The Phosphorus Cycle Differs From the Biogeochemical Cycles in That

The Phosphorus Cycle Differs From the Biogeochemical Cycles in That

Phosphorus Cycle

The phosphorus cycle therefore plays an important role in regulating principal productivity, the process in which radiant energy is used by primary producers to form organic substances equally food for consumers, every bit in photosynthesis.


Encyclopedia of Microbiology (Tertiary Edition)

Phosphorus Cycle

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Encyclopedia of Microbiology (Third Edition), 2009

Phosphorus Sources, Sinks, and Transport Pathways

phosphorus cycle
encompasses numerous living and nonliving environmental reservoirs and various ship pathways. In tracing the movement of phosphorus in the environment, the interplay between physical and biological processes becomes apparent. In addition to interim equally reservoirs of phosphorus in the environment (as discussed in this section), microbes contribute to the transformation of phosphorus inside other reservoirs such every bit in soil or aquatic environments (see ‘

Microbially mediated processes’).

Inside the Globe’due south chaff, the abundance of phosphorus is 0.10–0.12% (on a weight ground), with the bulk of phosphorus existing every bit inorganic phosphate minerals and phosphorus-containing organic compounds. A phosphate mineral is any mineral in which phosphate anion groups form tetrahedral complexes in clan with cations, although arsenate (



) and vanadanate (



) may also be substituted in the crystalline structure. Apatite is the nigh abundant group of phosphate minerals, comprising hydroxyapatite, fluorapatite, and chlorapatite (
Table 1
). These 3 forms of apatite share nearly identical crystalline structures, but differ in their relative proportions of hydroxide, fluoride, and chloride, each being named for the anion that is virtually abundant in the mineral. Phosphate minerals mostly form in the environment in magmatic processes or through precipitation from solution (which may be microbially mediated), and the chemical composition of the minerals depends on the ion or ions present in solution at the fourth dimension of precipitation. For this reason, it is not uncommon for natural deposits of phosphate minerals to be heterogeneous, rather than equanimous of ane homogeneous type of phosphate mineral. These natural deposits of phosphate minerals are collectively chosen ‘phosphorites’ to reflect variations in their chemical compositions.

Tabular array 1.
Phosphate minerals and their chemical compositions. Apatite is the general term for the three minerals hydroxylapatite, fluorapatite, and chlorapatite

Apatite Ca5(PO4)3(F,Cl,OH)
Hydroxylapatite Ca5(POfour)threeOH
Fluorapatite Ca5(PO4)3F
Chlorapatite Ca5(POiv)3Cl
Frankolite Caten−ab

(PO4)half dozen−x
Lazulite (Mg,Fe)Al2(PO4)2(OH)2
Monazite (Ce,La,Y,Th)POiv
Pyromorphite Pb5(POfour)threeCl
Strengite FePO4·2H2O
Triphylite Li(Atomic number 26,Mn)PO4
Turquoise CuAl6(POfour)4(OH)8·5H2O
Variscite AlPO4·2HtwoO
Vauxite FeAl2(PO4)2(OH)ii·6H2O
Vivianite Fe3(PO4)2·8H2O
Wavellite Al3(PO4)2(OH)3·5H2O

Soils and lake sediments are another terrestrial reservoir of phosphorus, comprising primarily inorganic phosphorus from weathered phosphate minerals, along with organic phosphorus from the decomposition, excretion, and lysis of biota (
Effigy one
). The beliefs of phosphorus in soils largely depends on the detail characteristics of each soil, and besides microbial action, factors such as temperature, pH, and the degree of oxygenation all influence phosphorus mobility. In soils, inorganic phosphorus is typically associated with Al, Ca, or Fe, and each compound has unique solubility characteristics that decide the availability of phosphate to plants. The mobility and bioavailability of phosphate in soils are limited primarily by adsorption (the physical adherence or bonding of phosphate ions onto the surfaces of other molecules), and the rate of microbially mediated mineralization of organic forms of phosphorus. Mineralization is discussed in detail in the section titled ‘Microbially mediated processes’.

Figure 1.
Schematic diagram of the phosphorus cycle showing phosphorus reservoirs (living in green boxes; nonliving in grayness boxes), physical transport pathways (bluish arrows), and microbially mediated transformations (green arrows).

Marine sediments likewise represent an important phosphorus reservoir, but because the physical and chemic factors affecting marine sediment differ considerably from those on state, processes controlling phosphorus dynamics in marine sediments are somewhat different from that of soils. In marine sediment, phosphate tin can be present in insoluble inorganic phosphates minerals (such equally phosphorites), which are relatively immobile. Phosphate can likewise be sorbed onto iron or manganese oxyhydroxides. The sorbed phosphate can regain mobility in response to changes in the redox potential at the sediment–water interface and thus is considered more mobile. Every bit in terrestrial sediments, phosphorus in marine detrital organic affair tin also become remobilized as decomposition progresses through microbially mediated processes.

Biota (i.e., microbes, plants, and animals) serve equally another reservoir of phosphorus in the environs, equally they assimilate phosphorus within their cellular biomass. Biota can contribute significantly to environmental phosphorus levels; for example, microbial communities contribute 0.5–7.5% of total phosphorus in grassland and pasture topsoil, and upwardly to 26% in indigenous forests. Microbes are too responsible for generating the myriad of organic phosphorus compounds plant throughout the environment. In detail, microbes and primary producers play an important role in providing nutrition, including phosphorus, to higher trophic levels by making information technology biologically available (bioavailable). Phosphorus absorption is a microbially mediated procedure, which is discussed in the section titled ‘Transitory immobilization’.

