Marine particle forensics frequently uncovers information on particle composition, age, size, and ecological history. Zooplankton fecal pellets (FPs) are also interrogated for process-related data, such as grazing rates and carbon sequestration potential. Here, flow cytometric analyses of FP contents revealed intact phytoplankton with photophysiological characteristics mirroring those of free-living cells. Mapping the cytometrically derived properties of Synechococcus cells inside FPs onto vertical profiles from free-living cells revealed the potential to estimate depth-specific grazing by individual zooplankton. An experiment conducted at sea confirmed that the photophysiological characteristics of free-living phytoplankton from multiple depths, consumed by zooplankton, and egested within FPs are retained for at least 24 h after grazing is initiated. These results have implications for high-resolution modeling of individual or group-specific zooplankton grazing dynamics that are critical for accurately linking zooplankton grazing in the surface ocean with the mesopelagic and deep ocean food webs and carbon export.
Climate-driven warming is projected to intensify wildfires, increasing their frequency and severity globally. Wildfires are an increasingly significant source of atmospheric deposition, delivering nutrients, organic matter, and trace metals to coastal and open ocean waters. These inputs have the potential to fertilize or inhibit microbial growth, yet their ecological impacts remain poorly understood. This study examines how ash leachate, derived from the 2017 Thomas Fire in California and lab-produced ash from Oregon vegetation, affects coastal plankton communities. Shipboard experiments off the California coast examined how pre-existing plankton biomass concentrations mediate responses to ash leachates. We found that ash leachate contained dissolved organic matter (DOM) that significantly increased bacterioplankton specific growth rates and DOM remineralization rates but had a negligible effect on bacterioplankton growth efficiency, suggesting low DOM bioavailability. Furthermore, ash-derived DOM had a higher potential to accumulate in high biomass water, where pre-existing DOM substrates may better support bacterial metabolism. Ash leachate had a neutral to negative effect on phytoplankton division rates and decreased microzooplankton grazing rates, particularly in low biomass water, leading to increased phytoplankton accumulation. Nanoeukaryotes accumulated in low biomass water, whereas picoeukaryotes and Synechococcus accumulated in high biomass water. Our findings suggest that the influence of ash deposition on DOM cycling, phytoplankton accumulation, and broader marine food web dynamics depends on pre-existing biomass levels. Understanding these interactions is critical for predicting the biogeochemical consequences of increasing wildfire activity on marine ecosystems.
Phytoplankton photoacclimation is a well‐documented response to changes in light and nutrient availability, with the Chlorophyll a to phytoplankton Carbon ratio ( = Chl: ) increasing at low light and decreasing under high light to optimize growth rate. Accurate estimation of phytoplankton growth rates and Net Primary Production (NPP) from space requires knowledge of , but cloud cover creates gaps. Current NPP models fill in the gaps by interpolating Chl (and other inputs) from clear‐sky pixels, ignoring the possibility of photoacclimation underneath clouds. Using data from 9,000 matchups between BioGeoChemical‐Argo floats and cloud cover from the Moderate Resolution Imaging Spectroradiometer, we compared the response of to irradiances under cloudy and clear skies. We found that phytoplankton photoacclimate similarly regardless of sky conditions at the global scale. This study highlights an incorrect assumption in current NPP estimates and suggests ways to improve global assessments of both chlorophyll and NPP.
Ocean color satellites have provided a synoptic view of global phytoplankton for over 25 years through near surface measurements of the concentration of chlorophyll a. While remote sensing of ocean color has revolutionized our understanding of phytoplankton and their role in the oceanic and freshwater ecosystems, it is important to consider both total phytoplankton biomass and changes in phytoplankton community composition in order to fully understand the dynamics of the aquatic ecosystems. With the upcoming launch of NASA's Plankton, Aerosol, Clouds, ocean Ecosystem (PACE) mission, we will be entering into a new era of global hyperspectral data, and with it, increased capabilities to monitor phytoplankton diversity from space. In this paper, we analyze the needs of the user community, review existing approaches for detecting phytoplankton community composition in situ and from space, and highlight the benefits that the PACE mission will bring. Using this three-pronged approach, we highlight the challenges and gaps to be addressed by the community going forward, while offering a vision of what global phytoplankton community composition will look like through the “eyes” of PACE.
