By:Daniel M. Sigman(Department that Geosciences, Princeton University)&Mathis P. Hain(Department the Geosciences, Princeton University)©2012lifwynnfoundation.org Education
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Citation:Sigman,D.M.&Hain,M.P.(2012)The biological Productivity of the Ocean.lifwynnfoundation.org education and learning Knowledge3(10):21
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Productivity fuel life in the ocean, drives its chemistry cycles, and also lowers atmospheric carbon dioxide. Nutrient uptake and also export communicate with circulation come yield unique ocean regimes.

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What is s Productivity?

ocean productivity mostly refers come the production of organic issue by "phytoplankton," tree suspended in the ocean, many of which are single-celled. Phytoplankton space "photoautotrophs," harvesting light to convert inorganic to organic carbon, and they supply this necessary carbon to varied "heterotrophs," biology that attain their power solely native the respiration of organic matter. Open ocean heterotrophs incorporate bacteria and more complicated single- and multi-celled "zooplankton" (floating animals), "nekton" (swimming organisms, consisting of fish and also marine mammals), and also the "benthos" (the seafloor ar of organisms).

The plenty of nested cycles that carbon linked with ocean productivity space revealed through the following meanings (Bender et al. 1987) (Figure 1). "Gross major production" (GPP) describes the full rate that organic carbon production by autotrophs, while "respiration" describes the energy-yielding oxidation the organic carbon back to carbon dioxide. "Net major production" (NPP) is GPP minus the autotrophs" very own rate the respiration; that is thus the price at i beg your pardon the complete metabolism the phytoplankton produce biomass. "Secondary production" (SP) generally refers to the growth rate of heterotrophic biomass. Just a small fraction of the organic matter ingested by heterotrophic biology is provided to grow, the majority being respired ago to dissolved not natural carbon and also nutrients that deserve to be reused through autotrophs. Therefore, SP in the ocean is little in comparison come NPP. Fisheries depend on SP; hence they rely on both NPP and also the efficiency with i m sorry organic matter is moved up the foodweb (i.e., the SP/NPP ratio). "Net ecosystem production" (NEP) is GPP minus the respiration by all organisms in the ecosystem. The worth of NEP counts on the boundaries defined for the ecosystem. If one considers the sunlit surface ocean down to the 1% light level (the "euphotic zone") end the food of an entire year, then NEP is tantamount to the particulate organic carbon sinking into the dark ocean inner plus the liquified organic carbon being circulated the end of the euphotic zone. In this case, NEP is likewise often described as "export production" (or "new production" (Dugdale & Goering 1967), as disputed below). In contrast, the NEP because that the whole ocean, including its shallow sediments, is about equivalent to the slow interment of organic matter in the sediments minus the rate of organic matter entering native the continents.


Productivity in the surface ocean, the definitions used to describe it, and also its connections to nutrient cycling. The blue cycle for “net ecosystem production” (NEP) (i.e. “new” or “export” production) includes the “new” nutrient it is provided from the ocean interior, its absorb by autotrophic phytoplankton growth, packaging into large particles by heterotrophic grazing organisms, and also sinking that organic issue out the the surface ocean. The red bike illustrates the fate that the majority of essential matter developed in the surface ar ocean, i beg your pardon is to it is in respired by heterotrophic biology to accomplish their energy requirements, in order to releasing the nutrients back into the surface ar water whereby they can be taken increase by phytoplankton once again come fuel “regenerated production.” The environment-friendly cycle to represent the inner respiration of phytoplankton themselves, that is, their own use the the products of photosynthesis for purposes various other than growth. This nested cycles integrate to yield (1) “gross major production” (GPP) representing the pistol photosynthesis and (2) “net major production” (NPP) the represents phytoplankton biomass production that develops the basis of the food net plus a lot smaller rate of organic issue export indigenous the surface. When the brand-new nutrient supply and export production are ultimately linked by fixed balance, there might be inequalities on small scales of an are and time, permitting for short accumulations that biomass.

