Description: Nitrogen (N) is the major limiting nutrient for phytoplankton growth and productivity in large parts of the world's oceans. Differential preferences for specific N substrates may be important in controlling phytoplankton community composition. To date, there is limited information on how specific N substrates influence the composition of naturally occurring microbial communities. We investigated the effect of nitrate ( math formula), ammonium ( math formula), and urea on microbial and phytoplankton community composition (cell abundances and 16S rRNA gene profiling) and functioning (photosynthetic activity, carbon fixation rates) in the oligotrophic waters of the North Pacific Ocean. All N substrates tested significantly stimulated phytoplankton growth and productivity. Urea resulted in the greatest (〉300%) increases in chlorophyll a (〈0.06 μg L−1 and ∼0.19 μg L−1 in the control and urea addition, respectively) and productivity (〈0.4 μmol C L−1 d−1 and ∼1.4 μmol C L−1 d−1 in the control and urea addition, respectively) at two experimental stations, largely due to increased abundances of Prochlorococcus (Cyanobacteria). Two abundant clades of Prochlorococcus, High Light I and II, demonstrated similar responses to urea, suggesting this substrate is likely an important N source for natural Prochlorococcus populations. In contrast, the heterotrophic community composition changed most in response to math formula. Finally, the time and magnitude of response to N amendments varied with geographic location, likely due to differences in microbial community composition and their nutrient status. Our results provide support for the hypothesis that changes in N supply would likely favor specific populations of phytoplankton in different oceanic regions and thus, affect both biogeochemical cycles and ecological processes.

Description: To investigate the biogeochemistry of iron in the waters of the European continental margin, we determined the dissolved iron distribution and redox speciation in filtered (〈0.2 μm) open-ocean and shelf waters. Depth profiles were sampled over the shelf slope southeast of the Chapelle Bank area (47.61°N, 4.24°W to 46.00°N, 8.01°W) and a horizontal surface-water transect over the shelf and through the English Channel (la Manche) and the southern North Sea (46°N, 8°W to 52°N, 4°E). An abrupt trace-metal front was found near the shelf slope, indicated by a horizontal gradient of dissolved iron (DFe) and aluminium (DAl), which correlated with changing salinities (r2 = 0.572 and 0.528, respectively, n = 92). Labile Fe(II) concentrations varied from 〈12 pmol L-1 in North Atlantic surface waters to >200 pmol L-1 in the near bottom waters of the shelf break. Labile Fe(II) accounted for ∼5 of the dissolved iron species in surface shelf waters (mean 5.0 ± 2.7), whereas higher Fe(II) fractions (i.e., >8) were observed near the sea bottom on the shelf break and during a midday solar maximum in surface waters in the vicinity of the Scheldt river plume. Benthic processes (resuspension and diagenesis) constituted important sources of Fe(II) and DFe in this region, and photoreduction of Fe(III) species in shelf waters caused enhanced labile Fe(II) concentrations. These processes increased the lability of iron and its potential availability to marine organisms in the shelf ecosystem. © 2007, by the American Society of Limnology and Oceanography, Inc.

Description: A compilation of more than 30 studies shows that adult Antarctic krill (Euphausia superba) may frequent benthic habitats year-round, in shelf as well as oceanic waters and throughout their circumpolar range. Net and acoustic data from the Scotia Sea show that in summer 2-20 of the population reside at depths between 200 and 2000 m, and; that large aggregations can form above the seabed. Local differences in the vertical distribution of krill indicate that reduced feeding success in surface waters, either due to predator encounter or food shortage, might initiate such deep migrations and results in benthic feeding. Fatty acid and microscopic analyses of stomach content confirm two different foraging habitats for Antarctic krill: the upper ocean, where fresh phytoplankton is the main food source, and deeper water or the seabed, where detritus and copepods are consumed. Krill caught in upper waters retain signals of benthic feeding, suggesting frequent and dynamic exchange between surface and seabed. Krill contained up to 260 nmol iron per stomach when returning from seabed feeding. About 5 of this iron is labile, i.e., potentially available to phytoplankton. Due to their large biomass, frequent benthic feeding, and acidic digestion of particulate iron, krill might facilitate an input of new iron to Southern Ocean surface waters. Deep migrations and foraging at the seabed are significant parts of krill ecology, and the vertical fluxes involved in this behavior are important for the coupling of benthic and pelagic food webs and their elemental repositories. © 2011, by the American Society of Limnology and Oceanography, Inc.