Bio-optical properties of marine phytoplankton retrieved through satellite remote sensing are used to estimate ocean productivity and carbon cycling. Daily activity of phytoplankton is attuned to the predictable light fluctuations of the diel cycle. Field and laboratory studies have documented diel changes in phytoplankton growth, division, and respiration, carbon and pigment content, cell size, photosynthetic efficiency and rate, and DNA replication and transcription. Many of these physiological changes can alter cellular optical properties and contribute to diel variations in bulk absorption and scattering properties. Consequently, understanding phytoplankton contributions to diel optical cycles is essential for improving algorithms that convert remote sensing data to biological rates and stocks. Here, we describe time-resolved cellular, photophysiological, and bio-optical properties for three cultured phytoplankton ranging in cell size from (+/-) 1 to 6 mu m: Ostreococcus lucimarinus, Synechococcus (WH8102), and Thalassiosira pseudonana, all of which can significantly contribute to phytoplankton abundance and/or biomass in the open and coastal ocean. This work is the first to characterize complete diel cycles in absorption and attenuation for O. lucimarinus and backscattering for all these species. Results show that the percent increase from the minimum to maximum values over the diel cycle ranged between (+/-) 24-121%, (+/-) 31-235%, and (+/-) 25-198% for particulate absorption, attenuation, and backscattering, respectively. Diel changes in bio-optical properties also differed in both timing and magnitude across phytoplankton species, demonstrating the importance of contextualizing remote sensing observations with phytoplankton community composition.
The composition of the marine phytoplankton community has been shown to impact many biogeochemical processes and marine ecosystem services. A variety of methods exist to characterize phytoplankton community composition (PCC), with varying degrees of taxonomic resolution. Accordingly, the resulting PCC determinations are dependent on the method used. Here, we use surface ocean samples collected in the North Atlantic and North Pacific Oceans to compare high performance liquid chromatography (HPLC) pigment-based PCC to four other methods: quantitative cell imaging, flow cytometry, and 16S and 18S rRNA amplicon sequencing. These methods allow characterization of both prokaryotic and eukaryotic PCC across a wide range of size classes. PCC estimates of many taxa resolved at the class level (e.g., diatoms) show strong positive correlations across methods, while other groups (e.g., dinoflagellates) are not well captured by one or more methods. Since variations in phytoplankton pigment concentrations are related to changes in optical properties, this combined dataset expands the potential scope of ocean color remote sensing by associating PCC at the genus- and species-level with group- or class-level PCC from pigments. Quantifying the strengths and limitations of pigment-based PCC methods compared to PCC assessments from amplicon sequencing, imaging, and cytometry methods is the first step toward the robust validation of remote sensing approaches to quantify PCC from space.
This manuscript presents an overview of NASA’s EXport Processes in the Ocean from Remote Sensing 2021 Field Campaign in the North Atlantic (EXPORTS NA) and provides quantitative and dynamical descriptions of the physical processes modulating water transformations during the study. A major programmatic goal was to conduct the sampling in a Lagrangian mode so that ocean ecological and biogeochemical changes can be observed independent from physical advective processes. To accomplish this goal, EXPORTS NA conducted a multi-ship, multi-asset field sampling program within a retentive, anticyclonic mode water eddy. Beneath depths of ~100 m, Lagrangian sampling assets remained within the eddy core waters (ECWs) throughout the experiment, demonstrating that the ECWs within the mode water eddy were retentive. However, strong westerly winds from four storm events deepened the mixed layer (ML) of the surface core waters (SCWs) above the eddy’s mode water core by 25-40 m and exchanged some of the SCWs with surface waters outside of the eddy via Ekman transport. Estimates of flushing times ranged from 5-8 days, with surface exchange fractions ranging from 20-75% and were consistent with particle tracking advected by combined geostrophic and Ekman velocities. The relative contributions of horizontal and vertical advection on changes in SCW tracers depend on the horizontal and vertical gradients of that tracer. For example, in the surface waters, horizontal advection played a large role in salinity fluxes, yet vertical entrainment played a larger role in the fluxes of nutrients into the ML. Each storm injected nutrients and low oxygen waters into the ML, after which the surface ocean ecosystem responded by reducing nutrient concentrations and increasing %O2 saturation levels. Overall, SCW values of chlorophyll and POC were the largest at the onset of the field program and decreased throughout the campaign. The analysis presented provides a physical oceanographic context for the many measurements made during the EXPORTS NA field campaign. Illustrated are the many challenges of conducting a production-flux experiment even in a Lagrangian frame and the inherent uncertainties of interpreting biological carbon pump observations that were collected in a Eulerian frame of reference.