There are no accumulations of life biomass in the marine atmosphere that compare with the forests and grasslands on floor (Sarmiento & Bender 1994). Nevertheless, s biology is responsible for the warehouse of more carbon far from the environment than is the terrestrial biosphere (Broecker 1982). This is accomplished by the sinking the organic matter out that the surface ar ocean and also into the ocean interior before it is returned to liquified inorganic carbon and dissolved nutrient by bacterial decomposition. Oceanographers frequently refer to this process as the "biological pump," as it pumps carbon dioxide (CO2) out of the surface ocean and atmosphere and into the voluminous deep s (Volk & Hoffert 1985).

Only a portion of the organic matter produced in the surface ar ocean has actually the fate of gift exported to the deep ocean. The the organic matter produced by phytoplankton (NPP), most is respired ago to dissolved inorganic creates within the surface ocean and thus recycled for use by phytoplankton (Eppley & Peterson 1979) (Figure 1). Most phytoplankton cells room too small to sink individually, so sinking occurs only as soon as they aggregate into larger particles or are packaged into "fecal pellets" by zooplankton. The continues to be of zooplankton are additionally adequately big to sink. While sinking is a reasonably rare fate for any kind of given fragment in the surface ar ocean, biomass and also organic issue do no accumulate in the surface ocean, therefore export that organic matter by sinking is the ultimate fate for every one of the nutrients that enter into the surface ocean in dissolved form — v the exceptions that (1) dissolved nutrients have the right to be went back unused come the inner by the circulation in some polar areas (see below), and (2) circulation additionally carries dissolved organic issue from the surface ocean into the interior, a far-reaching process (Hansell et al. 2009) the we will certainly not attend to further. Together organic issue settles through the ocean interior and onto the seafloor, the is nearly entirely decomposed earlier to dissolved chemicals (Emerson & Hedges 2003, martin et al. 1987). This high efficiency of decomposition is as result of the truth that the organisms moving out the decomposition count upon it as their sole source of chemistry energy; in most of the open up ocean, the heterotrophs just leave behind the organic issue that is as well chemically resistant for it to be precious the invest to decompose. ~ above the whole, just a tiny fraction (typically much less than 1%) that the organic carbon from NPP in the euphotic zone survives to be buried in deep sea sediments.

Productivity in coastal ecosystems is often distinct from the of the open ocean. Along the coasts, the seafloor is shallow, and also sunlight can sometimes permeate all the way through the water tower to the bottom, thus allowing bottom-dwelling ("benthic") biology to photosynthesize. Furthermore, sinking organic issue isintercepted by the seabed, wherein it supports growing benthic faunal communities, in the process being recycled back to dissolved nutrients that room then immediately obtainable for main production. The proximity come land and its nutrient sources, the interception that sinking organic matter by the shallow seafloor, and also the propensity for seaside upwelling all result in highly productive ecosystems. Here, us mainly deal with the performance of the large open ocean; nevertheless, countless of the exact same concepts, albeit in modified form, apply to seaside systems.

What Does ocean Productivity Need?

Phytoplankton call for a suite the chemicals, and also those through the potential to it is in scarce in surface ar waters are commonly identified as "nutrients." Calcium is an instance of an element that is swiftly assimilated by part plankton (for production of calcium carbonate "hard parts") however is not generally considered a nutrient due to the fact that of that is uniformly high concentration in seawater. Dissolved not natural carbon, which is the feedstock for organic carbon manufacturing by photosynthesis, is also abundant and also so is not typically listed among the nutrients. However, its acidic form dissolved CO2 is regularly at adequately short concentrations to influence the expansion of at the very least some phytoplankton.

