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Final JGOFS Newsletter Volume 12, Number 4 November 2004 U.S. JGOFS: A Component of the U.S. Global Change Research Program U.S. JGOFS NEWS Volume 12, Number 4 November 2004 Do We Now Understand The Ocean’s Biological Pump? by Jorge L. Sarmiento, John Dunne and Robert A. Armstrong core problem for global carbon- layer or increases in irradiance. It is a measure of its effectiveness in Acycle research is to explain the fundamentally unstable, disappear- reducing surface nutrients relative to role of the oceanic “biological pump” ing rapidly when resources become subsurface values. We defi ne it more in determining levels of dissolved limited, and has a high ef-ratio.-ratio. WWee specifi cally as the average nitrate inorganic carbon (DIC) in surface wa- now think that a large fraction of level at depths in the range of 100 to ters and the exchange of carbon be- oceanic variability in the ef-ratio-ratio dde-e- 200 meters, minus the average in the tween ocean and atmosphere. Surface pends upon whether a given region top 100 meters, divided by the aver- nutrient concentrations are key to supports just the regeneration system age in the 100- to 200-meter range predicting this exchange because un- or the export pathway as well. (Figure 1). We use nitrate observa- used nutrients in the upper ocean al- Is knowledge of the relationship tions in the upper 100 meters from low associated carbon dioxide (CO2) of ef-ratio-ratio ttoo tthehe sstructuretructure ooff pplank-lank- summertime, when surface nutrients to escape into the atmosphere. What tonic food webs either necessary are at a minimum and the effi ciency have we learned during the JGOFS or suffi cient for understanding the is at its maximum. Thus a region in era that can help us understand what biological pump? We would argue which summer surface-layer nutrient controls the strength and effi ciency that it is neither. levels are equal to deep nutrient of the biological pump? A key concept for analyzing the levels has an effi ciency of 0%, and A new paradigm of ecological biological pump is the distinction be- one in which summer surface-layer systems in the surface ocean has tween its strength and its effi ciency. nutrients are totally depleted has an emerged during the last several years, We defi ne the strength of the pump effi ciency of 100%. focused on the relative roles of the as the magnitude of the export of or- Figure 2a shows that the strength regeneration of nutrients versus their ganic matter. For example, a location of the biological pump and its effi - export. The regeneration system such as the Bermuda Atlantic Time ciency are generally inversely related includes the smallest autotrophs, Series (BATS) station in the Sargasso to each other. For example, the picoplankton that are generally Sea where the export of organic mat- high-latitude regions of the Southern only able to utilize ammonium; ter is 4 moles of carbon per square Ocean have a strong but ineffi cient a microbial loop that consists of meter per year (mol C/m2/yr), has a biological pump. Conversely, the heterotrophic bacteria and the dis- stronger biological pump than one low-latitude subtropical gyres have a solved organic matter on which they with an export of 2 mol C/m2/yr, weak but effi cient biological pump. live, and small protists that graze such as the Hawaii Ocean Time-series While the strength of the biologi- on bacteria and picoplankton. This station (HOT) in the north Pacifi c cal pump is important in determin- system is ubiquitous and stable. It oligotrophic gyre (Figure 1b). ing the export of organic matter has a low ef-ratio,-ratio, defidefi nneded aass tthehe ffrac-rac- The maximum possible strength and thus the biological activity in tion of productivity that is supported of the biological pump at a given the deep ocean, the effi ciency of the by nutrients supplied from external location is limited by the supply of biological pump is the key factor in sources such as the nutricline or the macronutrients in the upwelling determining the balance of carbon atmosphere. water and the stoichiometric ratio of between the atmosphere and the The export pathway, on the carbon to nitrogen to phosphorus in ocean, at least in regions where other hand, comprises the classical the organic matter exported from the the supply of nutrients and carbon diatom-based food chain, including surface. Nitrogen fi xation and aeo- from below is high. This is because copepods and other large zooplank- lian fi xed nitrogen also contribute, it is ultimately the effi ciency of the ton predators. It is opportunistic, but to a lesser extent. biological