F., Arístegui, J., Gasol, J. M., Sintes, E., and Herndl, G. J.: Evidence of prokaryotic metabolism on suspended particulate organic matter in the dark waters of the subtropical North Atlantic, Limnol. Oceanogr., 54, 182–193, <a href="https://doi.org/10.4319/LO.2009.54.1.0182" target="_blank">https://doi.org/10.4319/LO.2009.54.1.0182</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib8"><label>8</label><mixed-citation>
      
Baumas, C., Fuchs, R., Garel, M., Poggiale, J.-C., Memery, L., Le Moigne, F. A. C., and Tamburini, C.: Reconstructing the ocean's mesopelagic zone carbon budget: sensitivity and estimation of parameters associated with prokaryotic remineralization, Biogeosciences, 20, 4165–4182, <a href="https://doi.org/10.5194/BG-20-4165-2023" target="_blank">https://doi.org/10.5194/BG-20-4165-2023</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib9"><label>9</label><mixed-citation>
      
Bishop, J. K. B., Lam, P. J., and Wood, T. J.: Getting good particles: Accurate sampling of particles by large volume in-situ filtration, Limnol. Oceanogr. Method., 10, 681–710, <a href="https://doi.org/10.4319/lom.2012.10.681" target="_blank">https://doi.org/10.4319/lom.2012.10.681</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib10"><label>10</label><mixed-citation>
      
Boyd, P. W., Sherry, N. D., Berges, J. A., Bishop, J. K. B., Calvert, S. E., Charette, M. A., Giovannoni, S. J., Goldblatt, R., Harrison, P. J., Moran, S. B., Roy, S., Soon, M., Strom, S., Thibault, D., Vergin, K. L., Whitney, F. A., and Wong, C. S.: Transformations of biogenic particulates from the pelagic to the deep ocean realm, Deep-Sea Res. Pt. II, 46, 2761–2792, <a href="https://doi.org/10.1016/S0967-0645(99)00083-1" target="_blank">https://doi.org/10.1016/S0967-0645(99)00083-1</a>, 1999.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib11"><label>11</label><mixed-citation>
      
Boyd, P. W., Claustre, H., Levy, M., Siegel, D. A., and Weber, T.: Multi-faceted particle pumps drive carbon sequestration in the ocean, Nature, 568, 327–335, <a href="https://doi.org/10.1038/s41586-019-1098-2" target="_blank">https://doi.org/10.1038/s41586-019-1098-2</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib12"><label>12</label><mixed-citation>
      
Buesseler, K. O. and Boyd, P. W.: Shedding light on processes that control particle export and flux attenuation in the twilight zone of the open ocean, Limnol. Oceanogr., 54, 1210–1232, <a href="https://doi.org/10.4319/lo.2009.54.4.1210" target="_blank">https://doi.org/10.4319/lo.2009.54.4.1210</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib13"><label>13</label><mixed-citation>
      
Buesseler, K. O., Boyd, P. W., Black, E. E., and Siegel, D. A.: Metrics that matter for assessing the ocean biological carbon pump, P. Natl. Acad. Sci. USA,  117, 9679–9687, <a href="https://doi.org/10.1073/PNAS.1918114117" target="_blank">https://doi.org/10.1073/PNAS.1918114117</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib14"><label>14</label><mixed-citation>
      
Burd, A. B., Hansell, D. A., Steinberg, D. K., Anderson, T. R., Arístegui, J., Baltar, F., Beaupré, S. R., Buesseler, K. O., DeHairs, F., Jackson, G. A., Kadko, D. C., Koppelmann, R., Lampitt, R. S., Nagata, T., Reinthaler, T., Robinson, C., Robison, B. H., Tamburini, C., and Tanaka, T.: Assessing the apparent imbalance between geochemical and biochemical indicators of meso- and bathypelagic biological activity: What the @$♯! is wrong with present calculations of carbon budgets?, Deep-Sea Res. Pt. II, 57, 1557–1571, <a href="https://doi.org/10.1016/j.dsr2.2010.02.022" target="_blank">https://doi.org/10.1016/j.dsr2.2010.02.022</a>, 2010.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib15"><label>15</label><mixed-citation>
      
Ceballos-Romero, E., De Soto, F., Le Moigne, F. A. C., García-Tenorio, R., and Villa-Alfageme, M.: <sup>234</sup>Th-derived particle fluxes and seasonal variability: when is the SS assumption reliable? Insights from a novel approach for carbon flux simulation, Geophys. Res. Lett., 45, 13414–13426, <a href="https://doi.org/10.1029/2018GL079968" target="_blank">https://doi.org/10.1029/2018GL079968</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib16"><label>16</label><mixed-citation>
      