Phosphorus is transported within the surroundings through various mass transfer pathways. For example, rivers are of import in the phosphorus bicycle as both reservoirs and transport pathways. Phosphorus that has weathered from minerals and has leached or eroded from soils enters rivers through a variety of vectors, including dissolved and particulate forms in water from overland flow and in groundwater, and particulates brought by current of air. Approximately 95% of phosphorus in rivers is particulate, and approximately 40% of that is spring within organic compounds. Rivers influence the distribution of phosphorus in soils and lakes by contributing or removing phosphorus, and riverine input is the single largest source of phosphorus to the oceans.

A number of outcomes are possible for phosphorus entering the ocean. Much of the riverine phosphorus flux is trapped in near-shore areas of the bounding main, such equally continental margins and estuaries, through immediate sedimentation and biological assimilation. The remaining phosphorus enters the dynamic surface ocean, also chosen the euphotic zone, in which virtually all bioavailable phosphorus is sequestered within biota through primary production. Upon expiry of the organisms, a fraction of the biologically sequestered phosphorus sinks below the euphotic zone and almost of it is regenerated into bioavailable forms like orthophosphate by heterotrophic organisms. This recycling is part of the so-called ‘microbial loop’. Physical processes such as upwelling and deep convective mixing draw the deep water, which, in most parts of the body of water, is nutrient-rich compared to the surface waters in the euphotic zone, where upward to 95% of it is reused in main production. The remainder is removed from the ocean reservoir through particulate sedimentation, mineral germination (which may exist microbially mediated), and scavenging by fe and manganese oxyhydroxides, all of which eolith phosphorus as a component of ocean sediment.

The phosphorus cycle differs from the cycles of other biologically of import elements, such every bit carbon, nitrogen, and sulfur, in that information technology lacks a significant gaseous component; virtually all phosphorus in the environment resides either in solid or in aqueous forms. The one exception to this rule is the volatile chemical compound phosphine (PH3, too called phosphane), a colorless, poisonous gas formed in the surround from the breakdown of alkali metal or alkali world metal phosphides with h2o. This procedure is poorly characterized and likely comprises various multistage chemical reactions. Microbially mediated phosphine production can exist a major source of the gas in engineered systems (east.chiliad., sewage treatment facilities and constructed wastewater treatment wetlands) where organic phosphorus is abundant and reducing conditions are common, suggesting that microbes could too play a function in phosphine formation in natural systems (although the direct enzymatic production of phosphine has non all the same been identified). Although phosphorus can exist equally phosphine, the gas does not persist in the environs owing to rapid autoxidation, precluding significant accumulation of phosphine in the atmosphere. Phosphine is therefore a pocket-sized component of the environmental phosphorus pool.

The absence of a significant gaseous stage does non eliminate the atmosphere as an important reservoir in the phosphorus bike. When weathering and erosion of soils generate inorganic and organic particulate phosphorus, current of air transports some of the particles from their source to a new location. These particles can include mineral dust, pollen and institute debris, insect fragments, and organic phosphorus bound to larger particles. This distribution of terrestrial particulate phosphorus, termed eolian degradation, plays an important role in delivering nutrients to the oceans. In oligotrophic ocean waters where nutrient levels are naturally low, such as in the open up ocean gyres where riverine inputs do not extend and pregnant upwelling does not occur, eolian deposition may contain a large portion of the food flux that is available for primary production. The eolian phosphorus flux to the oceans is approximately 1
1000 twelvemonth−1, of which approximately half is organic and the other one-half is inorganic. The solubility, and therefore bioavailability, of the phosphorus in eolian particulate matter differs significantly depending on its source; however, estimates suggest that approximately 15–l% is typically soluble.

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Phosphorus Cycle

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Encyclopedia of Bounding main Sciences, 2001


The global
phosphorus cycle
has four major components: (i) tectonic uplift and exposure of phosphorus-bearing rocks to the forces of weathering; (ii) concrete erosion and chemical weathering of rocks producing soils and providing dissolved and particulate phosphorus to rivers; (iii) riverine send of phosphorus to lakes and the ocean; and (four) sedimentation of phosphorus associated with organic and mineral matter and burial in sediments (

Effigy 1The cycle begins anew with uplift of sediments into the weathering government.

