Climate change dominates the interannual variation of carbon export efficiency in each season in the Northern Gulf of Mexico during 1998-2011.

Carbon export efficiency is a key indicator of the capacity of biological pump, but the controlling mechanism of the efficiency remains unclear. Our findings revealed that interannual variations in seasonal carbon export efficiency are determined by direct factors including riverine nutrient fluxes, stratification, residence time. These direct factors are finally attributed to two indirect factors (human activities and climate change). We quantified the absolute contributions of direct and indirect factors to carbon export efficiency. The results showed that the carbon export efficiency in the northern Gulf of Mexico in spring (summer; autumn; winter) was driven by human activities, which accounted for an absolute contribution of 16.02% (7.20%; 4.00%; 8.49%, respectively) through riverine nutrient fluxes, and by climate change, which accounted for an absolute contribution of 33.51% (21.43%; 25.73%; 15.80%, respectively) through stratification and water residence time. Moreover, carbon export efficiency could be predicted by MEI of 8 months earlier.

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To explore the temporal and spatial variations in phytoplankton community in small estuaries, we collected surface water samples from Yongjiang River estuary during wet, normal, and dry seasons and determined the main driving factors of phytoplankton community. A total of 358 species belonging to nine phyla and 123 genera were identified in all seasons. During wet, normal, and dry seasons, species number was 276, 154 and 151, and the abundance was (170.45±225.43)×103, (51.92±30.28)×103 and (31.65±12.79)×103 cells·L-1, respectively. Diatoms dominated the phytoplankton community, and the main dominant species were Cyclotella meneghiniana, Skeletonema costatum, and Paralia sulcata. Shannon diversity and Pielou evenness indices decreased from inside mouth to outside mouth in wet season, but there was no obvious spatial difference in normal season or dry season. Results of non-metric multidimensional scaling analysis and analysis of similarities showed that phytoplankton community composition differed significantly among different regions (inside, at and outside mouth) and different seasons. In wet season, phytoplankton abundance was significantly positively correlated with temperature, dissolved inorganic nitrogen, and dissolved reactive phosphorus, but significantly negatively correlated with salinity. In normal season, phytoplankton abundance was significantly negatively correlated with temperature. In dry season, it was not significantly correlated with environmental factors. Results of redundancy analysis showed that temperature, salinity, ammonium and dissolved reactive phosphorus explained the variations in phytoplankton community by 19.5%, 11.9%, 9.4% and 8.2%, respectively. These results revealed high dominance of diatoms and the main driving factors (temperature, salinity and nutrients) of phytoplankton community in Yongjiang River estuary.

Elemental geochemical evidence for the river-derived sources of trace metals in surface sediments from Hangzhou Bay, East China Sea.

Coastal estuaries are often heavily subject to riverine influences by the inputs of sediment from terrestrial sources. Hangzhou Bay (HZB) is threatened by the riverine derived trace metals from two large rivers of Qiantang River (QTR) and Yangtze River (YZR). However, previous studies mainly focused on the incidental transport from the largest river in China (YZR) and failed to simultaneously evaluate the contributions of these two rivers, especially the directly flowing river of QTR, by their trace elemental geochemical composition and distribution. Herein, a comprehensive study identified the river-derived sources of multiple trace metals in surface sediments which transported from both of the rivers. The sampling stations were separated into three regions of YZR, HZB, and QTR based on their spatial distributions of sediment grain size and components. The significant variations for most of the trace metals concentrations, except for Cd, Th, and U, were found among three regions (χ2 ≥ 8.22, p ≤ 0.016). The highest concentrations in HZB were mainly resulted from the grain size effect (68.82% of the total variance), while the highest concentrations of Sr, Cd, and Ba in YZR and Zr and Hf in QTR were attributed to the anthropogenic source (11.90%) and mineral composition (6.21%) of river basins. After normalized the diversity of multiple trace metals concentrations and the influence of grain size by ratios of Igeo and EFLi, three regions were effectively distinguished. It was indicated that As, Cd, and Sb were enriched in the sediments of rivers by anthropogenic source (EFLi > 1.5 and/or Igeo > 1). The results evidenced that, after removing the influence of grain size, elemental geochemical composition of the surface sediments confidently identified the river-derived anthropogenic sources of the enriched trace metals from two major rivers, and largely from YZR.