Cisternas-Novoa, C., Romanelli, E., and Passow, U.: Differences between suspended and sinking particles regulate carbon flux in the upper mesopelagic during a <i>Phaeocystis</i> Bloom, preprint, <a href="https://doi.org/10.64898/2026.06.09.731151" target="_blank">https://doi.org/10.64898/2026.06.09.731151</a>, 2026.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib17"><label>17</label><mixed-citation>
      
Claustre, H., Legendre, L., Boyd, P. W., and Levy, M.: The Oceans' Biological Carbon Pumps: Framework for a Research Observational Community Approach, Front. Mar. Sci., 8, <a href="https://doi.org/10.3389/fmars.2021.780052" target="_blank">https://doi.org/10.3389/fmars.2021.780052</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib18"><label>18</label><mixed-citation>
      
Clevenger, S. J., Benitez-Nelson, C. R., Drysdale, J., Pike, S., Puigcorbé, V., and Buesseler, K. O.: Review of the analysis of <sup>234</sup>Th in small volume (2–4&thinsp;L) seawater samples: improvements and recommendations, J. Radioanal. Nucl. Chem., 329, 1–13, <a href="https://doi.org/10.1007/s10967-021-07772-2" target="_blank">https://doi.org/10.1007/s10967-021-07772-2</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib19"><label>19</label><mixed-citation>
      
Clevenger, S. J., Benitez-Nelson, C. R., Roca-Martí, M., Bam, W., Estapa, M., Kenyon, J. A., Pike, S., Resplandy, L., Wyatt, A., and Buesseler, K. O.: Carbon and silica fluxes during a declining North Atlantic spring bloom as part of the EXPORTS program, Mar. Chem., 258, 104346, <a href="https://doi.org/10.1016/J.MARCHEM.2023.104346" target="_blank">https://doi.org/10.1016/J.MARCHEM.2023.104346</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib20"><label>20</label><mixed-citation>
      
Coppola, L., Roy-Barman, M., Wassmann, P., Mulsow, S., and Jeandel, C.: Calibration of sediment traps and particulate organic carbon export using <sup>234</sup>Th in the Barents Sea, Mar. Chem., 80, 11–26, <a href="https://doi.org/10.1016/S0304-4203(02)00071-3" target="_blank">https://doi.org/10.1016/S0304-4203(02)00071-3</a>, 2002.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib21"><label>21</label><mixed-citation>
      
de Boyer Montégut, C., Madec, G., Fischer, A. S., Lazar, A., and Iudicone, D.: Mixed layer depth over the global ocean: An examination of profile data and a profile-based climatology, J. Geophys. Res., 109, C12003, <a href="https://doi.org/10.1029/2004JC002378" target="_blank">https://doi.org/10.1029/2004JC002378</a>, 2004.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib22"><label>22</label><mixed-citation>
      
Devred, E., Wilson, K. L., Perry, T., Hardy, M., Brosnahan, M., and Ringuette, M.: Identification and validation of phytoplankton taxonomic assemblages derived from pigment signatures using samples collected in the Labrador Sea from 2014 to 2022, Dartmouth, Nova Scotia, Canada, 37 pp., ISBN: 978-0-660-70848-5, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib23"><label>23</label><mixed-citation>
      
Devred, E., Clay, S., Ringuette, M., Perry, T., Amirian, M., Irwin, A., and Finkel, Z.: Net primary production in the Labrador Sea between 2014 and 2022 derived from ocean colour remote sensing based on ecological regimes, Remote Sens. Environ., 323, 114713, <a href="https://doi.org/10.1016/J.RSE.2025.114713" target="_blank">https://doi.org/10.1016/J.RSE.2025.114713</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib24"><label>24</label><mixed-citation>
      
Dybwad, C., Assmy, P., Olsen, L. M., Peeken, I., Nikolopoulos, A., Krumpen, T., Randelhoff, A., Tatarek, A., Wiktor, J. M., and Reigstad, M.: Carbon Export in the Seasonal Sea Ice Zone North of Svalbard From Winter to Late Summer, Front. Mar. Sci., 7, 525800, <a href="https://doi.org/10.3389/fmars.2020.525800" target="_blank">https://doi.org/10.3389/fmars.2020.525800</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib25"><label>25</label><mixed-citation>
      
Finkel, Z. V., Beardall, J., Flynn, K. J., Quigg, A., Rees, T. A. V., and Raven, J. A.: Phytoplankton in a changing world: cell size and elemental stoichiometry, J. Plankton Res., 32, 119–137, <a href="https://doi.org/10.1093/PLANKT/FBP098" target="_blank">https://doi.org/10.1093/PLANKT/FBP098</a>, 2010.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib26"><label>26</label><mixed-citation>
      
