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Chapter 43 : The Marine Phosphorus Cycle

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Abstract:

Field investigation of the marine phosphorus (P) cycle requires the use of a variety of methods to measure the ambient concentrations of total dissolved and particulate (both organic and inorganic) matter to assess local inventories of P and to estimate P fluxes. The latter include the delivery to and losses from the ecosystem in question and the rates of microbial P uptake and microbial decomposition of P-containing organic matter. In seawater, the most commonly measured P pools are (i) soluble reactive P (SRP), (ii) total dissolved P (TDP), and (iii) particulate P (PP). The structure and function of the marine P cycle are both time and space (vertical and horizontal) variable; a comprehensive understanding will require a four-dimensional resolution of the key P inventories and fluxes. Measurements of P-ATP in marine ecosystems date back to the classic work of Holm-Hansen and Booth, who developed the theoretical basis for its use in total microbial biomass estimation. Marine microorganisms assimilate D-ATP for P and for purine salvage. The central portions of all major ocean basins are characterized as “marine deserts” because of the small standing stocks of photosynthetic organisms and small nutrient inventories.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43

Key Concept Ranking

Bacteria and Archaea
0.5680074
Mesopelagic Zone
0.4555832
Marine Ecosystem
0.42479113
Horizontal Gene Transfer
0.4128723
0.5680074
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Figures

Image of FIGURE 1
FIGURE 1

Schematic representation of the open ocean P cycle showing the major pools (boxes) and transformations and fluxes (arrows). The large box in the center, TDP, is comprised of phosphate (P), inorganic polyphosphate (poly-P), and DOP. The last group is usually the largest proportion of TDP in near-surface waters. Although the large DOP pool is poorly characterized, at least a portion of it is available as a source of P for microbial growth. Particulate P includes both living and nonliving organic P (POP) and PIP; POP is the dominant component. Ectoenzymatic activity (Ecto) may be important for the microbial assimilation of DOP into POP. Whereas euphotic-zone processes are dominated by the net conversion of P to organic P, subeuphotic-zone (>200 m) P cycle processes are dominated by the net remineralization of organic P to P. A number of physical processes, as shown, are responsible for redistributing P throughout the global oceans. In the marine environment, P is almost exclusively in the pentava-lent state (+5) as PO ; only under highly reduced conditions is P transformed to phosphine gas (PH).

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 2
FIGURE 2

Schematic representation of the stepwise analysis of P in seawater. From three direct measurements, SRP, TDP, and PP, one can derive total P (TDP + PP) and DOP (TDP – SRP). Additional analytical procedures exist for the measurement of inorganic polyphosphates, for the separation of PP into organic and inorganic components, and for partial chemical and/or molecular characterization of the TDP and PP pools. However, SRP, TDP, and PP are the most commonly measured P cycle constituents.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 3
FIGURE 3

Annual mean phosphate (SRP) concentrations at the ocean’s surface, based on the worldwide data set held at NODC. Surface SRP concentrations range from <0.1 µM in the oligotrophic, central gyres of the major ocean basins to >1 µM at high latitudes, especially in the Southern Ocean and North Pacific Ocean. (From the Data are publicly available at http://www.nodc.noaa.gov.)

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 4
FIGURE 4

Relationship between vertical distributions of SRP and dissolved oxygen (O) at Station ALOHA in the North Pacific gyre. (Left) Graph of SRP (µM) versus depth (m) showing the characteristic “nutrient-like” distribution of SRP, with regions of net P uptake and export near the surface and net P remineralization at greater depths. The inset shows these main P cycle processes, which are most intense in the upper 1,000 m of the water column. (Right, top) SRP and O concentration-versus-depth profile of the 300- to 700-m region of the water column at Station ALOHA showing the effects of net remineralization of organic matter. (Right, bottom) Model 2 linear regression analysis of SRP versus O suggesting an average consumption of 80 µmol liter O for each 1 µmol of P regenerated.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 5
FIGURE 5

SRP-versus-depth profiles for the North Atlantic Ocean (Bermuda Atlantic Time-Series Study [BATS]) and the North Pacific Ocean (HOT) showing significant interocean differences, including a steeper phosphocline gradient (i.e., a larger change in SRP concentration per meter in the upper mesopelagic zone region) and higher deep-water (>4,000 m) SRP concentrations for HOT. These differences in SRP inventories and gradients have significant implications for SRP fluxes into the euphotic zone.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 6
FIGURE 6

Map showing the locations of World Ocean Circulation Experiment program transects A-16 (Atlantic) and P-15 (Pacific). Data from these cruises were obtained from http://woce.nodc.noaa.gov, averaged over the depth range of 500 to 1,500 m, and then combined into 10° latitude bins and plotted as mean SRP concentrations (± 1 standard deviation). The resultant plot shows a systematic increase in SRP concentrations at midwater depths “down” the Atlantic and “up” the Pacific. This result is in accordance with the known pathways of deep water circulation and the increasing age of the deep water mass. The accumulation of SRP is a result of the time-integrated decomposition of organic matter. Deg, degrees.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 7
FIGURE 7

Concentration-versus-depth profiles for SRP (µM) and DOP (µM) at Station ALOHA. Also shown is DOP as a percentage of TDP.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 8
FIGURE 8

Concentration-versus-depth profiles of particulate P (nM) and biomass P (nM) at Station ALOHA. Biomass P was derived from P-ATP measurements, assuming a biomass P-to-ATP ratio of 6.1 (see the text for details). Also shown is biomass P as a percentage of PP.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 9
FIGURE 9

Changes in SRP inventories for samples collected at Station ALOHA during the period from 1989 to 2004. (Top) Contour plot of SRP concentrations in the upper 0 to 60 m of the water column. (Bottom) Depth-integrated (0 to 60 m) inventories of SRP showing a systematic loss of SRP from this ecosystem over the past 15 years.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 10
FIGURE 10

Changes in PP inventories and C:N:P stoichiometry of particulate matter collected at Station ALOHA during the period from 1989 to 2004. The solid trend lines are the best-fit linear regression analyses for each data set. The dashed lines in the PC:PP and PN:PP plots show the “ideal” elemental stoichiometry of 106 C:16 N:1 P described by Redfield et al. ( ).

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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Image of FIGURE 11
FIGURE 11

Temporal changes in the export of particulate matter at Station ALOHA during the period from 1989 to 2004, as measured by drifting sediment traps deployed at a reference depth of 150 m. The heavy lines represent the three-point running means of approximately monthly observations. Presented in the upper two plots are the PC and PN export fluxes showing coherent temporal trends. Presented in the lower plot is the PP export flux, showing a fundamentally different pattern, specifically a systematic decrease in PP export over time. d, day.

Citation: Karl D. 2007. The Marine Phosphorus Cycle, p 523-539. In Hurst C, Crawford R, Garland J, Lipson D, Mills A, Stetzenbach L (ed), Manual of Environmental Microbiology, Third Edition. ASM Press, Washington, DC. doi: 10.1128/9781555815882.ch43
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