Effigy 1.
Cartoon illustrating the major reservoirs and fluxes of phosphorus described in the text and summarized in Tables
Table 1 and 2. The oceanic photic zone, idealized in the cartoon, is typically thinner in littoral environments owing to turbidity from continental terrigenous input, and deepens as the h2o cavalcade clarifies with distance away from the continental margins. The distribution of phosphorus amid different chemical/mineral forms in marine sediments is given in the pie diagrams, where the abbreviations used are: Porg,organic phosphorus; PAtomic number 26, iron-bound phosphorus; Pdetr, detrital apatite; Pauth, authigenic/biogenic apatite. The Porg, PFe, and Pauth
reservoirs represent potentially reactive phosphorus pools (run across text and
Tables 2 and 5
for discussion), whereas the Pdetr
pool reflects mainly detrital apatite weathered off the continents and passively deposited in marine sediments (notation that Pdetr
is non an important sedimentary phosphorus component in abyssal sediments, far from continents).Continental margin phosphorus speciation information were compiled from Louchouarn P, Lucotte M, Duchemin E and de Vernal A (1997) Early diagenetic processes in recent sediments of the Gulf of St-Lawrence: Phosphorus, carbon and iron burial rates.
Marine Geology
139(1/four): 181}200, and Ruttenberg KC and Berner RA (1993) Authigenic apatite germination and burial in sediments from not-upwelling continental margin environments.
Geochimica et Cosmochimica Acta
57: 991}1007. Abyssal sediment phosphorus speciation data were compiled from Filippelli GM and Delaney ML (1996) Phosphorus geochemistry of equatorial Pacific sediments.
Geochimica et Cosmochimica Acta
threescore: 1479}1495, and Ruttenberg KC (1990)
Diagenesis and burying of phosphorus in marine sediments: implications for the marine phosphorus budget. PhD thesis, Yale University. The global phosphorus cycle cartoon is from Ruttenberg (2000). The global phosphorus cycle. In:
The Encyclopedia of Global Change, Oxford University Printing. (in Press), with permission. The vertical water column distributions of phosphate typically observed in the three ocean basins are shown in the panel to the right of the global phosphorus cycle cartoon, and are from Sverdrup HV, Johnson MW and Fleming RH (1942)
The Oceans, Their Physics, Chemical science and General Biology. New York: Prentice Hall, (1942 Prentice Hall; used with permission.

Phosphorus is an essential nutrient for all life forms. It is a key player in fundamental biochemical reactions involving genetic material (Deoxyribonucleic acid, RNA) and energy transfer (adenosine triphosphate, ATP), and in structural support of organisms provided by membranes (phospholipids) and bone (the biomineral hydroxyapatite). Photosynthetic organisms utilize dissolved phosphorus, carbon, and other essential nutrients to build their tissues using energy from the sunday. Biological productivity is contingent upon the availability of phosphorus to these organisms, which constitute the base of the nutrient concatenation in both terrestrial and aquatic systems.

Phosphorus locked up in bedrock, soils, and sediments is non directly available to organisms. Conversion of unavailable forms to dissolved orthophosphate, which can be directly alloyed, occurs through geochemical and biochemical reactions at various stages in the global phosphorus cycle. Production of biomass fueled past phosphorus bioavailability results in the degradation of organic matter in soil and sediments, where information technology acts as a source of fuel and nutrients to microbial communities. Microbial activity in soils and sediments, in turn, strongly influences the concentration and chemic form of phosphorus incorporated into the geological record.

This article begins with a brief overview of the various components of the global phosphorus bicycle. Estimates of the mass of important phosphorus reservoirs, send rates (fluxes) betwixt reservoirs, and residence times are given in
Tables 1 and 2. As is clear from the big uncertainties associated with these estimates of reservoir size and flux, there remain many aspects of the global phosphorus wheel that are poorly understood. The 2nd one-half of the article describes electric current efforts underway to advance our understanding of the global phosphorus cycle. These include (i) the apply of phosphate oxygen isotopes (δ
18O-PO4) as a tool for identifying the part of microbes in the transformer of phosphate from one reservoir to another; (2) the use of naturally occurring cosmogenic isotopes of phosphorus (32P and
33P) to provide insight into phosphorus-cycling pathways in the surface ocean; (3) critical evaluation of the potential role of phosphate limitation in coastal and open up ocean ecosystems; (iv) reevaluation of the oceanic residence time of phosphorus; and (v) rethinking the global phosphorus-bike on geological timescales, with implications for atmospheric oxygen and phosphorus limitation primary productivity in the bounding main.

Table 1.
Major reservoirs active in the global phosphorus cycle and associated residence times

Tabular array 2.
Fluxes between the major phosphorus reservoirs

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Phosphorus Bicycle

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Encyclopedia of Environmental, 2008

Organic Cycles

Imposed on the inorganic cycle are 2 organic cycles which move phosphorus through living organisms as part of the food concatenation. These are a land-based
phosphorus cycle
which transfers it from soil to plants, to animals, and dorsum to soil again; and a water-based organic wheel which circulates information technology among the creatures living in rivers, lakes, and seas. The land-based bike takes a twelvemonth on average and the water-based wheel organic bicycle simply weeks. It is the amount of phosphorus in these two cycles that governs the biomass of living forms that land and sea can sustain.

The amount of phosphorus in the world’southward soils is roughly 90–200
MMT P according to various estimates. While the total phosphorus content of soils is big, merely a pocket-sized fraction is available to biota in most soils. This constitutes an available phosphorus pool containing 1805–3000 MMT P, almost likely 2000–2600 MMT P. A larger amount, in the range 27–840
MMT P, tin can be found in the oceans. The seawater contains 80–120
MMT P and the rest is accumulated in sediments.

The ocean water loses phosphorus continually in a steady drizzle of detritus to the lesser, where it builds up in the sediments every bit insoluble calcium phosphate. Despite the geological remobilization, there is a net annual loss of millions of tons of phosphate a year from the marine biosphere. Thus the ocean sediments are by far the largest stock in the biogeochemical cycles of phosphorus.

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Improving food management in agriculture to reduce eutrophication, acidification and climatic change

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Ecology Assessment and Management in the Nutrient Industry, 2010



Similarly to nitrogen, the
phosphorus bicycle
is now too profoundly influenced past human activities. Today the almanac production of phosphate is around 53 million tonnes P

(IFA, 2009), of which around eighty% is used every bit fertilisers and some other five% in mineral feed (Steen, 1998). Product and consumption of nutrient is thereby virtually exclusively responsible for the phosphorus trouble, not only leading to emissions of nutrifying pollutants but also to depletion of a non-renewable resources.