Giering, S. L. C., Sanders, R., Lampitt, R. S., Anderson, T. R., Tamburini, C., Boutrif, M., Zubkov, M. V, Marsay, C. M., Henson, S. A., Saw, K., Cook, K., and Mayor, D. J.: Reconciliation of the carbon budget in the ocean's twilight zone, Nature, 507, 480–483, <a href="https://doi.org/10.1038/nature13123" target="_blank">https://doi.org/10.1038/nature13123</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib27"><label>27</label><mixed-citation>
      
Giering, S. L. C., Sanders, R., Martin, A. P., Lindemann, C., Möller, K. O., Daniels, C. J., Mayor, D. J., and St. John, M. A.: High export via small particles before the onset of the North Atlantic spring bloom, J. Geophys. Res.-Ocean., 121, 6929–6945, <a href="https://doi.org/10.1002/2016JC012048" target="_blank">https://doi.org/10.1002/2016JC012048</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib28"><label>28</label><mixed-citation>
      
Graff, J. R., Nelson, N. B., Roca-Martí, M., Romanelli, E., Kramer, S. J., Erickson, Z., Cetinic, I., Buesseler, K. O., Passow, U., Zhang, X., Benitez-Nelson, C., Bisson, K., Close, H. G., Crockford, T., Fox, J., Halewood, S., Lam, P., Roesler, C., Sweet, J., VerWey, B., Xiong, Y., and Siegel, D. A.: Reconciliation of total particulate organic carbon and nitrogen measurements determined using contrasting methods in the North Pacific Ocean as part of the NASA EXPORTS field campaign, Elem. Sci. Anthr., 11, <a href="https://doi.org/10.1525/elementa.2022.00112" target="_blank">https://doi.org/10.1525/elementa.2022.00112</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib29"><label>29</label><mixed-citation>
      
Hama, T., Miyazaki, T., Ogawa, Y., Iwakuma, T., Takahashi, M., Otsuki, A., and Ichimura, S.: Measurement of photosynthetic production of a marine phytoplankton population using a stable <sup>13</sup>C isotope, Mar. Biol., 73, 31–36, <a href="https://doi.org/10.1007/BF00396282" target="_blank">https://doi.org/10.1007/BF00396282</a>, 1983.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib30"><label>30</label><mixed-citation>
      
Henson, S. A., Yool, A., and Sanders, R.: Variability in efficiency of particulate organic carbon export: A model study, Global Biogeochem. Cy., 29, 33–45, <a href="https://doi.org/10.1002/2014GB004965" target="_blank">https://doi.org/10.1002/2014GB004965</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib31"><label>31</label><mixed-citation>
      
Henson, S. A., Laufkötter, C., Leung, S., Giering, S. L. C., Palevsky, H. I., and Cavan, E. L.: Uncertain response of ocean biological carbon export in a changing world, Nat. Geosci. 2022 154, 15, 248–254, <a href="https://doi.org/10.1038/s41561-022-00927-0" target="_blank">https://doi.org/10.1038/s41561-022-00927-0</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib32"><label>32</label><mixed-citation>
      
Kiørboe, T., Hansen, J. L. S., Alldredge, A. L., Jackson, G. A., Passow, U., Dam, H. G., Drapeau, D. T., Waite, A., and Garcia, C. M.: Sedimentation of phytoplankton during a diatom bloom: Rates and mechanisms, J. Mar. Res., 54, 1123–148, 1996.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib33"><label>33</label><mixed-citation>
      
Knap, A., Michaels, A., Close, A., Ducklow, H., and Dickson, A.: Protocols for the Joint Global Ocean Flux Study (JGOFS) Core Measurements, JGOFS Report Nr. 19, 1–210,  1996.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib34"><label>34</label><mixed-citation>
      
Lalande, C., Moran, S. B., Wassmann, P., Grebmeier, J. M., and Cooper, L. W.: <sup>234</sup>Th-derived particulate organic carbon fluxes in the northern Barents Sea with comparison to drifting sediment trap fluxes, J. Mar. Syst., 73, 103–113, <a href="https://doi.org/10.1016/j.jmarsys.2007.09.004" target="_blank">https://doi.org/10.1016/j.jmarsys.2007.09.004</a>, 2008.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib35"><label>35</label><mixed-citation>
      
Lalande, C., Bauerfeind, E., and Nöthig, E. M.: Downward particulate organic carbon export at high temporal resolution in the eastern Fram Strait: influence of Atlantic Water on flux composition, Mar. Ecol. Prog. Ser., 440, 127–136, <a href="https://doi.org/10.3354/MEPS09385" target="_blank">https://doi.org/10.3354/MEPS09385</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib36"><label>36</label><mixed-citation>
      