Description: We investigated the effect of elevated partial pressure of CO2 (pCO2) on the photosynthesis and growth of four phylotypes (ITS2 types A1, A13, A2, and B1) from the genus Symbiodinium, a diverse dinoflagellate group that is important, both free-living and in symbiosis, for the viability of cnidarians and is thus a potentially important model dinoflagellate group. The response of Symbiodinium to an elevated pCO2 was phylotype-specific. Phylotypes A1 and B1 were largely unaffected by a doubling in pCO2; in contrast, the growth rate of A13 and the photosynthetic capacity of A2 both increased by ~ 60. In no case was there an effect of ocean acidification (OA) upon respiration (dark- or light-dependent) for any of the phylotypes examined. Our observations suggest that OA might preferentially select among free-living populations of Symbiodinium, with implications for future symbioses that rely on algal acquisition from the environment (i.e., horizontal transmission). Furthermore, the carbon environment within the host could differentially affect the physiology of different Symbiodinium phylotypes. The range of responses we observed also highlights that the choice of species is an important consideration in OA research and that further investigation across phylogenetic diversity, for both the direction of effect and the underlying mechanism(s) involved, is warranted. © 2011, by the American Society of Limnology and Oceanography, Inc.

Description: The high-latitude North Atlantic (HLNA) is characterized by a marked seasonal phytoplankton bloom, which removes the majority of surface macronutrients. However, incomplete nitrate depletion is frequently observed during summer in the region, potentially reflecting the seasonal development of an iron (Fe) limited phytoplankton community. In order to investigate the seasonal development and spatial extent of iron stress in the HLNA, nutrient addition experiments were performed during the spring (May) and late summer (July and August) of 2010. Grow-out experiments (48–120 h) confirmed the potential for iron limitation in the region. Short-term (24 h) incubations further enabled high spatial coverage and mapping of phytoplankton physiological responses to iron addition. The difference in the apparent maximal photochemical yield of photosystem II (PSII) (Fv : Fm) between nutrient (iron) amended and control treatments (D(Fv : Fm)) was used as a measure of the relative degree of iron stress. The combined observations indicated variability in the seasonal cycle of iron stress between different regions of the Irminger and Iceland Basins of the HLNA, related to the timing of the annual bloom cycle in contrasting biogeochemical provinces. Phytoplankton iron stress developed during the transition from the prebloom to peak bloom conditions in the HLNA and was more severe for larger cells. Subsequently, iron stress was reduced in regions where macronutrients were depleted following the bloom. Iron availability plays a significant role in the biogeochemistry of the HLNA, potentially lowering the efficiency of one of the strongest biological carbon pumps in the ocean.