Phosphorus utilisation (i.e. uptake in meat, milk and eggs in relation to input with feed) in beast product varies between different livestock groups but is normally in the range of 15–forty% (Damgaard Poulsen and Holton Rubaek, 2005). Consequently, the ascendant part of phosphorous in the feed ends upward in the manure, and therefore the ongoing trend on organising the globe’southward pork and poultry production in landless production systems poses a long-term problem also for the phosphorous wheel. In areas close to concentrated livestock operations, there volition be increasing soil P aggregating whereas the feed crops that are geographically separated for the creature farms must be fertilised with ‘new’ phosphorous from synthetic fertilisers. Nowadays P distribution on global farmland must exist improved and there are two of import arguments for that today: the eutrophication problem and resources depletion. Such an improvement must get-go with a disquisitional review of the effects of the present structure that is now developed in the globe’s livestock production system.

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Phosphorus in Soils—Biological Interactions☆

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Reference Module in Globe Systems and Environmental Sciences, 2019

The Phosphorus Bike

In some ways, the
phosphorus bicycle
is not as complex as the nitrogen or sulfur cycles because P does not typically undergo oxidation-reduction reactions or exist in gaseous forms. Yet, as in the N and S cycles, the biological processes of mineralization and immobilization besides as found uptake are present. Additionally, the microbial fraction is active in the solubilization of relatively insoluble inorganic P materials, making P available for establish or microbial uptake or other transformations.

Fig. 3
shows the P cycle, illustrating the various compartments of P in the soil environs. The P cycle is divided into a geochemical subcycle and a biological subcycle, with the inorganic solution P pool serving every bit the central point in the overall bicycle. This puddle of inorganic solution P serves as the source of orthophosphate for plants and soil microorganisms. The remainder of this article will deal with the biological subcycle of P.

Fig. 3

Fig. 3.
The phosphorus wheel showing inputs, losses, and transformations. This model illustrates the relationship of the chemical and biological P subcycles through the solution orthophosphate puddle.

Adapted with permission from Walbridge, M.R. (1991). Phosphorus availability in acid organic soils of the lower North Carolina coastal plainly.
72, 2083–2100.

In the biological subcycle, soluble orthophosphate is either taken up by plants or immobilized into the microbial biomass. When organic sources of P, such every bit crop residues, manures, and municipal biosolids, are land-applied, iii possible fates of the P tin can occur. The organic P, specially inositol phosphates, may be incorporated straight into stable humus, becoming relatively unavailable for plant and microbial use. If the balance has a relatively high P content, relative to C and N, a portion of the P will exist mineralized every bit orthophosphate during microbial decomposition of the residual. Finally, part of the P in the residue will be incorporated into the microbial biomass during decomposition. This immobilized P may go available as soluble P when the microbial biomass fraction dies back and is itself mineralized by other microorganisms.

While this model of the P wheel segregates biological activity from geochemical action, it should be noted that the dissolution of apatite and other secondary P minerals can exist enhanced by the activities of microorganisms and plants. Dissolution or solubilization of these minerals releases orthophosphate to solution P, where it tin then exist utilized by plants or microorganisms. Of course, a portion of this P will over again precipitate equally secondary P minerals, or be adsorbed past Al- and Fe-oxides and clay surfaces.

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Treatise on Geochemistry, 2003


Historical perspective: the marine phosphorus budget

The current vision of the marine
phosphorus bicycle
differs substantially from that which prevailed as of the early 1980s. These changes accept been driven past methodological developments, which have made new observations possible, also as past challenges fabricated to accepted paradigms every bit new studies have worked to reconcile new information with old, and sometimes entrenched, views.

The beginning comprehensive global marine phosphorus budget took the approach of quantifying phosphorus removal from the body of water by characterizing P burial rates in different depositional environments (Froelich
et al., 1982
). These researchers took the important approach of separately quantifying P-burying rates for different depositional environments, and different sediment types, recognizing that the processes dominating in unlike environments would be distinct, and therefore subject to dissimilar decision-making factors. This early incarnation of the marine phosphorus budget focused nearly exclusively on the deep sea; the only sea margin data included were from areas characterized by upwelling apportionment and phosphorite germination, equally the latter were recognized as hot-spots for phosphorus burial. Notably, upwelling margins represent only 2% of the total marginal area of the bounding main (Ganeshram
et al., 2002
Berner, 1982).

The emphasis on the pelagic realm was the effect of the prevailing bias at the time, in both research and funding, toward blueish water oceanography, whereas continental margins received far less research attending. The pelagic budget described by
et al.

was viewed equally cocky-consistent, because the P-removal flux with sediments was balanced by the dissolved phosphate input flux with rivers, with a residence fourth dimension of 80
kyr, similar to the approved value of 100
kyr, the accustomed residence time estimate at that time (Broecker and Peng, 1982).

Adopting the budgetary model of
et al.
Ruttenberg (1993)
presented a revised budget, this time taking into account the importance of continental margins,
in toto, in the global marine P-cycle. Inclusion of the margins was motivated by a growing recognition of continental margins as extremely important depocenters for organic affair (Berner, 1982), and the cognition that organic matter is one of the most, if not the nigh important vector for delivery of phosphorus to the seabed. An additional motivator for inclusion of continental margins was the recognition that the early diagenetic regime in organic-matter-rich margin sediments of all types, and not exclusively those underlying upwelling regimes, make these depositional environments likely places for authigenic carbonate fluorapatite (CFA) formation.