Broadly crucial nutrients include nitrogen (N), phosphorus (P), iron (Fe), and silicon (Si). There appear to be fairly uniform demands for N and also P among phytoplankton. In the early on 1900s, oceanographer Alfred Redfield discovered that plankton develop their biomass with C:N:P stoichiometric ratios the ~106:16:1, to which we currently refer together the Redfield ratios (Redfield 1958). As Redfield noted, the liquified N:P in the deep s is close to the 16:1 ratio of plankton biomass, and we will certainly argue below that plankton i charged this ratio on the deep, no vice versa. Steel is uncovered in biomass just in trace amounts, yet it is supplied for varied essential objectives in organisms, and also it has come to be clear over the last 25 years the iron"s scarcity often limits or affects performance in the open up ocean, specifically those regions where high-N and also -P deep water is lugged rapidly come the surface ar (Martin & Fitzwater 1988). Research is ongoing to understand the role of other trace facets in efficiency (Morel et al. 2003). Silicon is a nutrient just for particular plankton taxa-diatoms (autotrophic phytoplankton), silicoflaggellates, and radiolaria (heterotrophic zooplankton) — which usage it to do opal tough parts. However, the usual dominance of cat birds in Si-bearing waters, and the propensity of diatom-associated organic issue to sink the end of the surface ocean, make Si accessibility a major factor in the broader ecology and also biogeochemistry of surface waters.

Sunlight is the can be fried energy resource — straight or indirectly — for almost all life top top Earth, consisting of in the deep ocean. However, light is took in and scattered such the very tiny of the penetrates below a depth that ~80 m (as deep together 150 m in the least abundant subtropical regions, however as shallow together 10 m in highly abundant and seaside regions) (Figure 2). Thus, photosynthesis is largely restricted to the top light-penetrated skin the the ocean. Moreover, across most that the ocean"s area, consisting of the tropics, subtropics, and also the temperate zone, the absorb of sunlight reasons surface water come be much warmer 보다 the basic deep ocean, the latter being filled v water that sank from the surface ar in the high latitudes . Warmth water is more buoyant than cold, which causes the top sunlit layer to float on the denser deep ocean, v the change between the two recognized as the "pycnocline" (for "density gradient") or "thermocline" (the upright temperature gradient that drives density stratification throughout most that the ocean, number 2). Wind or another resource of power is compelled to journey mixing throughout the pycnocline, and also so the carry of water with its dissolved chemicals between the sunlit surface and also the dark internal is sluggish. This double effect of light on photosynthesis and also seawater buoyancy is an important for the success of s phytoplankton. If the s did not have a thin buoyant surface layer, mix would bring algae out of the light and thus away from your energy source for most of the time. Rather of nearly neutrally buoyant single celled algae, larger, positively buoyant photosynthetic organisms (e.g., pelagic seaweeds) might dominate the open up ocean. This hypothetical case aside, although viable phytoplankton cell are discovered (albeit at low concentrations) in depths waters, photosynthesis limits active phytoplankton development to the top skin of the ocean, while upper ocean density stratification prevents them indigenous being blended down into the dark abyss. Thus, many open ocean biomass, including phytoplankton, zooplankton, and nekton, is found within ~200 m the the ocean surface.


Typical problems in the subtropical ocean, as indicated by data collected at the Bermuda Atlantic Time-series terminal in July, 2008. The thermocline (vertical temperature gradient) stratifies the upper water column. Throughout this certain station occupation, the shallow wind-mixed surface layer is not well defined, presumably due to the fact that of strong insolation and also a lack of wind that allowed consistent stratification every the means to the surface. Very small sunlight penetrates deeper 보다 ~100 m. New supply of the major nutrients N and also P is restricted by the slow mixing across the top thermocline (showing below only the N nutrient nitrate, NO3-). In ~ the top euphotic zone, the slow nutrient it is provided is fully consumed through phytoplankton in your growth. This expansion leads to the accumulation of particulate necessary carbon in the surface ocean, several of which is respired by bacteria, zooplankton, and also other heterotrophs, and also some of i m sorry is exported as sinking material. The deep chlorophyll preferably (DCM) occurs at the contact where over there is adequate light because that photosynthesis and yet significant nutrient it is provided from below. The DCM have to not it is in strictly taken as a depth best in phytoplankton biomass, as the phytoplankton at the DCM have actually a an especially high interior chlorophyll concentration. The data shown here is made obtainable the Bermuda institute of s Sciences (http://bats.bios.edu) and also the Bermuda Bio Optics task (http://www.icess.ucsb.edu/bbop/).

