The low-latitude ocean regions, considered as the expanse between the subtropical convergence zones (30°S-30°N), are important for both their rich biodiversity as well as for fisheries and food security1-3. Both marine primary productivity (PP) and export over this region account for approximately half of their respective global totals4. Large PP and export occur in low-latitude oceans despite thermocline macronutrient concentrations being less than half of what is found in subpolar/circumpolar regions, indicating vigorous decadal renewal timescales for thermocline waters through the shallow overturning circulation5-7. An Earth system model has previously been considered under strong anthropogenic forcing (historical/RCP85) to argue that low-latitude (30°S-30°N) export and PP could decrease by 41% and 24% respectively by 2300 relative to the present climate state8. This collapse was attributed to a cascade of processes, initially triggered by sea ice retreat over the Southern Ocean, which in turn induced Southern Ocean subsurface nutrient trapping. Reduced surface nutrient concentrations in the Southern Ocean in turn lead to reduced nutrients exported northward to the low-latitude thermocline in mode and intermediate waters. This work is important in emphasizing a first-order role for a low-latitude response in PP and export under multi-century climate perturbations. The interpretation, however, invoked the mechanisms presented in an earlier analysis9, which found that 75% of low-latitude new production is contingent on a sustained supply of nutrients from the Southern Ocean using an ocean biogeochemistry model. A subsequent study with an inverse ocean biogeochemical model identified a contribution of 44% to PP to the north of 40°S as being due to Southern Ocean nutrient trapping10, thereby also arguing for a first-order role for the Southern Ocean. However, independent observational analyses indicate that at least for the case of the Pacific basin, the Subantarctic Mode Water water mass cannot readily access regions north of 20°S due to strong potential vorticity barriers11.
We are thereby motivated to propose that low-latitude PP and export are sustained primarily by processes local to the tropics, and that the sensitivity to non-local perturbations over the Southern Ocean is more modest than that previously proposed 9. In particular, we propose that remineralization within the low-latitude mesopelagic domain (LLMD) (defined here to span 30°S-30°N and 150m-870m) and rapid subsequent re-emergence of nutrients through the shallow overturning structures within the low-latitudes are the dominant driver of low-latitude PP and export. This is investigated through the combined use of in-situ observations and a new suite of model sensitivity studies with a fully prognostic global ocean biogeochemistry model.
We examine the extent to which local nutrient recycling in the tropics contributes to sustaining low-latitude PP and export by quantifying its contribution to the total low-latitude mesopelagic nutrient inventory (here we use phosphate, or PO43-) using an observational product based on hydrographic measurements, GLODAPv212,13. We invoke a well-known diagnostic that decomposes total PO43- concentrations14,15 into two components: a preformed component that represents the PO43- concentrations at the surface when a new water mass is transferred from the surface into the interior by subduction, and a regenerated component that reflects the cumulative impact of respiration of organic matter that increases PO43- concentrations as the water mass is transported in the ocean interior.