Identification of CFA in nonupwelling environments required the utilise of new, and indirect methods of detection, considering dilution of this authigenic phase by the big brunt of terrigenous material in many continental margin settings makes identification by direct methods, such as XRD, impossible. Ruttenberg (1992) adjusted existing methods for sequential extraction of phosphorus and trace metals from a variety of disciplines, including soil science, limnology, and marine geochemistry, resulting in the SEDEX method, which is able to separately quantify CFA. Application of a coupled SEDEX–pore water approach to continental margin sediments revealed that formation of authigenic CFA is not restricted to margin environments characterized past upwelling (Ruttenberg and Berner, 1993, and others, see
Section 8.13.3.iii.2). Inclusion of continental margins in the global marine phosphorus budget increased the P-removal flux past 2–6 times (depending upon which burial flux estimates are used), due to high rates of burial in body of water margin sediments of organic phosphorus, authigenic CFA, and iron-bound phosphorus, the latter particularly important in deltaic marginal environments (Table 3;
Ruttenberg, 1993).

More recent studies take further refined the estimated burial fluxes of phosphorus in the global marine phosphorus budget (Table 4)
, including meliorate estimates of P-removal with iron oxyhydroxides at MORs (Wheat
et al., 1996
), inclusion of burial fluxes for authigenic rare globe element- and thorium-phosphates (Rasmussen, 2000), phosphates buried in clan with hydroxyapatite from fish bones, scales, and teeth (using a modified SEDEX method to separately quantify hydroxyapatite as distinct from CFA:
et al., 2000
); and taking into account the return benthic flux of phosphate out of sediments (Colman and Holland, 2000). These and other studies (Compton
et al., 2000
Filippelli and Delaney, 1996) concur with
Ruttenberg (1993)
that the before pelagic-focused budget of
et al.

underestimated global ocean P-burying fluxes, and therefore overestimated P-residence times (Tabular array iii).

Table 4.
Geochemical sectionalization of reactive-P burying fluxes

Phosphorus reservoir Phosphorus burial flux (10x
mol year−1)
Method of determination
Organic-P Delaney, 1998
(calculated from Corg
burying charge per unit and Porg/Corg
1.v Froelich
et al., 1982

(calculated from Corg
burial rate and Porg/Corg
1.6 Froelich, 1984 (calculated from Corg
burial rate and Porg/Corg
2.0 Mach
et al., 1987

(calculated from Corg
burial rate and Porg/Corg
4.1 Ruttenberg, 1993
(measured Porg
via SEDEX method and estimated sediment delivery flux to the oceans)
Variability in first four estimates results from unlike Corg
burying fluxes and different Porg/Corg
ratios chosen for estimated Porg
burial fluxes (see
Delaney, 1998
for summary).
Authigenic CFA, Biogenic HAP, 0.4 Froelich
et al., 1982
(based on diagenetic modeling of sediment pore h2o)
CaCOthree-P 8.0 Filippelli and Delaney, 1996
(assuming 80–90% of Ptotal
measured via SEDEX method is PCFA
combined with Ptotal
accumulation rates)
nine.1 (2.2) Ruttenberg, 1993
(measured PCFA
via SEDEX method and estimated sediment delivery flux to the oceans; value in parentheses is minimum guess, see comments)
et al.
estimates are for upwelling, classical phosphogenic provinces only.
Ruttenberg’s (1993)
maximum estimate assumes all phosphorus measured in pace Iii of the SEDEX method is truly authigenic. The minimum estimate assumes only the portion observed to increment above the concentration in the shallowest sediment interval is truly authigenic, accounting for the possibility that there may be a nonauthigenic background component to this reservoir (run across
Ruttenberg and Berner (1993)
for an expanded discussion). HAP=hydroxyapatite (fish basic, teeth, scales).
Ferric atomic number 26-bound P:
Hydrothermal MOR Processes:
High-temperature ridge centrality
0.01 Wheat
et al.
Low-temperature ridge flank
0.65 Wheat
et al.

basalt seawater reactions during convective circulation of seawater in sediments and crust of flanks of MORs
Hydrothermal plumage scavenging
0.77 Wheat
et al.
; Feeley
et al.
Total hydrothermal
Non-hydrothermal scavenging onto Atomic number 26-oxyhydroxides: 1.5 Froelich
et al., 1982

(in the
et al.