At the same time, the presence of a slim buoyant surface ar layer conspires with other processes to impose nutrient limitation on s productivity. The export the organic matter to depth depletes the surface ocean of nutrients, resulting in the nutrients to accumulate in deep waters whereby there is no light obtainable for photosynthesis (Figure 2). Due to the fact that of the thickness difference between surface water and also the deep sea across most the the ocean, ocean circulation can only an extremely slowly reintroduce liquified nutrients come the euphotic zone. By driving nutrients out the the sunlit, buoyant surface ar waters, s productivity effectively limits itself.

Phytoplankton growth limitation has traditionally been understood in the context of Liebig"s regulation of the Minimum, which states that plant growth will be as good as allowed by the least accessible resource, the "limiting nutrient" that sets the productivity of the device (de Baar 1994). If this watch is powerful, interactions amongst nutrients and between nutrients and light can additionally control productivity. A basic but crucial example the this potential for "co-limitation" originates from polar regions, wherein oblique solar insolation combines through deep mixing of surface waters come yield short light availability. In together environments, higher iron supply can increase the performance with i m sorry phytoplankton record light energy (Maldonado et al. 1999, Sunda & Huntsman 1997). More broadly, it has been suggested that phytoplankton should typically seek a state of co-limitation by every the chemistry they require, consisting of the plenty of trace metal nutrients (Morel 2008).

Who are the significant Players in s Productivity?

In comparison to the terrestrial biosphere, most naval photosynthesis is performed by single-celled organisms, and also the much more abundant the the multicellular forms are structurally much easier than the vascular plants on land. Throughout much that the twentieth century, the was believed that cells in the selection of ~5 come ~100 microns diameter account for most phytoplankton biomass and productivity. This size range is composed mostly of eukaryotes, organisms whose cell contain complex membrane-bound structures ("organelles"), consisting of the cell"s nucleus and also chloroplasts. Well-studied develops of eukaryotic phytoplankton include the opal-secreting diatoms, prymnesiophytes (including the CaCO3-secreting coccolithophorids), and also the necessary wall-forming dinoflagellates. The centrality of these organisms in beforehand oceanographic assumed was as result of their ease of access by typical light microscopy.

Only v recent technical advances have smaller organisms come to be readily observable, revolutionizing our view of the plankton. In particular, the cyanobacteria, which space prokaryotes (lacking a nucleus and also most other organelles uncovered in eukaryotes), are now well-known to it is in important amongst the phytoplankton. Initially, the cyanobacteria were identified largely with early american forms such together Trichodesmium that play the an essential role of "fixing" nitrogen (see below). However, significant discoveries over the last thirty years have revealed the prevalence across the global ocean of imreparable cells cyanobacteria of ~0.5 to ~1.5 microns diameter. That is now well-known that 2 cyanobacterial genera — Synechoccocus and also Prochlorococcus — dominate phytoplankton numbers and also biomass in the nutrient-poor tropical and subtropical ocean (Waterbury et al. 1979, Chisholm et al. 1988). In addition, brand-new methods, both microscopic and genetic, room revealing a formerly unappreciated diversity of smaller eukaryotes in the open up ocean.

Mapping ecological and biogeochemical functions onto the genetic diversity that the phytoplankton is an energetic area in biological and also chemical oceanography. Based on observations and theory, the smaller phytoplankton such together the uncellular cells cyanobacteria room thought to conquer regenerated production in countless systems, conversely, the bigger eukaryotes show up to beat a more important duty in brand-new production (i.e., NEP, number 1; see below).

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Just as large eukaryotes were once thought to conquer the phytoplankton, it to be long believed that multicell zooplankton of ≥200 microns dominate heterotrophy — the tiny crustaceans recognized as copepods space the prototypical example. We now know that heterotrophy is often dominated by single-celled standard scale ("microzooplankton," the ~1 come ~200 microns) and also by bacteria (of ~0.3 to ~1 microns), the last carrying out many of the organic carbon decomposition in the ocean.