The distribution of PO43- on the mid-thermocline density horizon s0=25.5 kg m-3 in the ocean interior (Fig. 1a) reveals concentrations that are maximum in the equatorial band. Over the same density horizon the regenerated fraction of PO43 (Fig. 1b) is larger than 50 percent over much of the region between 20°S and 20°N, with exceptions (minimum values) neighboring subduction regions. On the density layer associated with the upper boundary of Subantarctic Mode Water (SAMW) formed in the Southern Ocean and entering the Pacific Ocean just below the thermocline (s0=26.8 kg m-3)16, the PO43- concentrations are higher than in the thermocline (Fig. 1c). Additionally, the regenerated fraction of PO43- is modestly higher than 50% throughout most of the tropics (Fig. 1d). Viewed north-south along 150°W in the Pacific, despite the large vertical gradients in PO43- concentrations across the equatorial thermocline (Fig. 1e), the regenerated fraction exhibits remarkable homogeneity with values just above 50% over the region 10°S-10°N and 150m-700m. Further sensitivity analysis with varying stochiometric ratios of O2: PO43- indicates that the substantial regenerated pool of PO43- identified here is relatively insensitive to our choice of a ratio (Extended Data Fig. 1),
Here we hypothesize that the large regenerated nutrient pool in the low-latitudes provides a dominant control on the elevated low-latitude PP and export through local processes. To test this hypothesis, we apply a set of perturbation simulations with the PISCES-v2 biogeochemistry model17 (see methods) that all use identical circulation fields but that vary by perturbing nutrient remineralization in distinct latitude bands. In PISCES-v2, the export of organic material from the euphotic zone to the interior of the ocean is represented by the gravitational sedimentation of two particulate pools that differ by their mean size, and thus by their sinking speed. These particulate pools are continuously solubilized to dissolved organic matter according to a temperature-dependent rate. This dissolved pool is then ultimately remineralized to inorganic nutrients. In addition to a standard experiment (STD)17, we performed a series of perturbation experiments in which remineralization of nutrients and solubilization of organic carbon is disallowed within specific latitude-bounded regions (see Methods for model perturbation details). For PERT_TROP, solubilization was disallowed within the tropics over the depths comprising mode and intermediate waters (30°S-30°N, 150m-1500m). For PERT_SOUTH and PERT_NORTH, solubilization was disallowed between 150m and the next-to-bottom grid-box of the model domain, over 90°S-30°S and 30°N-90°N, respectively.
A comparison of the STD and TROP_PERT experiments reveals a large reduction in PP and export over the low-latitude domain when local mesopelagic remineralization is disallowed (Fig. 2). For STD, PP and export are 21.5 PgC yr-1 and 2.73 PgC yr-1, respectively, when integrated over 30°S-30°N. For PERT_TROP, PP and export are 7.3 and 1.37 PgC yr-1, respectively, over the same domain. Thus PERT_TROP represents a 66% drop in PP and a 50% drop in export relative to STD over 30°S-30°N. These changes are larger than the 17% reduction in PP and 38% decrease in export found for the PERT_SOUTH case (maps showing perturbation patterns for PERT_SOUTH and PERT_NORTH are shown in Extended Data Figs. 2 and 3, respectively). Proper attribution of the strong sensitivities under PERT_TROP relative to STD is facilitated by a coarse-graining of the modeled PO43- budget in the LLMD spanning 30°S-30°N and 150m-870m (Fig. 3; the full budgets for all four simulations are shown in Extended Data Fig. 4). For STD (budgets in black) PP is 21.5 PgC yr-1, the vertical flux of soft tissue and dissolved organic phosphorous (DOP) into the LLMD is 2.73 PgC yr-1, while the soft tissue flux out of the LLMD at 870m is 0.88 PgC yr-1 (DOP fluxes are negligible), with a net solubilization source within the LLMD (largest in the shallower thermocline layers) of 1.72 PgC yr-1. As such, the regenerated PO4 represents 63% of the net upward flux of PO43- across the 150m horizon within the tropics. The net meridional fluxes (the sum of both northward and southward fluxes) of PO43- across 30°S and 30°N, on the other hand, present a net divergence that depletes PO43- over the LLMD. For the PERT_TROP case (red numbers) low-latitude remineralization was disallowed between 150m-1500m, and the -0.1 PgC yr-1 of interior production corresponds to a degree of PP occurring just below 150m depth. The increase in the upward flux of PO43- across 870m by 0.12 PgC yr-1 and the net reduction of meridional divergence across 30°S and 30°N by 0.2 PgC yr-1 indicate that there are feedbacks that reduce the loss of PO43- and thereby dampen the drop in export for PERT_TROP.