The gravitational settling of organic particles from the surface to the deep ocean is an important export pathway and one of the largest components of the marine biological carbon pump (BCP). The strength and efficiency of the gravitational pump is often measured using metrics reliant on reference depths and empirical formulations that parameterize the relationship between depth and flux or concentration. Here, BGC-Argo profiles were used to identify the isolume where POC concentration starts to decline, revealing attenuation trends below this isolume that are remarkably consistent across the global ocean. We developed a semi-mechanistic approach that uses observations from the first optical depth to predict POC concentration from the surface ocean to the base of the mesopelagic (1000 m), allowing assessments of spatial and temporal variability in BCP efficiencies. We find that rates of POC attenuation are high in areas of high biomass and low in areas of low biomass, supporting the view that bloom events sometimes result in a relatively weak deep biological pump characterized by low transfer efficiency to the base of the mesopelagic. Our isolume-based attenuation model was applied to satellite data to yield the first remote sensing-based estimate of integrated global POC stock of 3.02 Pg C for the upper 1000 m, with 1.27 Pg C of this global carbon stock located above the reference isolume where POC begins to attenuate.
Microbial respiration is a critical component of the marine carbon cycle, determining the proportion of fixed carbon that is subject to remineralization as opposed to being available for export to the ocean depths. Despite its importance, methodological constraints have led to an inadequate understanding of this process, especially in low-activity oligotrophic and mesopelagic regions. Here, we quantify respiration rates as low as 0.2 µmol O2 L-1 d-1 in contrasting ocean productivity provinces using oxygen optode sensors to identify size-fractionated respiration trends. In the low productivity region of the North Pacific Ocean at Station Papa, surface whole water microbial respiration was relatively stable at 1.2 µmol O2 L-1 d-1. Below the surface, there was a decoupling between respiration and bacterial production that coincided with increased phytodetritus and small phytoplankton. Size-fractionated analysis revealed that cells <5 µm were responsible for the majority of the respiration in the Pacific, both at the surface and below the mixed layer. At the North Atlantic Porcupine Abyssal Plain, surface whole water microbial respiration was higher (1.7 µmol O2 L-1 d-1) than in the Pacific and decreased by 3-fold below the euphotic zone. The Atlantic size-fraction contributions to total respiration shifted on the order of days during the evolution of a phytoplankton bloom with regular storm disturbances. The high-resolution optode method used in the Atlantic captured these significant shifts and is consistent with coinciding stain-based respiration methods and historical site estimates. This study highlights the dynamic nature of respiration across vertical, temporal, and size-fractionated factors, emphasizing the need for sensitive, high-throughput techniques to better understand ocean ecosystem metabolism.