Long-term response of interspecific competition among three typical bloom-forming species to changes in phosphorus and temperature.

Phosphorus and temperature play an important role in the succession of diatom-dinoflagellate blooms. However, there is little long-term research on interspecific competition based on phosphorus source and temperature. Here, interspecific competition among Skeletonema costatum, Prorocentrum donghaiense and Karenia mikimotoi was studied using trialgal laboratory co-cultures under different phosphorus and temperature conditions. These results suggest that S. costatum and P. donghaiense alternated as competing dominant species during the experimental period, which coincides with the different phosphorus conditions. However, K. mikimotoi growth was significantly inhibited throughout the experiment. We suggest that this may be due to different algal requirements for phosphorus, optimal growth temperatures, and possible allelopathic effects. This study provides a comprehensive mechanism of interspecific competition between diatom-dinoflagellate in response to phosphorus and temperature and elucidates the seasonal succession of diatom-dinoflagellate from late spring to early summer in the Changjiang River Estuary and the adjacent East China Sea.

Decadal changes of macrofauna community in a semi-enclosed Bay of Yueqing in East China Sea.

To reveal the long-term variation of macrofauna community in Yueqing Bay, an aquacultural bay famous for its shellfish culturing in the East China Sea, macrofauna samples were collected in three period from 2002 to 2003 and 2006-2007 to 2020-2021. The results show that macrofaunal community structure in this area has changed significantly (ANOSIM, p < 0.01) in nearly two decades with significant decreases in species number, biodiversity index and average biomass. Meanwhile, the taxa composition also changed significantly as the dominance of annelid increased while that of mollusks, echinoderms and vertebrates decreased. As a consequence of the variation of taxa composition and total biomass, macrofauna community showed a tendency of miniaturization as individuals with smaller body size and lower biomass dominated the community. According to the results of CCA analysis, temperature, salinity and dissolved oxygen content were the main environmental factors that restricted the species composition of macrofauna community. Further studies still needed to reveal the main reasons that cause the variation of macrofauna community. Overall, the results of this study suggest that the present status of Yueqing Bay benthic ecosystem is concerning from a macrobenthos perspective, as the biodiversity index and biomass of macrofauna decreased significantly. Effective measures should be taken in urgently to restrain the safety and function of coastal ecosystems.

Characterization and risk assessment of microplastics in laver from the Yueqing Bay.

Microplastics (MPs) pollution is regarded as a global challenge for ocean. As an important food source of human, macroalgae could suffer MP pollution and transmit MPs into human via food web. However, few studies have revealed the relationship of MP pollution between macroalgae and its habitat. In order to evaluate the trapping and accumulation of MPs in macroalgae and surface water, the present study investigated MP pollution in a typical aquaculture macroalgae species, laver (Porphyra haitanensis) in the Yueqing Bay. The results indicated MP abundance in laver (1.45 ± 0.26 items/g) was at a medium level while MP abundance in surface water (0.21 ± 0.15 item/m3) was at a relatively low level worldwide. Distribution trend and characteristics of MPs in laver and surface water showed highly similarity. Besides, heavy metal elements (Fe and Zr) were detected on the surface of MPs trapped by laver. Pollution load index (PLI) in surface water of the whole bay was low, indicating MP pollution was not serious in the Yueqing Bay. Due to the discharging of domestic sewage in recent years, fiber-shaped, textile MPs accounted for most in laver and surface water of the Yueqing Bay. These results indicated that MPs in surface water could be trapped by P. haitanensis, thus macroalgae cultivation might be a potential way to alleviate seawater MP pollution in the nearshore areas.

Sustainably developing global blue carbon for climate change mitigation and economic benefits through international cooperation.

Blue carbon is the carbon storage in vegetated coastal ecosystems such as mangroves, salt marshes, and seagrass. It is gaining global attention as its role in climate change mitigation and local welfare growth. However, a global assessment on the long-term spatiotemporal sustainable development status of blue carbon has not been conducted, and the relations among blue carbon ecosystems, driving forces for climate change mitigation, and socioeconomic interventions for development capacity on a global scale are still unclear. Here, we constructed a blue carbon development index (BCDI), comprising three subsystems: driving force, resource endowment, and development capacity, to assess the sustainable development level of 136 coastal countries' blue carbon over 24 consecutive years and explore the relationship among subsystems. We further propose a cooperation model to explore the feasibility of global blue carbon cooperation and quantify benefit allocation to specific countries. The results showed an upward trend in BCDI scores with variations in regional performance over the past two decades, and we found a positive correlation between development capacity and blue carbon resource endowment. Based on the scenario simulations of global cooperation, we found that coastal countries could improve the global average BCDI score, add 2.96 Mt of annual carbon sequestration, and generate $136.34 million in 2030 under Global Deep Cooperation scenario compared with the Business-As-Usual scenario.