written report, this quantity was attributed to burial with CaCO3, determined past dissolving foram and coccolith tests from deep-body of water cores. The work of Sherwood
et al.
(1987) and Palmer (1985) demonstrated conclusively that phosphorus associated with CaCO3
tests in the deep bounding main is nearly all associated with Fe-oxyhydroxide coatings on the tests. This quantity is therefore more accurately attributable to phosphorus burial with reactive Fe-oxyhydroxide phases (see also
Ruttenberg, 1993)
iv.0 (0.4) Ruttenberg, 1993
(measured PFe
via SEDEX method and estimated sediment delivery flux to the oceans; value in parentheses is minimum judge, see comments)
Ruttenberg’s (1993)
maximum estimate assumes all phosphorus measured in stride 2 of the SEDEX method is truly reactive-P. The bulk of the phosphorus reported in
Ruttenberg (1993)
was observed in deltaic sediments, however, and the possibility exists that some portion of this reservoir is detrital, and therefore does not represent phosphorus removed from seawater. Thus, the minimum estimate assumes that all of the deltaic PAtomic number 26
is detrital (this is an extreme, and probable unrealistic minimum end-fellow member)
Loosely sorbed P 1.3 Ruttenberg, 1993
(measured Pexchg
via SEDEX method and estimated sediment delivery flux to the oceans)
Calcium carbonate <0.009 Delaney, 1998
(calculated from Holocene CaCO3
burial flux and maximum foraminiferal CaCO3-P content from Palmer, 1985)
Any phosphorus associated with CaCO3
tests, not surface metal oxyhydroxide coatings, is included in the quantity measured by pace 3 of the SEDEX method; this estimate serves every bit a measure out of the fraction of the step III, or the burial flux of authigenic apatite+biogenic apatite+CaCO3, that can be accounted for by burial of CaCO3.
REE phosphates and aluminum-phosphates 6.56 Rasmussen, 2000
(determined for ancient marine sandstones via microscopy, where sedimentary textures are the master evidence use to argue that they are authigenic, and therefore representative of a reactive-P burying flux.
These phases have withal to be observed in modern marine sediments, and their role equally reactive phosphorus sinks in the ocean has yet to be verified. Verification of authigenic REE–P and Al–P formation in the modern bounding main would strengthen arguments for the authigenic nature of these phases in ancient sediments.

Ruttenberg (1993),
Delaney (1998), and
et al.

The college P-burying charge per unit estimates that these various groups have converged upon prepare upwards an imbalance in the global marine phosphorus budget when contrasted with the riverine dissolved P-input charge per unit. This imbalance can be reconciled if some fraction of the phosphorus associated with riverine particulate matter is solubilized upon entry into the bounding main (Table two). Estimates of the quantity of phosphorus that might be liberated upon commitment from rivers to the oceans, or “releasable-P,” have been made in several studies (Ruttenberg, 1993;
Colman and Kingdom of the netherlands, 2000;
Ruttenberg and Canfield, 1994;
Berner and Rao, 1994;
et al., 2000
et al., 1995
Ramirez and Rose, 1992;
Froelich, 1988). Estimates made on the ground of P-inputs that include this “releasable” riverine particulate-P yield residence times that autumn inside the range of residence fourth dimension estimates derived from P-burial fluxes (Table 3). Despite the big uncertainties associated with these numbers, equally evidenced by the maximum and minimum values derived from both input and removal fluxes (Table three), these updated residence times are all significantly shorter than the canonical value of 100
kyr. Residence times on the order of 10–17
kyr make feasible a office for phosphorus in perturbations of the body of water–atmosphere CO2
reservoir on the timescale of glacial–interglacial climate change.

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Food depletion and pesticide use

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Ecophysiology of Pesticides, 2019


Urease and phosphatase enzymes

Hydrolases are of particular importance on account of their role in the soil nitrogen, phosphorus, carbon, and sulfur cycles (Megharaj et al., 1999). Urease is an enzyme that catalyzes the hydrolysis of urea into CO2
and NH3
and is a cardinal component in the nitrogen cycle in soils. Urease activity is found in a large number of soil bacteria and fungi (Sarathchandra et al., 1984). Phosphatase is an exocellular enzyme produced by many soil microorganisms and is responsible for the hydrolysis of organic P compounds to inorganic P (Monkiedje et al., 2002). Several researchers accept shown unchanged, increase, or decrease in urease activity following diverse pesticide applications (Antonious, 2003; Chen et al., 2001b; Ingram et al., 2005; Nowak et al., 2004). The urease activity in soil is often correlated with the size of the microbial population and activeness (Roberge and Knowles, 1967). In soil, hydrolysis of urea, an important fertilizer, with urease yields NH4
ion that is taken up past plants. Nitrification of NH4
ion into NO
ion in soil is rapid and