Marine particle forensics frequently uncover information on composition, age, size, and ecological history. Zooplankton fecal pellets are also studied for process-related data, such as grazing rates and carbon sequestration potential. Here, flow cytometric analyses of fecal pellet contents revealed intact phytoplankton with photophysiological characteristics mirroring those of free-living cells. Mapping the cytometrically derived properties of cells inside fecal pellets onto vertical profiles from free-living cells revealed the potential to estimate depth specific grazing by individual zooplankton. An experiment conducted at sea confirmed that the photophysiological characteristics of free-living phytoplankton from multiple depths, consumed by zooplankton, and excreted within fecal pellets are retained for at least 24 hours after grazing is initiated. These results have implications for high resolution modeling of individual or group specific zooplankton grazing dynamics that are critical for accurately linking zooplankton grazing in the surface ocean with the mesopelagic and deep ocean food webs and carbon export. ### Competing Interest Statement The authors have declared no competing interest.
Viral lysis of phytoplankton is one of the most common forms of death on Earth. Building on an assay used extensively to assess rates of phytoplankton loss to predation by grazers, lysis rates are increasingly quantified through dilution-based techniques. In this approach, dilution of viruses and hosts are expected to reduce infection rates and thus increase host net growth rates (i.e., accumulation rates). The difference between diluted and undiluted host growth rates is interpreted as a measurable proxy for the rate of viral lytic death. These assays are usually conducted in volumes ≥ 1 L. To increase throughput, we implemented a miniaturized, high-throughput, high-replication, flow cytometric microplate dilution assay to measure viral lysis in environmental samples sourced from a suburban pond and the North Atlantic Ocean. The most notable outcome we observed was a decline in phytoplankton densities that was exacerbated by dilution, instead of the increased growth rates expected from lowered virus-phytoplankton encounters. We sought to explain this counterintuitive outcome using theoretical, environmental, and experimental analyses. Our study shows that, while die-offs could be partly explained by a ‘plate effect’ due to small incubation volumes and cells adhering to walls, the declines in phytoplankton densities are not volume-dependent. Rather, they are driven by many density- and physiology-dependent effects of dilution on predation pressure, nutrient limitation, and growth, all of which violate the original assumptions of dilution assays. As these effects are volume-independent, these processes likely occur in all dilution assays that our analyses show to be remarkably sensitive to dilution-altered phytoplankton growth and insensitive to actual predation pressure. Incorporating altered growth as well as predation, we present a logical framework that categorizes locations by the relative dominance of these mechanisms, with general applicability to dilution-based assays.
Food webs trace the flow of organic matter and energy among producers and consumers; for pelagic marine food webs, network complexity directly influences the amount and form of carbon exported to the deep ocean via the biological pump. Here we present a synoptic view of mixed layer food web dynamics observed during the late summer 2018 EXport Processes in the Ocean from Remote Sensing (EXPORTS) field campaign in the subarctic Northeast Pacific at the long-running time-series site, Ocean Station Papa. Carbon biomass reservoirs of phytoplankton, microzooplankton, and bacterioplankton, were approximately equal while mesozooplankton biomass was 70% lower. Live organisms composed ∼40% of the total particulate organic carbon within the mixed layer: the remainder was attributed to detritus. Rates of carbon transfer among reservoirs indicated production and assimilation rates were well balanced by losses, leaving little organic carbon available for export. The slight positive net community production rate generated organic carbon that was exported from the system in the form of food web byproducts, such as large fecal pellets generated by mesozooplankton. This characteristically regenerative food web had relatively slow turnover times with small-magnitude transfers of carbon relative to standing stocks that occurred amidst a high background concentration of detrital particles and dissolved organic matter. The concurrent estimation of food web components and rates revealed that separated processes dominated the transfer of carbon within the food web compared to those that contributed to export. Plain Language Summary The biological carbon pump drives a downward flux of organic matter from the sunlit surface ocean to the vast ocean interior. Ecological interactions in the surface ocean directly affect the amount and type of carbon that is exported to the deep ocean. In this study, we present a synthesis of the late summer mixed layer food web in the Northeast Pacific that was extensively characterized during the 2018 EXport Processes in the Ocean from Remote Sensing (EXPORTS) field campaign. We found the majority of carbon was recycled within the mixed layer by microbes through multiple transfers between producers and consumers. Larger organisms, mesozooplankton and salps, only consumed a small amount of carbon but through the formation of sinking fecal pellets were the main mechanism of transporting carbon out of the system. The study highlights the need to concurrently study microbial and large organism dynamics to develop a predictive understanding of the fate of organic carbon in the oceans. Key Points The microbial loop dominated carbon flow in the late summer mixed layer food web of the North Pacific, most net production was respired leaving little carbon available for export. Active production and consumption of organic carbon occurred amid a high background of detrital particulate organic carbon (58% of total) with slow turnover time, 66 d. Mesozooplankton which had relatively minor carbon consumption rates created the majority of export production due to efficient repackaging of consumed material.