Size-fractionated N2 fixation off the Changjiang Estuary during summer.

Recent evidence has shown active N2 fixation in coastal eutrophic waters, yet the rate and controlling factors remain poorly understood, particularly in large estuaries. The Changjiang Estuary (CE) and adjacent shelf are characterized by fresh, nitrogen-replete Changjiang Diluted Water (CDW) and saline, nitrogen-depletion intruded Kuroshio water (Taiwan Warm Current and nearshore Kuroshio Branch Current), where N2 fixation may be contributed by different groups (i.e., Trichodesmium and heterotrophic diazotrophs). Here, for the first time, we provide direct measurement of size-fractionated N2 fixation rates (NFRs) off the CE during summer 2014 using the 15N2 bubble tracer method. The results demonstrated considerable spatial variations (southern > northern; offshore > inshore) in surface and depth-integrated NFRs, averaging 0.83 nmol N L-1 d-1 and 24.3 µmol N m-2 d-1, respectively. The highest bulk NFR (99.9 µmol N m-2 d-1; mostly contributed by >10 µm fraction) occurred in the southeastern East China Sea, where suffered from strong intrusion of the Kuroshio water characterized by low N/P ratio (<10) and abundant Trichodesmium (up to 10.23 × 106 trichomes m-2). However, low NFR (mostly contributed by <10 µm fraction) was detected in the CE controlled by the CDW, where NOx concentration (up to 80 µmol L-1) and N/P ratio (>100) were high and Trichodesmium abundance was low. The >10 µm fraction accounted for 60% of depth-integrated bulk NFR over the CE and adjacent shelf. We speculated that the present NFR of >10 µm fraction was mostly supported by Trichodesmium. Spearman rank correlation indicated that the NFR was significantly positively correlated with Trichodesmium abundance, salinity, temperature and Secchi depth, but was negatively with turbidity, N/P ratio, NOx, and chlorophyll a concentration. Our study suggests that distribution and size structure of N2 fixation off the CE are largely regulated by water mass (intruded Kuroshio water and CDW) movement and associated diazotrophs (particularly Trichodesmium) and nutrient conditions.

Siting MPAs for multiple protecting purposes by co-consideration of ecological importance and anthropogenic impacts.

The global marine ecosystem has been significantly altered by the combined effects of multiple anthropogenic impacts. Systematic planning of marine protected areas (MPAs) is of paramount importance in alleviating conflicts between humans and the sea. Existing approaches, however, merely integrate both ecological and anthropogenic factors for multiple conservation purposes. By combining the three main anthropogenic impact factors with two main ecological importance factors, this study used a GIS-based AHP-OWA method to identify different levels of priority protection for MPAs in Zhejiang, China. Our results proved that: 1) the multi-objective MPA siting issues can be addressed by the GIS-based AHP-OWA method through scenario simulation; 2) the best locations for MPAs are in the northeast, central, and southern marine areas of Zhejiang; 3) considering the trade-off degree, spatial conservation efficiency, and spatial heterogeneity, an optimized MPA siting scheme can be developed for decision-makers. The proposed MPA siting method and case study may provide an effective technical reference for solving regional marine spatial planning (MSP) issues in the future.

A comprehensive evaluation of microplastic pollution in the Xiangshan Bay of China with special reference to seasonal variation.

Marine microplastic (MP) pollution has drawn global attention due to its potential risk to ecosystem. In the present study, we investigated MP pollution in surface water and sediment of a semi-closed bay: the Xiangshan Bay in the East China Sea in spring and summer. The results showed that MP abundance in surface water increased significantly in summer than spring (0.233 and 0.036 item/m3, respectively), while MP abundance in sediment was relatively steady. Meanwhile, the smaller size MPs (diameter < 1000 µm) and land-input fragment-shaped and film-shaped PP and PE increased in surface water in summer compared to spring. Surface microstructure of MPs showed that there were more cracks on MPs in summer comparing to spring. Based on diversity index, MP pollution in the Xiangshan Bay was at a low level and the composition was relatively uncomplicated. The source tracing analysis indicated main contributor of MPs were different in two seasons: textile industry was the dominate source of MPs in spring while fishery production were the dominate source in summer. Our results indicate that the pollution source of MPs could be various in different seasons due to the different climate and human activities, and provide a reference in the prevention and control of MP pollution in semi-closed bay ecosystems.