Lam, P. J., Ohnemus, D. C., and Auro, M. E.: Size-fractionated major particle composition and concentrations from the US GEOTRACES North Atlantic Zonal Transect, Deep-Sea Res. Pt. II, 116, 303–320, <a href="https://doi.org/10.1016/J.DSR2.2014.11.020" target="_blank">https://doi.org/10.1016/J.DSR2.2014.11.020</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib37"><label>37</label><mixed-citation>
      
Lam, P. J., Lee, J.-M., Heller, M. I., Mehic, S., Xiang, Y., and Bates, N. R.: Size-fractionated distributions of suspended particle concentration and major phase composition from the U.S. GEOTRACES Eastern Pacific Zonal Transect (GP16), Mar. Chem., 201, 90–107, <a href="https://doi.org/10.1016/j.marchem.2017.08.013" target="_blank">https://doi.org/10.1016/j.marchem.2017.08.013</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib38"><label>38</label><mixed-citation>
      
Laws, E. A. and Maiti, K.: The relationship between primary production and export production in the ocean: Effects of time lags and temporal variability, Deep-Sea Res. Pt. I, 148, 100–107, <a href="https://doi.org/10.1016/J.DSR.2019.05.006" target="_blank">https://doi.org/10.1016/J.DSR.2019.05.006</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib39"><label>39</label><mixed-citation>
      
Lemaitre, N.: Multi-proxy approach (Thorium-234, excess Barium) of export and remineralization fluxes of carbon and biogenic elements associated with the oceanic biological pump, Université de Bretagne Occidentale, 319 pp., <a href="https://doi.org/10.70675/7b3eb896z1f81z4818zb4d4za9cac3dc2592" target="_blank">https://doi.org/10.70675/7b3eb896z1f81z4818zb4d4za9cac3dc2592</a>,  2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib40"><label>40</label><mixed-citation>
      
Lemaitre, N., Planchon, F., Planquette, H., Dehairs, F., Fonseca-Batista, D., Roukaerts, A., Deman, F., Tang, Y., Mariez, C., and Sarthou, G.: High variability of particulate organic carbon export along the North Atlantic GEOTRACES section GA01 as deduced from <sup>234</sup>Th fluxes, Biogeosciences, 15, 6417–6437, <a href="https://doi.org/10.5194/bg-15-6417-2018" target="_blank">https://doi.org/10.5194/bg-15-6417-2018</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib41"><label>41</label><mixed-citation>
      
Maiti, K., Buesseler, K. O., Pike, S. M., Benitez-Nelson, C., Cai, P., Chen, W., Cochran, K., Dai, M., Dehairs, F., Gasser, B., Kelly, R. P., Masque, P., Miller, L. A., Miquel, J. C., Moran, S. B., Morris, P. J., Peine, F., Planchon, F., Renfro, A. A., van der Loeff, M. R., Santschi, P. H., Turnewitsch, R., Waples, J. T., and Xu, C.: Intercalibration studies of short-lived thorium-234 in the water column and marine particles, Limnol. Oceanogr. Methods, 10, 631–644, <a href="https://doi.org/10.4319/lom.2012.10.631" target="_blank">https://doi.org/10.4319/lom.2012.10.631</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib42"><label>42</label><mixed-citation>
      
Mari, X., Passow, U., Migon, C., Burd, A. B., and Legendre, L.: Transparent exopolymer particles: Effects on carbon cycling in the ocean, Prog. Oceanogr., 151, 13–37, <a href="https://doi.org/10.1016/J.POCEAN.2016.11.002" target="_blank">https://doi.org/10.1016/J.POCEAN.2016.11.002</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib43"><label>43</label><mixed-citation>
      
Le Moigne, F. A. C., Poulton, A. J., Henson, S. A., Daniels, C. J., Fragoso, G. M., Mitchell, E., Richier, S., Russell, B. C., Smith, H. E. K., Tarling, G. A., Young, J. R., and Zubkov, M.: Carbon export efficiency and phytoplankton community composition in the Atlantic sector of the Arctic Ocean, J. Geophys. Res.-Ocean., 120, 3896–3912, <a href="https://doi.org/10.1002/2015JC010700" target="_blank">https://doi.org/10.1002/2015JC010700</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib44"><label>44</label><mixed-citation>
      