Description: The duration and magnitude of the North Atlantic spring bloom impacts both higher trophic levels and oceanic carbon sequestration. Nutrient exhaustion offers a general explanation for bloom termination, but detail on which nutrients and their relative influence on phytoplankton productivity, community structure, and physiology is lacking. Here, we address this using nutrient addition bioassay experiments conducted across the midlatitude North Atlantic in June 2017 (late spring). In four out of six experiments, phytoplankton accumulated over 48–72 h following individual additions of either iron (Fe) or nitrogen (N). In the remaining two experiments, Fe and N were serially limiting, that is, their combined addition sequentially enhanced phytoplankton accumulation. Silicic acid (Si) added in combination with N + Fe led to further chlorophyll a (Chl a) enhancement at two sites. Conversely, addition of zinc, manganese, cobalt, vitamin B12, or phosphate in combination with N + Fe did not. At two sites, the simultaneous supply of all six nutrients, in combination with N + Fe, also led to no further Chl a enhancement, but did result in an additional 30–60% particulate carbon accumulation. This particulate carbon accumulation was not matched by a Redfield equivalent of particulate N, characteristic of high C:N organic exudates that enhance cell aggregation and sinking. Our results suggest that growth rates of larger phytoplankton were primarily limited by Fe and/or N, making the availability of these nutrients the main bottom‐up factors contributing to spring bloom termination. In addition, the simultaneous availability of other nutrients could modify bloom characteristics and carbon export efficiency.

Description: Bacterial productivity and biomass are thought to be limited by dissolved organic carbon (DOC) in much of the world’s oceans. However, the mixed layer of oligotrophic oceans is often depleted in dissolved inorganic nitrogen and phosphate, raising the possibility that macronutrients may also limit heterotrophic bacterial growth. We used nutrient bioassay experiments to determine whether inorganic nutrients (N, P, Fe) and/or DOC could limit bacterial productivity and biomass in the central North Atlantic during the spring of 2004 (Mar–Apr). We observed that both heterotrophic bacterial productivity and biomass were co-limited by N and P in the oligotrophic North Atlantic, and additions of labile DOC (glucose) provided no stimulation unless N and P were also added. Flow cytometry results indicated that only a small subset of large cells high in nucleic acid content were responsible for the increased productivity in the combined NP amendments. In contrast, nutrient additions elicited no net change on the dominant component of the bacterial population, composed of small cells with relatively low nucleic acid content. In the combined NP treatments the relative increase in bacterial production was greater than that measured when phytoplankton productivity was relieved of nitrogen limitation. These results suggest that N and P co-limitation in the bacterial community results in increased competition between the heterotrophic and autotrophic components of the surface communities in the Central North Atlantic Ocean, and potentially impacts the cycling of organic matter by the bacterioplankton.

Description: Nutrient addition bioassay experiments were performed in the low-nutrient, low-chlorophyll oligotrophic subtropical North Atlantic Ocean to investigate the influence of nitrogen (N), phosphorus (P), and/or iron (Fe) on phytoplankton physiology and the limitation of primary productivity or picophytoplankton biomass. Additions of N alone resulted in 1.5-2 fold increases in primary productivity and chlorophyll after 48 h, with larger (~threefold) increases observed for the addition of P in combination with N (NP). Measurements of cellular chlorophyll contents permitted evaluation of the physiological response of the photosynthetic apparatus to N and P additions in three picophytoplankton groups. In both Prochlorococcus and the picoeukaryotes, cellular chlorophyll increased by similar amounts in N and NP treatments relative to all other treatments, suggesting that pigment synthesis was N limited. In contrast, the increase of cellular chlorophyll was greater in NP than in N treatments in Synechococcus, suggestive of NP co-limitation. Relative increases in cellular nucleic acid were also only observed in Synechococcus for NP treatments, indicating co-limitation of net nucleic acid synthesis. A lack of response to relief of nutrient stress for the efficiency of photosystem II photochemistry, Fv :Fm, suggests that the low nutrient supply to this region resulted in a condition of balanced nutrient limited growth, rather than starvation. N thus appears to be the proximal (i.e. direct physiological) limiting nutrient in the oligotrophic sub-tropical North Atlantic. In addition, some major picophytoplankton groups, as well as overall autotrophic community biomass, appears to be co-limited by N and P.