Ecological risk assessment of trace metals in sediments and their effect on benthic organisms from the south coast of Zhejiang province, China.

To explore the ecological risks of trace metals in sediments and their relationship with benthic organisms, 12 trace metals were analyzed and the macrobenthos were identified in the sediments collected from the south coast of Zhejiang province which belongs to the East China Sea. Spatially, the concentrations of most trace metals were high in the estuary, except for Ba and Sr. There was no obvious enrichment for trace metals, except that the concentration of Cd slightly exceeded the coastal background. The ecological risks calculated by the concentrations of Cr, Cd, Cu, Zn, Pb, and Ni in sediments showed that the methods based on sediment quality guidelines could judge the ecological risk more intuitively than the methods based on background value (PN, PLI, RI). The significant correlations between ecological risks and benthos density and biomass revealed the negative impact of trace metals at high concentrations on macrobenthic survival in sediments.

Differential physiological response of marine and freshwater microalgae to polystyrene microplastics.

Effects of microplastics on microalgae have not been compared from different habitat. To answer this question, three marine microalgae species (Chlorella marined, Nannochloropsis oculate, and Phaeodactylum tricornutum) and two freshwater species (Chlorella vulgaris and Tetradesmus obliquus) were selected and exposed to the environment relevant concentrations of polystyrene microplastics. The results indicated that microplastics have a significant concentration effect on the growth of microalgae. The attachment of microalgae to microplastics surface and the aggregation of microalgae with each other were observed. Under exposure of microplastics, the photosynthesis of microalgae was inhibited while the antioxidant system was activated, indicating that microplastics had a negative impact on microalgae. At the end of exposure, the oxidative stress status caused by microplastics in marine microalgae were alleviated, but the antioxidant system of freshwater microalgae was still at high levels, indicating a stress response. In addition, integrated biomarker response (IBR) indicated that the effects of microplastics on freshwater microalgae were severer than marine microalgae, which might relate to their differences in removing reactive oxygen species (ROS) effectively and membrane structure. Our study provides a reliable data for understanding the complex effects of microplastics on microalgae, and especially for comparing the differential effects of microplastics among different microalgae.

Influence of biodeposition by suspended cultured oyster on the distributions of trace elements in multiple media in a semi-enclosed bay of China.

It remains unclear how the suspended non-fed bivalve mariculture will alter the coastal transfer and cleaning process of trace elements, the non-degradable contaminants, which have been reported to accumulate in sediment from bivalve mariculture areas. Herein, we set up a field in situ comparative test in the suspended oyster (Crassostrea plicatula) farming area (OF) and reference area (RA) of Xiangshan Bay to verify our hypothesis that the biodepositon of suspended oysters would consolidate trace elements from the water column and transport them to the sediment. Distribution of trace elements in multiple media of biodeposits (BDs), settling particles (SPs), sediments (SEs), and seawater demonstrate that the accelerated deposition of BDs which enriched trace elements from the water column by oysters filtering suspended particles led to trace elements accumulation in SEs from OF. Additionally, As, Cd, Co, Cr, Cu, Ni, V, and Zn were strongly regulated by this process with significant (p < 0.05) higher concentrations in SEs from OF (10.96, 0.20, 13.98, 82.40, 38.47, 38.22, 108.57, and 111.20 µg/g, repectively) than those from RA (9.43, 0.13, 11.76, 63.30, 30.34, 29.55, 86.59, and 100.24 µg/g, repectively), but the extent was different for Mn, Mo, Pb, and W with concentrations in SEs from OF (737.37, 0.81, 30.98, and 3.96 µg/g, repectively) and RA (765.25, 0.69, 31.27, and 3.34 µg/g, repectively), especially for Rb and Sr with concentrations in SEs from OF (131.13 and 96.24 µg/g, repectively) and RA (142.21 and 161.10 µg/g, repectively), due to their geochemical and geophysical properties. Moreover, the harvest of hyper-accumulated oysters as a sink for removing trace elements from water column cannot hide the impact of this process.