Moran, S. B., Weinstein, S. E., Edmonds, H. N., Smith, J. N., Kelly, R. P., Pilson, M. E. Q., and Harrison, W. G.: Does <sup>234</sup>Th&thinsp;∕&thinsp;<sup>238</sup>U disequilibrium provide an accurate record of the export flux of particulate organic carbon from the upper ocean?, Limnol. Oceanogr., 48, 1018–1029, <a href="https://doi.org/10.4319/lo.2003.48.3.1018" target="_blank">https://doi.org/10.4319/lo.2003.48.3.1018</a>, 2003.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib45"><label>45</label><mixed-citation>
      
Morris, P. J., Sanders, R., Turnewitsch, R., and Thomalla, S.: <sup>234</sup>Th-derived particulate organic carbon export from an island-induced phytoplankton bloom in the Southern Ocean, Deep-Sea Res. Pt. II, 54, 2208–2232, <a href="https://doi.org/10.1016/j.dsr2.2007.06.002" target="_blank">https://doi.org/10.1016/j.dsr2.2007.06.002</a>, 2007.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib46"><label>46</label><mixed-citation>
      
Ohashi, K., Laurent, A., Renkl, C., Sheng, J., Fennel, K., and Oliver, E.: DalROMS-NWA12 v1.0, a coupled circulation-ice-biogeochemistry modelling system for the northwest Atlantic Ocean: Development and validation, Geosci. Model Dev., 17, 8697–8733, <a href="https://doi.org/10.5194/GMD-17-8697-2024" target="_blank">https://doi.org/10.5194/GMD-17-8697-2024</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib47"><label>47</label><mixed-citation>
      
Owens, S. A., Buesseler, K. O., and Sims, K. W. W.: Re-evaluating the <sup>238</sup>U-salinity relationship in seawater: Implications for the <sup>238</sup>U–<sup>234</sup>Th disequilibrium method, Mar. Chem., 127, 31–39, <a href="https://doi.org/10.1016/j.marchem.2011.07.005" target="_blank">https://doi.org/10.1016/j.marchem.2011.07.005</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib48"><label>48</label><mixed-citation>
      
Owens, S. A., Pike, S., and Buesseler, K. O.: Thorium-234 as a tracer of particle dynamics and upper ocean export in the Atlantic Ocean, Deep-Sea Res. Pt. II, 116, 42–59, <a href="https://doi.org/10.1016/j.dsr2.2014.11.010" target="_blank">https://doi.org/10.1016/j.dsr2.2014.11.010</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib49"><label>49</label><mixed-citation>
      
Parsons, T. R., Maita, Y., and Lalli, C. M.: A manual of chemical and biological methods for seawater analysis, Pergamon Press, Oxford, UK, 173 pp., <a href="https://doi.org/10.25607/OBP-1830" target="_blank">https://doi.org/10.25607/OBP-1830</a>, 1984.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib50"><label>50</label><mixed-citation>
      
Passow, U.: Transparent exopolymer particles (TEP) in aquatic environments, Prog. Oceanogr., 55, 287–333, <a href="https://doi.org/10.1016/S0079-6611(02)00138-6" target="_blank">https://doi.org/10.1016/S0079-6611(02)00138-6</a>, 2002.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib51"><label>51</label><mixed-citation>
      
Passow, U. and Carlson, C.: The biological pump in a high CO<sub>2</sub> world, Mar. Ecol. Prog. Ser., 470, 249–271, <a href="https://doi.org/10.3354/meps09985" target="_blank">https://doi.org/10.3354/meps09985</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib52"><label>52</label><mixed-citation>
      
Passow, U. and Wassmann, P.: On the trophic fate of <i>Phaeocystis pouchetii</i> (Hariot): IV. The formation of marine snow by <i>P. pouchetii</i> on JSTOR, Mar. Ecol. Prog. Ser., 104, 153–161, 1994.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib53"><label>53</label><mixed-citation>
      
Passow, U. and Weber, T.: The biological carbon pump, in: Treatise on Geochemistry, 3rd edn., vol. 4, Elsevier, 333–369, <a href="https://doi.org/10.1016/B978-0-323-99762-1.00031-0" target="_blank">https://doi.org/10.1016/B978-0-323-99762-1.00031-0</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib54"><label>54</label><mixed-citation>
      
Puigcorbé, V., Roca-Martí, M., Masqué, P., Benitez-Nelson, C., Rutgers van der Loeff, M., Bracher, A., and Moreau, S.: Latitudinal distributions of particulate carbon export across the North Western Atlantic Ocean, Deep-Sea Res. Pt. I, 129, 116–130, <a href="https://doi.org/10.1016/J.DSR.2017.08.016" target="_blank">https://doi.org/10.1016/J.DSR.2017.08.016</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib55"><label>55</label><mixed-citation>
      