Measurements of particulate organic carbon (POC) are critical for understanding the ocean carbon cycle, including biogenic particle formation and removal processes, and for constraining models of carbon cycling at local, regional, and global scales. Despite the importance and ubiquity of POC measurements, discrepancies in methods across platforms and users, necessary to accommodate a multitude of needs and logistical constraints, commonly result in disparate results. Considerations of filter type and pore size, sample volume, collection method, and contamination sources underscore the potential for dissimilar measurements of the same variable assessed using similar and different approaches. During the NASA EXport Processes in the Ocean from RemoTe Sensing (EXPORTS) 2018 field campaign in the North Pacific Ocean, multiple methodologies and sampling approaches for determining POC were applied, including surface inline flow-through systems and depth profiles using Niskin bottles, in situ pumps, and Marine Snow Catchers. A comparison of results from each approach and platform often resulted in significant differences. Supporting measurements, however, provided the means to normalize results across datasets. Using knowledge of contrasting protocols and synchronous or near-synchronous measurements of associated environmental variables, we were able to reconcile dataset differences to account for undersampling of some particle types and sizes, possible sample contamination and blank corrections. These efforts resulted in measurement agreement between initially contrasting datasets and insights on long-acknowledged but rarely resolved discrepancies among contrasting methods for assessing POC concentrations in the ocean.
Synechococcus are the most abundant cyanobacteria in high latitude regions and are responsible for an estimated 17% of annual marine net primary productivity. Despite their biogeochemical importance, Synechococcus populations have been unevenly sampled across the ocean, with most studies focused on low-latitude strains. In particular, the near absence of Synechococcus genomes from high-latitude, High Nutrient Low Chlorophyll (HNLC) regions leaves a gap in our knowledge of picocyanobacterial adaptations to iron limitation and their influence on carbon, nitrogen, and iron cycles. We examined Synechococcus populations from the subarctic North Pacific, a well-characterized HNLC region, with quantitative metagenomics. Assembly with short and long reads produced two near complete Synechococcus metagenome-assembled genomes (MAGs). Quantitative metagenome-derived abundances of these populations matched well with flow cytometry counts, and the Synechococcus MAGs were estimated to comprise >99% of the Synechococcus at Station P. Whereas the Station P Synechococcus MAGs contained multiple genes for adaptation to iron limitation, both genomes lacked genes for uptake and assimilation of nitrate and nitrite, suggesting a dependence on ammonium, urea, and other forms of recycled nitrogen leading to reduced iron requirements. A global analysis of Synechococcus nitrate reductase abundance in the TARA Oceans dataset found nitrate assimilation genes are also lower in other HNLC regions. We propose that nitrate and nitrite assimilation gene loss in Synechococcus may represent an adaptation to severe iron limitation in high-latitude regions where ammonium availability is higher. Our findings have implications for models that quantify the contribution of cyanobacteria to primary production and subsequent carbon export.
