Nutrient supply regulates the activity of phytoplankton, but the global biogeography of nutrient limitation and co-limitation is poorly understood. Prochlorococcus adapt to local environments by gene gains and losses, and we used genomic changes as an indicator of adaptation to nutrient stress. We collected metagenomes from all major ocean regions as part of the Global Ocean Ship-based Hydrographic Investigations Program (Bio-GO-SHIP) and quantified shifts in genes involved in nitrogen, phosphorus, and iron assimilation. We found regional transitions in stress type and severity as well as widespread co-stress. Prochlorococcus stress genes, bottle experiments, and Earth system model predictions were correlated. We propose that the biogeography of multinutrient stress is stoichiometrically linked by controls on nitrogen fixation. Our omics-based description of phytoplankton resource use provides a nuanced and highly resolved description of nutrient stress in the global ocean.
Climate-driven depletion of ocean oxygen strongly impacts the global cycles of carbon and nutrients as well as the survival of many animal species. One of the main uncertainties in predicting changes to marine oxygen levels is the regulation of the biological respiration demand associated with the biological pump. Derived from the Redfield ratio, the molar ratio of oxygen to organic carbon consumed during respiration (i.e., the respiration quotient, r−O2:C) is consistently assumed constant but rarely, if ever, measured. Using a prognostic Earth system model, we show that a 0.1 increase in the respiration quotient from 1.0 leads to a 2.3% decline in global oxygen, a large expansion of low-oxygen zones, additional water column denitrification of 38 Tg N/y, and the loss of fixed nitrogen and carbon production in the ocean. We then present direct chemical measurements of r−O2:C using a Pacific Ocean meridional transect crossing all major surface biome types. The observed r−O2:C has a positive correlation with temperature, and regional mean values differ significantly from Redfield proportions. Finally, an independent global inverse model analysis constrained with nutrients, oxygen, and carbon concentrations supports a positive temperature dependence of r−O2:C in exported organic matter. We provide evidence against the common assumption of a static biological link between the respiration of organic carbon and the consumption of oxygen. Furthermore, the model simulations suggest that a changing respiration quotient will impact multiple biogeochemical cycles and that future warming can lead to more intense deoxygenation than previously anticipated.
A past global synthesis of marine particulate organic matter (POM) suggested latitudinal variation in the ratio of surface carbon (C): nitrogen (N): phosphorus (P). However, this synthesis relied on compiled datasets that may have biased the observed pattern. To demonstrate latitudinal shifts in surface C:N:P, we combined hydrographic and POM observations from 28°N to 69°S in the eastern Pacific Ocean (GO-SHIP line P18). Both POM concentrations and ratios displayed distinct biome-associated changes. Surface POM concentrations were relatively low in the North Pacific subtropical gyre, increased through the Equatorial Pacific, were lowest in the South Pacific subtropical gyre, and increased through the Southern Ocean. Stoichiometric elemental ratios were systematically above Redfield proportions in warmer regions. However, C:P and N:P gradually decreased across the Southern Ocean despite an abundance of macro-nutrients. Here, a size-fraction analysis of POM linked increases in the proportion of large plankton to declining ratios. Subsurface N* values support the hypothesis that accumulated remineralization products of low C:P and N:P exported POM helps maintain the Redfield Ratio of deep nutrients. We finally evaluated stoichiometric models against observations to assess predictive accuracy. We attributed the failure of all models to their inability to capture shifts in the specific nature of nutrient limitation. Our results point to more complex linkages between multinutrient limitation and cellular resource allocation than currently parameterized in models. These results suggest a greater importance of understanding the interaction between the type of nutrient limitation and plankton diversity for predicting the global variation in surface C:N:P. Plain Language SummaryCompiled observations of particulate organic matter elemental ratios indicate conservation of N and P where nutrients are scarce, and vice versa in nutrient-rich upwelling and polar regions. However, because the compiled datasets vary in methodology, mesoscale trends are unable to be resolved. In the current study, we observe strong gradients in particulate organic matter (POM) C:N:P ratios using consistent methods for a latitudinal transect in the eastern Pacific Ocean. Single environmental factors were unable to predict variation in C:N:P across regions suggesting a more complex regulation. Ratios of C:N and C:P in the South Pacific Subtropical Gyre were unexpectedly high for a subtropical gyre in the southern hemisphere. A single-nutrient model (nitrate or phosphate) produced significant regional biases, leading us to hypothesize multiple-nutrient models as necessary under conditions of severe nutrient stress. In the Southern Ocean, we measured total and small size fractions to estimate significantly lower C:N:P ratios of larger POM. The N:P ratio of large POM are nearest to the N:P ratio of exported organic matter estimated from remineralized nutrients in the subsurface. This analysis will help evaluate the regional importance of temperatures, n...
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