Description: Nutrients limiting phytoplankton growth in the ocean are a critical control on ocean productivity and can underpin predicted responses to climate change. The extensive western subtropical North Pacific is assumed to be under strong nitrogen limitation, but this is not well supported by experimental evidence. Here, we report the results of 14 factorial nitrogen–phosphorus–iron addition experiments through the Philippine Sea, which demonstrate a gradient from nitrogen limitation in the north to nitrogen–iron co-limitation in the south. While nitrogen limited sites responded weakly to nutrient supply, co-limited sites bloomed with up to ~60-fold increases in chlorophyll a biomass that was dominated by initially undetectable diatoms. The transition in limiting nutrients and phytoplankton growth capacity was driven by a gradient in deep water nutrient supply, which was undetectable in surface concentration fields. We hypothesize that this large-scale phytoplankton response gradient is both climate sensitive and potentially important for regulating the distribution of predatory fish.

Description: Promoting effects of aluminum addition on chlorophyll biosynthesis and growth of two cultured iron‐limited marine diatoms Linbin Zhou CAS Key Laboratory of Tropical Marine Bio‐resources and Ecology, Guangdong Provincial Key Laboratory of Applied Marine Biology, South China Sea Institute of Oceanology Chinese Academy of Sciences Guangzhou China Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany University of Chinese Academy of Sciences Beijing China https://orcid.org/0000-0001-7230-4116 Fengjie Liu Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany Grantham Institute—Climate Change and the Environment, Department of Life Sciences Imperial College London London UK Eric P. Achterberg Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany Anja Engel Marine Biogeochemistry Division GEOMAR Helmholtz Centre for Ocean Research Kiel Germany https://orcid.org/0000-0002-1042-1955 Peter G.C. Campbell Institut National de la Recherche Scientifique Centre Eau Terre Environnement Quebec Canada https://orcid.org/0000-0001-7160-4571 Claude Fortin Institut National de la Recherche Scientifique Centre Eau Terre Environnement Quebec Canada https://orcid.org/0000-0002-2479-1869 Liangmin Huang CAS Key Laboratory of Tropical Marine Bio‐resources and Ecology, Guangdong Provincial Key Laboratory of Applied Marine Biology, South China Sea Institute of Oceanology Chinese Academy of Sciences Guangzhou China University of Chinese Academy of Sciences Beijing China Yehui Tan CAS Key Laboratory of Tropical Marine Bio‐resources and Ecology, Guangdong Provincial Key Laboratory of Applied Marine Biology, South China Sea Institute of Oceanology Chinese Academy of Sciences Guangzhou China University of Chinese Academy of Sciences Beijing China Abstract Aluminum (Al) may play a role in the ocean's capacity for absorbing atmospheric CO 2 via influencing carbon fixation, export, and sequestration. Aluminum fertilization, especially in iron (Fe)‐limited high‐nutrient, low‐chlorophyll ocean regions, has been proposed as a potential CO 2 removal strategy to mitigate global warming. However, how Al addition would influence the solubility and bioavailability of Fe as well as the physiology of Fe‐limited phytoplankton has not yet been examined. Here, we show that Al addition (20 and 100 nM) had little influence on the Fe solubility in surface seawater and decreased the Fe bio‐uptake by 11–22% in Fe‐limited diatom Thalassiosira weissflogii in Fe‐buffered media. On the other hand, the Al addition significantly increased the rate of chlorophyll biosynthesis by 45–60% for Fe‐limited T. weissflogii and 81–102% for Fe‐limited Thalassiosira pseudonana , as well as their cell size, cellular chlorophyll content, photosynthetic quantum efficiency ( F v / F m ) and growth rate. Under Fe‐sufficient conditions, the Al addition still led to an increased growth rate, though the beneficial effects of Al addition on chlorophyll biosynthesis were no longer apparent. These results suggest that Al may facilitate chlorophyll biosynthesis and benefit the photosynthetic efficiency and growth of Fe‐limited diatoms. We speculate that Al addition may enhance intracellular Fe use efficiency for chlorophyll biosynthesis by facilitating the superoxide‐mediated intracellular reduction of Fe(III) to Fe(II). Our study provides new evidence and support for the iron–aluminum hypothesis.