Microbial respiration plays a pivotal role in the marine carbon cycle, influencing the fraction of fixed carbon that undergoes remineralization versus export to depth. Despite its importance, methodological constraints have led to an inadequate understanding of this process, especially in low-activity oligotrophic and mesopelagic regions. Here, we quantify respiration rates as low as 0.2 µmol O2 L-1 d-1 in contrasting ocean productivity provinces using optical oxygen sensors to identify size-fractionated respiration trends. At the low productivity North Pacific Ocean Station Papa, surface microbial respiration was relatively stable at 1.2 µmol O2 L-1 d-1. Below the surface there was a decoupling of respiration and bacterial production potentially driven by phytodetritus remineralization. Size-fractionated rates showed cells <5 µm contributed the most to Pacific respiration. At the North Atlantic Porcupine Abyssal Plain, the optode measurement frequency was drastically increased. Surface microbial respiration was higher (1.7 µmol O2 L-1 d-1) and decreased by 3-fold below the euphotic zone. The Atlantic filtered fraction contributions to total respiration shifted with the phytoplankton bloom evolution. The high resolution optode method used in the Atlantic is consistent with coinciding in vivo 2-para-(iodophenyl)-3(nitrophenyl)-5(phenyl) tetrazolium chloride respiratory stain measurements and historical site estimates. We estimate that 58% of gross primary production was respired at the Pacific site and 34% at the Atlantic site, demonstrating that the Atlantic had a higher carbon export potential. This study highlights the dynamic nature of respiration across vertical, temporal, and size-fractionated factors, emphasizing the need for sensitive, high-throughput techniques to better understand ocean ecosystem metabolism.
Phytoplankton play a major role on Earth, impacting the global distribution and cycles of carbon, oxygen, nitrogen, sulfur, and other elements, and structuring marine food webs. One fundamental trait of phytoplankton with direct biogeochemical implications is their size, as it governs metabolic and sinking rates as well as prey–predator interactions. Phytoplankton size spans approximately 3.5 orders of magnitude (when expressed as an equivalent spherical diameter), and thus measuring the full range in size distribution of phytoplankton is challenging and rarely attempted. Here, we constructed phytoplankton size spectra by merging state‐of‐the‐art cytometry and imaging cytometry measurements that were collected in the western North Atlantic Ocean, along a latitudinal gradient (36°N to 55°N) and during different phases of the annual cycle of phytoplankton. The derived spectra show a seasonal pattern that parallels changes in phytoplankton biomass, and do not always follow a commonly assumed power‐law model. Shifts in size spectra were more pronounced in the sub‐Arctic and temperate subregions, compared to the subtropical region of the study area. We evaluated the relationships between different size groups and environmental parameters to derive ecologically meaningful size groups. Finally, to simulate Ocean Color remote‐sensing algorithms of phytoplankton size, we compared temporal variations in descriptors of the size spectra (median particle size, phytoplankton size distribution exponent) with optical size proxies derived from light absorption and attenuation; good agreement was observed in the northern sections of the study area where temporal changes in community size structure were more pronounced.
Dissolved organic carbon (DOC) produced by primary production in the sunlit ocean can be physically transported to the mesopelagic zone. The majority of DOC exported to this zone is remineralized by heterotrophic microbes over a range of timescales. Capturing a deep convective mixing event is rare, as is observing how microbes respond in situ to the exported DOC. Here, we report ship and Argo float observations from hydrostation North Atlantic Aerosols and Marine Ecosystems Study (NAAMES) 2 Station 4 (N2S4; 47.46°N, 38.72°W), a retentive anticyclonic eddy in the subtropical region of the western North Atlantic. Changes in biogeochemistry and bacterioplankton responses were tracked as the water column mixed to approximately 230 m and restratified over the subsequent 3 days. Over this period, rapid changes in bacterioplankton production (BP) and cell abundance were observed throughout the water column. BP increased by 91% in the euphotic zone (0–100 m) and 55% in the upper mesopelagic zone (100–200 m), corresponding to 33% and 103% increases in cell abundance, respectively. Within the upper mesopelagic, BP upon the occupation of N2S4 (20 ± 4.7 nmol C L–1 d–1) was significantly greater than the average upper mesopelagic BP rate (2.0 ± 1.6 nmol C L–1 h–1) at other stations that had been stratified for longer periods of time. BP continued to increase to 31 ± 3.0 nmol C L–1 d–1 over the 3-day occupation of N2S4. The rapid changes in BP in the upper mesopelagic did not coincide with rapid changes in community composition, but the taxa that increased in their relative contribution included those typically observed in the epipelagic zone. We interpret the subtle but significant community structure dynamics at N2S4 to reflect how injection of labile organic matter into the upper mesopelagic zone by physical mixing supports continued growth of euphotic zone-associated bacterioplankton lineages on a timescale of days.