The food resource of a given type of zooplankton is frequently driven by its own size, through microzooplankton grazing ~ above the prokaryotes and smaller eukaryotes and multicellular zooplankton grazing on larger eukaryotes, both phytoplankton and microzooplankton. Since of their family member physiological simplicity, microzooplankton are thought to be highly reliable grazers that strongly border the biomass accumulation of your prey. In contrast, the multicell zooplankton, due to the fact that they frequently have more complicated life histories, can lag behind the proliferation of their prey, permitting them come bloom and also sometimes stop predation altogether and sink directly. The multicellular zooplankton additionally often facilitate the manufacturing of sinking organic matter, because that example, with the manufacturing of fecal pellets by copepods.


The diversity the the plankton interacts with open ocean environmental problems to affect the performance of the bigger ecosystem (Michaels & silver 1988, Morel et al. 1991, Buesseler 1998) (Figure 3). In the nutrient-poor tropical and also subtropical ocean, the (small) cyanobacteria tend to it is in numerically dominant, perhaps because they specialize in acquisition up nutrient at low concentrations. Tiny phytoplankton have a greater surface area-to-volume proportion than do big phytoplankton. A better proportional surface ar area disclosure the absorb of nutrients throughout the cell boundary, a an important process when nutrients space scarce, most likely explaining why small phytoplankton conquer the biomass in the nutrient-poor ocean. The microzooplankton properly graze these tiny cells, staying clear of their biomass indigenous accumulating and sinking directly. Moreover, these single-celled microzooplankton lack a digestive tract, so they do not produce the fecal pellets that stand for a major mechanism the export. Instead, any residual organic issue remains in the upper ocean, to it is in degraded by bacteria. Every told, microzooplankton grazing that phytoplankton biomass leads to the remineralization of many of its had nutrients and also carbon in the surface ocean, and also thus rises recycling loved one to organic matter export. This an extremely efficient recycling elevates NPP loved one to NEP, yielding a short NEP:NPP ratio (~0.05–0.3) in nutrient-poor systems (Figure 3a). In contrast, bigger phytoplankton, such together diatoms, often conquer the nutrient-rich polar ocean, and also these deserve to be grazed directly by multicellular zooplankton. By growing adequately quickly to outstrip the grazing rates of these zooplankton, the diatoms have the right to sometimes accumulate come high concentrations and produce abundant sinking material. In addition, the zooplankton export organic issue as fecal pellets. In these abundant systems, the less extensive upper s recycling reasons NEP and also NPP come be an ext similar, through an NEP:NPP ratio frequently near 0.5 (Figure 3b).


The most extensively accepted paradigm for the controls on surface nutrient recycling efficiency. NPP is supported by both new nutrient supply from the deep ocean and also nutrients regenerated within the surface ocean. The fraction of NEP:NPP ratio appears to vary v the nutrient supply, because links come the ecology the the plankton. In the nutrient-poor tropical and subtropical s (a), the (small) cyanobacteria tend to be numerically dominant. The microzooplankton the graze these small cells do so effectively, preventing phytoplankton native sinking directly. Moreover, these single-celled microzooplankton perform not create sinking fecal pellets. Instead, any kind of residual organic issue remains to be degraded through bacteria. This rises recycling loved one to organic issue export, yielding a low NEP:NPP proportion (~0.1). In nutrient-rich areas (b), big phytoplankton are much more important, and also these can be grazed straight by multicell zooplankton. By farming adequately promptly to outstrip the grazing rates of zooplankton, the large phytoplankton can sometimes accumulate come high concentrations and produce abundant sinking material. In addition, the zooplankton violin organic matter as fecal pellets. In these fertile systems, the less intensive upper s recycling causes NEP and NPP come be more similar, through an NEP:NPP ratio regularly near 0.5. The relationships between nutrient supply, phytoplankton size, and sinking thus overcome this watch of upper s nutrient cycling.