tin can leach downwardly with water. Decreased urease activeness in soil with the application of pesticides reduces urea hydrolysis that is generally beneficial because it helps to maintain N in a form (NH4
+) less leachable (Antonious, 2003).
Yang et al. (2006)
showed that chlorimuron-ethyl– and furadan-activated urease in the four soils. The chlorimuron-ethyl– and furadan-enhanced urease activity upwards to xiv%–xviii% and thirteen%–21%, respectively. Contrarily, acetamiprid reduced up to 35% urease activity in soil at 43 days after crop sowing (Singh and Kumar, 2008). Similar observations have been reported in case of phosphatase activity in soil (Demanou et al., 2004; Madhuri and Rangaswamy, 2002; Monkiedje et al., 2002; Rangaswamy and Venkateswarlu, 1996). Acid and alkaline phosphatases are mostly found in microorganisms and animals (Tabatabai, 1982, 1994).
Demanou et al. (2004)
did not find pregnant effect of Ridomil fungicide on phosphatases activities in soil. These enzymes may be protected from degradation by adsorption to clays or to humic substances in soil (Boyd and Mortland, 1990). This protection of these exoenzymes may upshot their insensitivity toward the fungicide application (Demanou et al., 2004). Similarly,
Yao, et al. (2006)
revealed that the activity of dehydrogenase was increased after acetamiprid application for two weeks. They demonstrated that the enzyme activities in samples treated with 0.5, 5, and 50
mg kg−i
dry soil were nigh 2.five-, ane.5-, and 2-fold to that of the control at 28 days of application. Contrary to these findings,
Klose, et al. (2006)
reported that soil fumigation reduced the activeness of acid phosphatase to 22% over a menstruum of xc days. However, decrease in this enzyme may
exist ascribed to the suppression of a sensitive fraction of soil biota. b-Glucosidase, cellulase, invertase, and other enzymes are besides very important enzymes involved in the transformation/decomposition of organic matter in soil. b-Glucosidase catalyzes the hydrolysis of disaccharides in soil to class b-glucose. The hydrolysis products of b-glucosidases are believed to be of import energy sources for microorganisms (Tabatabai, 1994). Cellulase catalyzes hydrolysis of cellulose to
d-glucose. Cellulose is the near abundant polysaccharide of constitute cell walls and represents a significant input to soils (Richards, 1987). Invertase hydrolyzes sucrose to fructose and glucose. Invertase is ubiquitous enzyme that occurs in plant tissues and soil organisms (Skujins, 1976). Other enzymes such equally NR, aryl sulfatase, amylase, xylanase, catalase, and protease also play an of import office in biochemical reactions and nutrient cycling. Application of pesticides increased, decreased, or did non bear on activities of these enzymes in soils, depending on the nature and concentrations of pesticides used, incubation period, status of enzymes in soil, and soil atmospheric condition.
Ismail et al. (1999)
studied the effects of methamidophos on cellulolytic activeness in three soils (loamy sand, clay loam, and clay) and found a correlation between cellulase activity and clay content of the soils. No effect of methamidophos was observed on cellulase activeness in dirt soil, whereas 25% and 38% reduction in cellulase activity was observed in dirt loam and loam sand, respectively, after 6 weeks. It is usually reported that high enzymatic activities are associated with loftier organic matter contents, but different results have been reported by
Gianfreda et al. (1995), where low invertase concentration was recorded in soil containing loftier organic affair contents. They also observed greater invertase activity with college dirt content in soil. Recently,
Gundi, et al. (2007)
investigated the furnishings of the insecticides including monocrotophos and quinalphos (organophosphates), and cypermethrin (pyrethroid), on soil enzyme activities in ii agricultural soils—blackness vertisol soil and red alfisol soil for 30 days under laboratory conditions.

Individually applied three insecticides at 5, 10, and 25
mg g−1
to the soil markedly increased the activities of cellulase and amylase. Interestingly, combinations involving monocrotophos or quinalphos with cypermethrin demonstrated synergistic and combative effects on both enzymes in the soils. At low concentrations (5 and 10
mg g−i), insecticides in combination demonstrated synergistic furnishings while at higher concentrations (25
mg yard−1) demonstrated an antagonistic interaction effects on these enzymes. They reported that such diverse furnishings of insecticides on two enzyme activities
was in concomitant to populations of cellulolytic and amylolytic microbes in soils treated with insecticides. Similarly, in another study,
Klose, et al. (2006)
reported that soil fumigation reduced the action of aryl sulfatase (62%) and b-glucosidase (half-dozen%), implying that S-mineralization in soils and the full oxidative potential of microorganisms were more affected by fumigation. They also indicated that soil fumigation could alter microbial communities and important biochemical reactions involved in cycling of elements.

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Ecology Bear upon, Concept and Measurement of

Ellen W.
James R.
, in

Encyclopedia of Biodiversity, 2001


Altered Biogeochemical Cycles

All the substances found in living things, such as water, carbon, nitrogen, phosphorus, and sulfur, cycle through ecosystems in biogeochemical cycles. Human activities modify or accept the potential to alter all these cycles. Sometimes the results stalk from changing the amount or the precise chemistry of the cycled substance; in other cases, humans change biogeochemical cycles by changing the biota itself.

Freshwater use, dams, and other engineering feats alter the amount and charge per unit of river catamenia to the oceans and increase evaporation rates, directly affecting the water bike and indirectly impoverishing aquatic life. Direct homo modifications of living systems also perturb the water cycle. In the tall “deject forests” of redwoods in northern California or Douglas firs in the Pacific Northwest, the copse gather more than moisture direct from the clouds than falls as rain. Among the furnishings of logging in such forests is a depletion of stream flows, which shifts the h2o cycle. In South Africa, European settlers supplemented the treeless native scrub, or fynbos, with such trees as pines and Australian acacias from like Mediterranean climates. Because these copse are larger and thirstier than the native scrub, regional water tables have fallen sharply.

Human being action has contradistinct the global nitrogen cycle past greatly increasing the amount of nitrogen stock-still from the atmosphere (combined into compounds usable by living things). The increase comes mostly from deliberate addition of nitrogen to soils every bit fertilizer simply also equally a by-product of the burning of fossil fuels. Agriculture, livestock raising, and individual grand maintenance chronically add tons of excess nutrients, including nitrogen and phosphorus, to soils and water. The additions are often invisible; their biological impacts are ofttimes dramatic. Increased nutrients in littoral waters, for example, trigger blooms of toxic dinoflagellates, the algae that crusade cherry-red tides, fish kills, and tumors and other diseases in varied body of water creatures. When huge blooms of algae dice, they fall to the seafloor, where their decomposition so robs the water of oxygen that fish and other marine organisms tin can no longer live there. With nitrogen concentrations in the Mississippi River two to three times as loftier as they were fifty years ago, a gigantic expressionless zone forms in the Gulf of Mexico every summer; it covered a record 20,020 square kilometers in the summertime of 1999.