In this article, we present Bio-GO-SHIP, a new ocean observing program that will incorporate sustained and consistent global biological ocean observations into the Global Ocean Ship-based Hydrographic Investigations Program (GO-SHIP). The goal of Bio-GO-SHIP is to produce systematic and consistent biological observations during global ocean repeat hydrographic surveys, with a particular focus on the planktonic ecosystem. Ocean plankton are an essential component of the earth climate system, form the base of the oceanic food web and thereby play an important role in influencing food security and contributing to the Blue Economy. Despite its importance, ocean biology is largely under-sampled in time and space compared to physical and chemical properties. This lack of information hampers our ability to understand the role of plankton in regulating biogeochemical processes and fueling higher trophic levels, now and in future ocean conditions. Traditionally, many of the methods used to quantify biological and ecosystem essential ocean variables (EOVs), measures that provide valuable information on the ecosystem, have been expensive and labor- and time-intensive, limiting their large-scale deployment. In the last two decades, new technologies have been developed and matured, making it possible to greatly expand our biological ocean observing capacity. These technologies, including cell imaging, bio-optical sensors and 'omic tools, can be combined to provide overlapping measurements of key biological and ecosystem EOVs. New developments in data management and open sharing can facilitate meaningful synthesis and integration with concurrent physical and chemical data. Here we outline how Bio-GO-SHIP leverages these technological advances to greatly expand our knowledge and understanding of the constituents and function of the global ocean plankton ecosystem.
Abstract The abundance and productivity of phytoplankton constrain energy transfer through marine food webs and the export of organic carbon to the deep ocean. Bio‐optical measurements correlate well with phytoplankton carbon (Cphyto), but the effect of taxonomic variability on this relationship is still uncertain. Here, we explore how changes in phytoplankton community composition influence the relationship between the particulate backscatter coefficient (bbp) and Cphyto and present a new approach to estimate phytoplankton biomass more accurately using bbp. We found that using a fixed scaling factor for the conversion of bbp to Cphyto could lead to the underestimation or overestimation of biomass, depending on the dominant taxonomic group in the phytoplankton community. In addition, we demonstrate how a simple ratio of absorption at two wavelengths can be used to provide a coarse approximation of phytoplankton community composition when scaling bbp to Cphyto, thereby improving the estimation of net primary production.
To identify the effect of microzooplankton grazing on phytoplankton abundance and size structure, we quantified phytoplankton growth and herbivorous grazing rates throughout the euphotic zone and across a light gradient on the North Pacific EXport Processes in the Ocean from RemoTe Sensing (EXPORTS) cruise near Ocean Station Papa. During 30 days of continuous, Lagrangian observation in August and September of 2018, depth integrated chlorophyll a (Chl a) concentrations were stable and averaged 20 +/- 2 mgm(-2). Bottle-incubation experiments revealed that phytoplankton growth was balanced by microzooplankton grazing even when phytoplankton growth rates varied from 0 to 0.4 d(-1) in response to light manipulation. Microzooplankton grazing caused a decline in phytoplankton abundance that was balanced by increased phytoplankton cell size resulting in consistent phytoplankton biomass over time. Microzooplankton grazed phytoplankton at an average rate of 0.11 +/- 0.17 d(-1) which lead to an intrinsic phytoplankton growth rate of -0.07 +/- 0.26 d(-1). Predicted stocks from grazing experiments aligned closely (within 16%) with in situ Chl a dynamics and phytoplankton abundance, suggesting that the dominant loss process of phytoplankton was grazing by microzooplankton rather than physical mixing or sinking of phytoplankton. Consequently, microzooplankton played a critical role in regulating primary producer biomass and in transferring particulate organic carbon through the food web where a fraction could then be exported as byproducts of food web processes.