Puigcorbé, V., Masqué, P., and Le Moigne, F. A. C.: Global database of ratios of particulate organic carbon to thorium-234 in the ocean: improving estimates of the biological carbon pump, Earth Syst. Sci. Data, 12, 1267–1285, <a href="https://doi.org/10.5194/essd-12-1267-2020" target="_blank">https://doi.org/10.5194/essd-12-1267-2020</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib56"><label>56</label><mixed-citation>
      
Reigstad, M. and Wassmann, P.: Does <i>Phaeocystis</i> spp. contribute significantly to vertical export of organic carbon?, Biogeochemistry, 83, 217–234, <a href="https://doi.org/10.1007/s10533-007-9093-3" target="_blank">https://doi.org/10.1007/s10533-007-9093-3</a>, 2007.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib57"><label>57</label><mixed-citation>
      
Reinthaler, T., Van Aken, H., Veth, C., Arístegui, J., Robinson, C., Williams, P. J. L. B., Lebaron, P., and Herndl, G. J.: Prokaryotic respiration and production in the meso- and bathypelagic realm of the eastern and western North Atlantic basin, Limnol. Oceanogr., 51, 1262–1273, <a href="https://doi.org/10.4319/LO.2006.51.3.1262" target="_blank">https://doi.org/10.4319/LO.2006.51.3.1262</a>, 2006.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib58"><label>58</label><mixed-citation>
      
Resplandy, L., Martin, A. P., Le Moigne, F., Martin, P., Aquilina, A., Mémery, L., Lévy, M., and Sanders, R.: How does dynamical spatial variability impact <sup>234</sup>Th-derived estimates of organic export?, Deep-Sea Res. Pt. I, 68, 24–45, <a href="https://doi.org/10.1016/j.dsr.2012.05.015" target="_blank">https://doi.org/10.1016/j.dsr.2012.05.015</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib59"><label>59</label><mixed-citation>
      
Riley, J. S., Sanders, R., Marsay, C., Le Moigne, F. A. C., Achterberg, E. P., and Poulton, A. J.: The relative contribution of fast and slow sinking particles to ocean carbon export, Global Biogeochem. Cy., 26, <a href="https://doi.org/10.1029/2011GB004085" target="_blank">https://doi.org/10.1029/2011GB004085</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib60"><label>60</label><mixed-citation>
      
Roca-Martí, M. and Puigcorbé, V.: Combined Use of Short-Lived Radionuclides (<sup>234</sup>Th and <sup>210</sup>Po) as Tracers of Sinking Particles in the Ocean, Ann. Rev. Mar. Sci., 16, 551–575, <a href="https://doi.org/10.1146/annurev-marine-041923-013807" target="_blank">https://doi.org/10.1146/annurev-marine-041923-013807</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib61"><label>61</label><mixed-citation>
      
Roca-Martí, M., Puigcorbé, V., Iversen, M. H., Rutgers van der Loeff, M. M., Klaas, C., Cheah, W., Bracher, A., and Masqué, P.: High particulate organic carbon export during the decline of a vast diatom bloom in the Atlantic sector of the Southern Ocean, Deep-Sea Res. Pt. II, 138, 102–115, <a href="https://doi.org/10.1016/j.dsr2.2015.12.007" target="_blank">https://doi.org/10.1016/j.dsr2.2015.12.007</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib62"><label>62</label><mixed-citation>
      
Roca-Martí, M., Benitez-Nelson, C. R., Umhau, B. P., Wyatt, A. M., Clevenger, S. J., Pike, S., Horner, T. J., Estapa, M. L., Resplandy, L., and Buesseler, K. O.: Concentrations, ratios, and sinking fluxes of major bioelements at Ocean Station Papa, Elem. Sci. Anthr., 9, <a href="https://doi.org/10.1525/elementa.2020.00166" target="_blank">https://doi.org/10.1525/elementa.2020.00166</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib63"><label>63</label><mixed-citation>
      
Roca‐Martí, M., Healey, M., and Kienast, S. S.: Particulate Th-234, organic carbon, organic nitrogen and biogenic silica concentrations in size-fractionated particles in the Labrador Sea in spring 2022, PANGAEA [data set], <a href="https://doi.org/10.1594/PANGAEA.983961" target="_blank">https://doi.org/10.1594/PANGAEA.983961</a>, 2025a.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib64"><label>64</label><mixed-citation>
      
Roca‐Martí, M., Healey, M., and Kienast, S. S.: Total (dissolved&thinsp;+&thinsp;particulate) Th-234 and U-238 activities in seawater in the Labrador Sea in spring 2022, PANGAEA [data set], <a href="https://doi.org/10.1594/PANGAEA.983957" target="_blank">https://doi.org/10.1594/PANGAEA.983957</a>, 2025b.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib65"><label>65</label><mixed-citation>
      