The burning of fossil fuels is radically altering the carbon bike, primarily by greatly increasing the atmospheric concentration of carbon dioxide. With other greenhouse gases, such as methane and oxides of nitrogen, carbon dioxide helps keep the earth’s surface at a livable temperature and drives establish photosynthesis, just since the industrial revolution, atmospheric carbon dioxide concentrations have risen 30% and are now widely thought to be disrupting the planet’s climate.

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URL: discipline/commodity/pii/B0122268652001073

Thing and Thing Flows in the Biosphere

, in

Encyclopedia of Ecology, 2008

Food Webs

Although biological cycling of each biogenic element is characterized by its ain properties (run into
Carbon Cycle,
Oxygen Cycle,
Nitrogen Cycle,

Phosphorus Cycle
Calcium Cycle, and
Sulfur Cycle), all of the elements include migration of biomass in nutrient webs. Transfer of the matter in the course of the cycling involves the following main steps: absorption and accumulation by living organisms of elements from abiotic environs; distribution of the matter among organisms as a result of herbivory, predation, and parasitism; territorial migration of organisms; formation of dead organic affair (DOM or mortmass) as a event of excretion and death of organisms; decomposition of the mortmass and return of the elements to the abiotic environment.

In accordance with the identify, occupied by species in the food webs, they are usually divided into three main groups: producers (which utilize external energy, solar or inorganic chemical, and realize biosynthesis: generate organic matter), consumers (which apply chemical free energy of living tissue of other organisms), and reducers (which use chemic energy of mortmass and exercise its biodegradation: decomposition to simple inorganic agents). A nomenclature of such organisms was initially proposed past A. L. Lavoisier in 1792 so, in another form, was developed by West. Pfeffer in 1886.

The chief players in the nutrient cycling are producers and reducers. The former are an ‘engine’ of the cycling; they involve elements from the abiotic surround in the turnover and ship them further in the composition of permanently generating high-energetic organic affair.

The reducers ‘close’ the cycling; they render the elements to the abiotic environment, where they can exist used by producers once again. Abiotic decomposition takes identify, just its intensity is very depression. Without producers available elements would concentrate in the biomass and exit the surround; cycling would stop, and life development would end. The simplest bogus stable ecosystems, functioning in closed flasks, included populations of producers (unicellular algae) and reducers (bacteria and fungus).

Theoretically it is possible to envision producers, independently realizing the function of reducing with respect to their ain biomass. Just in reality this possibility is not realized. This fact can exist explained by the absence of evolutionary reasons of forming ‘self-sufficient’ organisms, if the hypothesis near the origin of heterotrophs (consumers and reducers) earlier producers is correct. Stable biological cycles could form gradually, during the procedure of co-adaptation of producers and reducers. Shut symbiosis and species peculiarity are typical for relations between producers and reducers.

The most common flows of matter in the biosphere, including food chains and abiotic topical means, are presented in

Figure 2
. There are four ecosystems of different nature in the

Figure 2
: terrestrial; shelf; open sea; abyssal black geyser. The core wheel for each ecosystem is cycling i: ‘producers–mortmass–reducers–inorganic salts–producers’. For the terrestrial ecosystem it tin be written: ‘producers–litter–reducers–soil–producers’; for the h2o 1: ‘producers–sediments–reducers–water–producers’. The cycles are initiated by the producer cake, which transform the solar energy to the chemic one in the class of the photosynthetic process. It is the well-nigh rapid and mainframe cycle in the biosphere; its dynamic properties are considered beneath.

Figure two.
Main matter flows in the biosphere.

The amount of different biogenic elements in cycling is determined mainly by features of the environment. The intensity of cycling depends on properties of the producer block. The additional loop 2 reflects the role of the consumer block. The energy and thing flow through information technology are approximately 10 times less than directly from the producers to the mortmass, just the consumers, influencing the producer block, ‘bootstrap’ the turnover. Similarly, loop 3, involving the block of consumers of 2d social club and also 10 times less intensive than loop two, contributes to increasing intensity of the full cycling. In

Figure 2

for the marine ecosystems the blocks of producers, reducers, and consumers are marked by the showtime letters P, R, and C, respectively.

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Sustainable diets for a nutrient-secure time to come

Claire Fitch
, in

Ecology Diet, 2019

Nutrient cycles

The rapid increase in the utilize of fertilizers has considerably increased the food supply over the past century (Bouwman et al., 2013
), but concurrently significantly modified nitrogen and
phosphorus cycles
in ways that can limit future production potential (

Steffen et al., 2015;
Erisman et al., 2013). Nitrate and phosphate runoff and groundwater contagion from synthetic fertilizers, brute manure, and discharges from some aquaculture systems cause toxic algal blooms that deplete oxygen levels in the h2o and kill fish, plants, and other aquatic life (Mallin and Cahoon, 2003;
Islam, 2005). Such eutrophication not only harms aquatic ecosystems and humans reliant on those water supplies, merely it too impairs fish farming in the region (Fry et al., 2016). Meanwhile, phosphorus in fertilizer is derived from mining global phosphate rock sources, which are vulnerable to geopolitical conflicts and expected to be depleted by the end of the century, threatening long-term food product capacities (Cordell et al., 2009).

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URL: discipline/article/pii/B9780128116609000163

The Phosphorus Cycle Differs From the Biogeochemical Cycles in That


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