The PERT_TROP perturbation relative to STD (Fig. 3) underscores that the dominant process sustaining elevated low-latitude PP and export is local resolubilization (mainly through remineralization) of PO43- locally within the local LLMD, in conjunction with the subsequent local re-emergence into the low-latitude euphotic zone. The upward net flux of PO43- across 870m into the LLMD plays a critical supporting role in balancing not only the downward particle flux across 870m, but also in compensating for the net meridional divergence of PO43- out of the LLMD across 30°S and 30°N. A previous study18 considered a role for remineralization in sustaining low latitude new production, but under the assumption that regenerated nutrients are a downstream legacy of new/preformed nutrients supplied through SAMW, thereby not accounting for a sizable low-latitude remineralized pool of PO43- sustained through local low-latitude processes. The same study also ruled out a role for supply from the deep ocean.
In our interpretation, the remineralized low-latitude pool is a legacy of the deeper low-latitude upwelling of preformed PO43- that is originally sourced in the deep-water formation regions of the Southern Ocean. The net upwelling flux of 8 Sv across 870m in the global tropics (Extended Data Fig. 5) in the model circulation state is consistent with a deep upwelling cell in the low-latitudes that has been identified with observational constraints19 and is also characteristic of a broad range of climate models20,21. In fact the main impact of our PERT_SOUTH experiment is to enhance the net divergence (0.4 PgC yr-1 relative to STD) of the regenerated PO43- pool from the low-latitudes (Extended Data Fig. 4).
Our results also motivate us to reconsider previous modeling work showing substantial low-latitude PP and export decreases under sustained warming to the year 2300 8. Here we consider five CMIP6 models: IPSL-CM6A22, CESM2-WACCM23, UKESM124, ACCESS-ESM1.525, and MIROC-ES2L26 , run to 2300 under high emissions forcing (SSP5-8.5)27. Anomalies of primary production for each model relative to the period 1850-1900 are shown in Fig. 4a (corresponding thermal fields are shown in Extended Data Fig. 6), revealing a pronounced divergence in not only the amplitude of PP but also the sign of the response. This divergence of the global response across CMIP6 models is best interpreted as a measure of the very large uncertainty in PP, under the assumption that each of the models presents an equally likely outcome under strong and persistent anthropogenic climate perturbations. The divergence of the globally integrated primary production (Fig. 4a) is in fact largely sustained over 30°S-30°N (Fig. 4b), with the low-latitude response reflecting an underlying diversity of low-latitude mechanisms or process-parameterizations28,29. As the increase in PP over 30°S-30°N for two of the models (IPSL-CM6A and ACCESS-ESM1.5) is not at face value consistent with the Southern Ocean-triggered nutrient trapping mechanism emphasized previously8, we address whether the low latitude nutrient recycling mechanism highlighted in Figs. 2 and 3 may be contributing.
To this end, we consider the fractional changes in zonal mean ocean interior PO43- concentrations for the cases of the IPSL-CM6A (Fig. 4c) (increasing low-latitude PP) and CESM2-WACCM (Fig. 4d) (decreasing low-latitude PP) models between the 1990s and 2290s. For a mean over the 1990s with both models, the transfer efficiency defined as the ratio of the soft tissue flux at 780m relative to 150m, is 27% for IPSL-CM6A and 29% for CESM2-WACCM over the tropics (Extended Data Tables 1 and 2). However, by the 2290s, transfer efficiency over the tropics has reduced to 21% for IPSL-CM6A (enhanced thermocline nutrient retention) and has increased to 34% in CESM2-WACCM (decreased thermocline nutrient retention), as is reflected in the perturbations to zonally averaged PO43- between the 1990s and 2290s for both models in Fig. 4. This difference between the two models reflects the fact that remineralization in PISCES-v2 is temperature-dependent and thereby enhanced under sustained warming, whereas for the MARBL biogeochemistry component of CESM2-WACCM the temperature dependence of remineralization remains static for CMIP6 transient simulations30. As such, these differences in representation of remineralization and ensuing thermocline nutrient retention contribute to a modest increase in export for IPSL-CM6A of 0.3% by the year 2300, and a reduction by 38% for CESM2-WACCM. This underscores the importance of improved process-understanding and model representation of remineralization for projecting future changes in marine biological production and export31, and as a consequence the ability of marine ecosystems to sustain food provision and other services32.