pump in stripping out as responding rapidly to injections of In contrast, we defi ne the ef- nutrients, shoaling of the mixed fi ciency of the biological pump as Continued on page 2 much carbon as possible from up- its link to the ef-ratio-ratio ttellell uuss aboutabout pump effi ciency. welling waters that determines how biological pump effi ciency? Given If the ef-ratio-ratio ddoesoes notnot indicateindicate tthehe much of the excess carbon stored in the relationship between the export effi ciency of the biological pump, the deep ocean can escape back into pathway and high ef-ratios,-ratios, oonene what does? While ocean circulation the atmosphere. In regions such as might expect high ef-ratio-ratio eenviron-nviron- plays an important role, effi ciency is the Southern Ocean where the bio- ments to have an effi cient biological clearly modulated by other factors. logical pump effi ciency is low, CO2 is pump and vice-versa. In fact, most The regions characterized by low able to escape from the deep waters of the world ocean exhibits the biological pump effi ciency in Figure that upwell there. Thus the average opposite relationship (Figure 2b). For 1d generally correspond to areas atmospheric concentration is higher example, the Southern Ocean has where the supply of nutrients from than it would be if the effi ciency of a high ef-ratio-ratio bbutut a llowow biologicalbiological upwelling and deep wintertime mix- the pump were high. pump effi ciency; the subtropical ing is high. However, there are areas gyres, on the other hand, have a low of high biological pump effi ciency What Governs Pump Effi ciency? -ratio but a highly effi cient biological that have extremely high nutrient What does the JGOFS-era para- pump. Thus knowledge of the ef-ratio-ratio supply, such as coastal regions. digm of the surface ecosystem and does not help us predict biological As a result of JGOFS-era discover- A) Primary Production (mmol C m-2 d-1) B) Particle Export Ratio 500 0.65 80°N 80°N 70 0.40 0.35 40°N 60 40°N 0.30 50 0.25 0° 40 0° 0.20 30 0.15 40°S 40°S 20 0.10 10 0.05 80°S 80°S 0 0.00 0° 100°E 160°W 60°W 0° 100°E 160°W 60°W C)C) Particle Export (mmol C m-2 d-1) D) Biological Pump Efficiency 100 1.0 80°N 80°N 20 0.9 0.8 15 40°N 40°N 0.7 10 0.6 0° 8 0° 0.5 6 0.4 0.3 40°S 4 40°S 0.2 2 0.1 80°S 80°S 0 0.0 0° 100°E 160°W 60°W 0° 100°E 160°W 60°W Figure 1: (a) Primary production estimated by a combination of empirical models using satellite-based estimates of chlorophyll. (b) Particle export ratio (particle export divided by primary production) estimated from satellite-based chlorophyll estimates and temperature distributions using an empirical model developed by John Dunne. (c) Particle export calculated by taking the product of the particle export ratio times the primary production. (d) Estimate of the effi ciency of the oceanic biological pump calculated from climatic average nitrate observations. Effi ciency is 100% when organisms deplete surface nutrients totally. It is 0% when no biological removal of upwelled nutrients occurs. 2 U.S. JGOFS Newsletter – November 2004 ies, we now generally believe that this hypothesis to the global of the region. Mitchell and his col- iron limitation plays a crucial role context, we fi nd that the overall leagues defi ned light limitation, not in limiting the uptake of nutrients, lower supply of iron to the ocean by the usual Sverdrup criterion of most particularly in the high-nutri- in the Southern Hemisphere may the threshold light supply necessary ent, low-chlorophyll areas that occur explain much of the remarkable to get a bloom started, but rather by predominantly in the Southern difference between the waters of the the light supply that is required to Ocean and the Southern Hemisphere Northern Hemisphere and those of deplete nutrients completely. subtropics (Figure 3a) and in the the Southern Hemisphere in terms of Given deep summertime mixing, North and equatorial Pacifi c (Figure biological pump strength, effi ciency frequent cloudiness, phytoplankton 3b). and ef-ratio-ratio (Figure(Figure 2).2). TToo cconfionfi rrmm self-shading and an extremely high What is the role of iron? Because of this hypothesis, we turn to seven nutrient supply, the Southern Ocean the inverse relationship between ef- major mesoscale iron fertilization might never experience total deple- ratio and export effi ciency observed experiments conducted during the tion of nutrients in surface waters in our analysis, we suggest that the JGOFS era. even if the iron supply were ad- critical role of iron is not associated These experiments demonstrate equate.
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