Exploring Variability of Trichodesmium Photophysiology Using Multi-Excitation Wavelength Fast Repetition Rate Fluorometry.

Fast repetition rate fluorometry (FRRf) allows for rapid non-destructive assessment of phytoplankton photophysiology in situ yet has rarely been applied to Trichodesmium. This gap reflects long-standing concerns that Trichodesmium (and other cyanobacteria) contain pigments that are less effective at absorbing blue light which is often used as the sole excitation source in FRR fluorometers-potentially leading to underestimation of key fluorescence parameters. In this study, we use a multi-excitation FRR fluorometer (equipped with blue, green, and orange LEDs) to investigate photophysiological variability in Trichodesmium assemblages from two sites. Using a multi-LED measurement protocol (447+519+634 nm combined), we assessed maximum photochemical efficiency (F v /F m ), functional absorption cross section of PSII (σ PSII ), and electron transport rates (ETRs) for Trichodesmium assemblages in both the Northwest Pacific (NWP) and North Indian Ocean in the vicinity of Sri Lanka (NIO-SL). Evaluating fluorometer performance, we showed that use of a multi-LED measuring protocol yields a significant increase of F v /F m for Trichodesmium compared to blue-only excitation. We found distinct photophysiological differences for Trichodesmium at both locations with higher average F v /F m as well as lower σ PSII and non-photochemical quenching (NPQ NSV ) observed in the NWP compared to the NIO-SL (Kruskal-Wallis t-test df = 1, p < 0.05). Fluorescence light response curves (FLCs) further revealed differences in ETR response with a lower initial slope (α ETR ) and higher maximum electron turnover rate ( E T R P S I I m a x ) observed for Trichodesmium in the NWP compared to the NIO-SL, translating to a higher averaged light saturation E K (= E T R P S I I m a x /α ETR ) for cells at this location. Spatial variations in physiological parameters were both observed between and within regions, likely linked to nutrient supply and physiological stress. Finally, we applied an algorithm to estimate primary productivity of Trichodesmium using FRRf-derived fluorescence parameters, yielding an estimated carbon-fixation rate ranging from 7.8 to 21.1 mgC mg Chl-a-1 h-1 across this dataset. Overall, our findings demonstrate that capacity of multi-excitation FRRf to advance the application of Chl-a fluorescence techniques in phytoplankton assemblages dominated by cyanobacteria and reveals novel insight into environmental regulation of photoacclimation in natural Trichodesmium populations.

3-Indoleacrylic acid from canola straw as a promising antialgal agent - Inhibition effect and mechanism on bloom-forming Prorocentrum donghaiense.

Harmful algal blooms (HABs) have induced severe damage worldwide. A novel high-efficient antialgal natural chemical, 3-indoleacrylic acid (3-IDC) with a 5-day half-maximal inhibitory concentration (IC50, 5d), was discovered from canola straw, and its algal inhibition mechanism was investigated. Adverse effects were observed on the growth of P. donghaiense with 3-IDC addition, following an increase in reactive oxygen species (ROS) production. 3-IDC also hindered the photosynthetic mechanism of P. donghaiense cells. Transcriptional results showed 3-IDC inhibiting the functions of all the nutrient assimilating genes, down-regulated ribulose-1,5-bisphosphate carboxylase/oxygenase II, and cytochrome f genes. The expression of heat shock protein (HSP) 70 and 90 and rhodopsin genes were also suppressed. The binding affinity of investigated receptors was observed. The conformational changes induced by the spatial microstructural alteration through 3-IDC may further contribute to the perturbation of those enzyme catalytic activities. The present results provide new insights on controlling HABs using 3-IDC.

The contribution of ocean-based solutions to carbon reduction in China.