Romanelli, E., Sweet, J., Giering, S. L. C., Siegel, D. A., and Passow, U.: The importance of transparent exopolymer particles over ballast in determining both sinking and suspension of small particles during late summer in the Northeast Pacific Ocean, Elementa, 11, <a href="https://doi.org/10.1525/elementa.2022.00122" target="_blank">https://doi.org/10.1525/elementa.2022.00122</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib66"><label>66</label><mixed-citation>
      
Romanelli, E., Stevens-Green, R., Cisternas-Novoa, C., LaRoche, J., Siegel, D. A., Carlson, C. A., and Passow, U.: Particle lability drives degradation dynamics and bacterial community assembly during a <i>Phaeocystis</i> bloom decline, preprint, <a href="https://doi.org/10.64898/2026.04.19.716305" target="_blank">https://doi.org/10.64898/2026.04.19.716305</a>, 2026.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib67"><label>67</label><mixed-citation>
      
Salter, I., Lampitt, R. S., Sanders, R., Poulton, A., Kemp, A. E. S., Boorman, B., Saw, K., and Pearce, R.: Estimating carbon, silica and diatom export from a naturally fertilised phytoplankton bloom in the Southern Ocean using PELAGRA: A novel drifting sediment trap, Deep-Sea Res. Pt. II, 54, 2233–2259, <a href="https://doi.org/10.1016/j.dsr2.2007.06.008" target="_blank">https://doi.org/10.1016/j.dsr2.2007.06.008</a>, 2007.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib68"><label>68</label><mixed-citation>
      
Sanders, R., Henson, S. A., Koski, M., De La Rocha, C. L., Painter, S. C., Poulton, A. J., Riley, J., Salihoglu, B., Visser, A., Yool, A., Bellerby, R., and Martin, A. P.: The Biological Carbon Pump in the North Atlantic, Prog. Oceanogr., 129, 200–218, <a href="https://doi.org/10.1016/j.pocean.2014.05.005" target="_blank">https://doi.org/10.1016/j.pocean.2014.05.005</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib69"><label>69</label><mixed-citation>
      
Savoye, N., Benitez-Nelson, C., Burd, A. B., Cochran, J. K., Charette, M., Buesseler, K. O., Jackson, G. A., Roy-Barman, M., Schmidt, S., and Elskens, M.: <sup>234</sup>Th sorption and export models in the water column: A review, Mar. Chem., 100, 234–249, <a href="https://doi.org/10.1016/j.marchem.2005.10.014" target="_blank">https://doi.org/10.1016/j.marchem.2005.10.014</a>, 2006.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib70"><label>70</label><mixed-citation>
      
Savoye, N., Trull, T. W., Jacquet, S. H. M., Navez, J., and Dehairs, F.: <sup>234</sup>Th-based export fluxes during a natural iron fertilization experiment in the Southern Ocean (KEOPS), Deep-Sea Res. Pt. II, 55, 841–855, <a href="https://doi.org/10.1016/j.dsr2.2007.12.036" target="_blank">https://doi.org/10.1016/j.dsr2.2007.12.036</a>, 2008.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib71"><label>71</label><mixed-citation>
      
Schoemann, V., Becquevort, S., Stefels, J., Rousseau, V., and Lancelot, C.: <i>Phaeocystis</i> blooms in the global ocean and their controlling mechanisms: a review, J. Sea Res., 53, 43–66, <a href="https://doi.org/10.1016/J.SEARES.2004.01.008" target="_blank">https://doi.org/10.1016/J.SEARES.2004.01.008</a>, 2005.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib72"><label>72</label><mixed-citation>
      
Smetacek, V., von Bodungen, B., Knoppers, B., Peinert, R., Pollehne, F., Stegmann, P., and Zeitzschel, B.: Seasonal stages characterizing the annual cycle of an inshore pelagic system, Rapp. Proces-Verbaux des Reun. Cons. Int. pour l'Exploration la Mer, 183, 126–135, 1984.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib73"><label>73</label><mixed-citation>
      
Smith, W. O. and Trimborn, S.: <i>Phaeocystis</i>: A Global Enigma, Ann. Rev. Mar. Sci., 16, 417–441, <a href="https://doi.org/10.1146/annurev-marine-022223-025031" target="_blank">https://doi.org/10.1146/annurev-marine-022223-025031</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib74"><label>74</label><mixed-citation>
      