Climate change has become a great challenge for humanity. However, the current climate change mitigation measures, primarily concentrate on land, more or less neglecting the vital role of the ocean-based solutions. While the ocean is a crucial regulator of the global climate, ocean-based solutions could also play an essential role in climate change mitigation and policymaking. This paper developed an Ocean-based Solutions Carbon Reduction Assessment Model (OSCRAM) that addresses coastal ecosystems, ocean energy, marine transportation, fishery, and seabed to estimate the oceanic contribution to climate change mitigation. It has been applied to evaluate the capacity of carbon emission reduction through oceans in China. We found that the total contribution for carbon emission reduction was about 6.86 Tg CO2 per year, and it may reach 139.39 Tg in 2030 under the target scenario. The results indicated that the ocean has huge potential to reduce carbon emissions. The development of marine energy and low-carbon marine shipping may have more potential for emission reduction in China, and the government should also protect and restore coastal wetlands for their enormous carbon storage. It can also provide a reference for the globe and other nations in achieving emission reduction goals.

Oxidative stress induced by nanoplastics in the liver of juvenile large yellow croaker Larimichthys crocea.

There are many toxicological studies on microplastics, but little is known about the effect of nanoplastics (NPs). Here, we evaluated the oxidative stress responses induced by NPs (10, 104 and 106 particles/l) in juvenile Larimichthys crocea during 14-d NPs exposure followed by a 7-d recovery. After exposure, the activities of antioxidant enzymes (SOD, CAT, GPx) and MDA levels increased in the liver of fish at the highest NPs concentration. SOD and CAT activities remained elevated above the baseline after recovery under high-concentration NPs but returned to the baseline in two other NP treatments. Although lipid peroxidation in liver was reversible, juvenile fish in NPs treatments exhibited a lower survival rate than the control during both exposure and recovery. Furthermore, IBR value and PCA analysis showed the potential adverse effects of NPs. Considering that NPs can reduce the survival of fish juveniles, impacts of NPs on fishery productivity should be considered.

Toxicity mechanisms of polystyrene microplastics in marine mussels revealed by high-coverage quantitative metabolomics using chemical isotope labeling liquid chromatography mass spectrometry.

Marine microplastic has become an important environmental issue of global concern due to its wide distribution and harmful impacts. However, there is still insufficient information on the toxicity mechanism of microplastics to marine organisms. In this study, we developed and applied a high-coverage quantitative metabolomics technique to investigate the toxicity mechanisms of the polystyrene microspheres (micro-PS) on marine mussels (Mytilus coruscus). A total of 3599 metabolites were quantified, including 163 positively identified metabolites, 318 high-confident putatively identified metabolites, and 2602 mass-matched metabolites from the hemolymph of mussels. Metabolomics analysis indicated that micro-PS disrupted the amino acid metabolism, particularly phenylalanine metabolism, which may lead to oxidative stress and neurotoxicity. Micro-PS at environmentally relevant concentrations induced oxidative stress and immunotoxicity in mussels. After 7 days of recovery, along with the significant clearance of micro-PS by mussels, both metabolite levels and biochemical indicators generally returned to the same level as the control group. Overall, the results showed that microplastics at environmentally-relevant concentrations can cause toxic effects on mussels but these influences are reversible. We envisage the usages of high-coverage metabolomics for investigating the toxicity of various types of microplastics under many different conditions, including those relevant to the marine environment.

Seaweed farms provide refugia from ocean acidification.

Seaweed farming has been proposed as a strategy for adaptation to ocean acidification, but evidence is largely lacking. Changes of pH and carbon system parameters in surface waters of three seaweed farms along a latitudinal range in China were compared, on the weeks preceding harvesting, with those of the surrounding seawaters. Results confirmed that seaweed farming is efficient in buffering acidification, with Saccharina japonica showing the highest capacity of 0.10 pH increase within the aquaculture area, followed by Gracilariopsis lemaneiformis (ΔpH = 0.04) and Porphyra haitanensis (ΔpH = 0.03). The ranges of pH variability within seaweed farms spanned 0.14-0.30 unit during the monitoring, showing intense fluctuations which may also help marine organisms adapt to enhanced pH temporal variations in the future ocean. Deficit in pCO2 in waters in seaweed farms relative to control waters averaged 58.7 ± 15.9 µatm, ranging from 27.3 to 113.9 µatm across farms. However, ΔpH did not significantly differ between day and night. Dissolved oxygen and Ωarag were also elevated in surface waters at all seaweed farms, which are benefit for the survival of calcifying organisms. Seaweed farming, which unlike natural seaweed forests, is scalable and is not dependent on suitable substrate or light availability, could serve as a low-cost adaptation strategy to ocean acidification and deoxygenation and provide important refugia from ocean acidification.