Steinberg, D. K., Van Mooy, B. A. S., Buesseler, K. O., Boyd, P. W., Kobari, T., and Karl, D. M.: Bacterial vs. zooplankton control of sinking particle flux in the ocean's twilight zone, Limnol. Oceanogr., 53, 1327–1338, <a href="https://doi.org/10.4319/lo.2008.53.4.1327" target="_blank">https://doi.org/10.4319/lo.2008.53.4.1327</a>, 2008.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib75"><label>75</label><mixed-citation>
      
Stephens, B. M., Roca-Martí, M., Maas, A. E., Amaral, V. J., Clevenger, S., Traylor, S., Benitez-Nelson, C. R., Boyd, P. W., Buesseler, K. O., Carlson, C. A., Cassar, N., Estapa, M., Fassbender, A. J., Huang, Y., Lam, P. J., Marchal, O., Menden-Deuer, S., Paul, N. L., Santoro, A. E., Siegel, D. A., and Nicholson, D. P.: An upper-mesopelagic-zone carbon budget for the subarctic North Pacific, Biogeosciences, 22, 3301–3328, <a href="https://doi.org/10.5194/bg-22-3301-2025" target="_blank">https://doi.org/10.5194/bg-22-3301-2025</a>, 2025.


    </mixed-citation></ref-html>
<ref-html id="bib1.bib76"><label>76</label><mixed-citation>
      
Stevens-Green, R., Romanelli, E., Cisternas-Novoa, C., Pollara, S., Jabre, L., Passow, U., Bertrand, E. M., LaRoche, J., Stevens-Green, R., Romanelli, E., Cisternas-Novoa, C., Pollara, S., Jabre, L., Passow, U., Bertrand, E. M., and LaRoche, J.: Exploring the Microbial Communities Involved in the Biological Carbon Pump Throughout the Decline of a Large <i>Phaeocystis pouchetii</i> Bloom in the Labrador Sea, in: Ocean Sciences Meeting (OSM) 2024,  2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib77"><label>77</label><mixed-citation>
      
Tesdal, J. E., Ducklow, H. W., Goes, J. I., and Yashayaev, I.: Recent nutrient enrichment and high biological productivity in the Labrador Sea is tied to enhanced winter convection, Prog. Oceanogr., 206, 102848, <a href="https://doi.org/10.1016/J.POCEAN.2022.102848" target="_blank">https://doi.org/10.1016/J.POCEAN.2022.102848</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib78"><label>78</label><mixed-citation>
      
Volk, T. and Hoffert, M. I.: Ocean carbon pumps: analysis of relative strengths and efficiencies in ocean-driven atmospheric CO<sub>2</sub> changes, in: The Carbon Cycle and Atmospheric CO<sub>2</sub>: Natural Variations Archean to Present, Vol. 32, edited by: Sundquist, E. T. and Broecker, W. S., American Geophysical Union, Washington, DC, 99–110, <a href="https://doi.org/10.1029/GM032" target="_blank">https://doi.org/10.1029/GM032</a>, 1985.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib79"><label>79</label><mixed-citation>
      
Wiedmann, I., Ceballos‐Romero, E., Villa‐Alfageme, M., Renner, A. H. H., Dybwad, C., Jagt, H., Svensen, C., Assmy, P., Wiktor, J. M., Tatarek, A., Różańska‐Pluta, M., and Iversen, M. H.: Arctic Observations Identify Phytoplankton Community Composition as Driver of Carbon Flux Attenuation, Geophys. Res. Lett., 47, e2020GL087465, <a href="https://doi.org/10.1029/2020GL087465" target="_blank">https://doi.org/10.1029/2020GL087465</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib80"><label>80</label><mixed-citation>
      
Wollenburg, J. E., Katlein, C., Nehrke, G., Nöthig, E.-M., Matthiessen, J., Wolf- Gladrow, D. A., Nikolopoulos, A., Gázquez-Sanchez, F., Rossmann, L., Assmy, P., Babin, M., Bruyant, F., Beaulieu, M., Dybwad, C., and Peeken, I.: Ballasting by cryogenic gypsum enhances carbon export in a <i>Phaeocystis</i> under-ice bloom, Sci. Rep., 8, 7703, <a href="https://doi.org/10.1038/s41598-018-26016-0" target="_blank">https://doi.org/10.1038/s41598-018-26016-0</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib81"><label>81</label><mixed-citation>
      
Yashayaev, I.: Intensification and shutdown of deep convection in the Labrador Sea were caused by changes in atmospheric and freshwater dynamics, Commun. Earth Environ., 5, <a href="https://doi.org/10.1038/S43247-024-01296-9" target="_blank">https://doi.org/10.1038/S43247-024-01296-9</a>, 2024.

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