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Home , Ecosystem, Ecosystem respiration

Ecosystems (2006) 9: 1–4 DOI: 10.1007/s10021-005-0036-3

COMMENTARY

Is Net Production Equal to Ecosystem Accumulation?

Gary M. Lovett,* Jonathan J. Cole, and Michael L. Pace

Institute of Ecosystem Studies, Millbrook, New York 12545, USA

ABSTRACT Net ecosystem production (NEP), defined as the conceptual coherence between NEP and net pri- difference between gross and mary production and that it is congruous with the total , represents the total widely accepted definitions of ecosystem autotro- amount of organic carbon in an ecosystem available phy and heterotrophy. Careful evaluation of NEP for storage, export as organic carbon, or nonbio- highlights the various potential fates of nonrespired logical oxidation to through fire or carbon in an ecosystem. ultraviolet oxidation. In some of the recent litera- ture, especially that on terrestrial , NEP Key words: net ecosystem production; carbon has been redefined as the rate of organic carbon accumulation; net primary production; gross pri- accumulation in the . Here we argue that mary production; total ecosystem respiration. retaining the original definition maintains the

INTRODUCTION Net ecosystem production (NEP) is a fundamental NEP for various ecosystems has increased in recent property of ecosystems. It was originally defined by years as ecosystem scientists investigate more fully Woodwell and Whittaker (1968) as the difference the controls on the C balance of the . between the amount of organic carbon (C) fixed by However, studies often esti- in an ecosystem (gross primary mate NEP as the rate of C accumulation, (for production, or GPP) and total ecosystem respiration example; Lichter 1998; Caspersen and others 2000; Re (the sum of autotrophic and heterotrophic res- Wirth and others 2002), and a recent paper and piration). Defined this way, NEP represents organic even a widely read textbook explicitly equate NEP C available for storage within the system or loss with C accumulation rate (Randerson and others from it by export or nonbiological oxidation. The 2002; Chapin and others 2002). The goal of this sign of NEP defines whether an ecosystem is essay is to explain why, in our view, this is incor- autotrophic (NEP greater than zero, as in a typical rect. or grassland) or heterotrophic (NEP less than NEP can best be understood within the context of zero, as in and many lakes and rivers). The a complete organic C balance for an ecosystem, original definition of NEP as the difference between which can be written as: GPP and R is conceptually parallel to the definition e DC ¼ GPP þ I À Re À E À Ox ð1Þ of net primary production (NPP), which is the dif- org nb ference between GPP and autotrophic respiration

(Woodwell and Whittaker 1968). The reporting of where GPP and Re are as defined above, DCorg is the change in organic C storage in the ecosystem, I

Received 18 March 2005; accepted 11 May 2005; published online j. is the import of organic C, E is the export of organic *Corresponding author; e-mail: [email protected] C, and Oxnb is nonbiological oxidation of C, for

1 2 G. M. Lovett and others

Figure 1. Fates of organic carbon (C) fixed in or imported into an ecosystem. Total ecosystem respiration (Re) is the sum of autotrophic respiration (Ra) and heterotrophic respiration (Rh). The shaded area contains the components of the NEP of the system. ‘‘Accumulation in ’’ represents all biomass (, , or microbial); the arrow is drawn from NPP in this diagram because plant biomass accumulation is generally the largest biomass term. NPP, net primary production; NEP, net ecosystem production; GPP, gross primary production; CO2, carbon dioxide; UV, ultraviolet.

instance by fire or ultraviolet (UV) oxidation. This Equation 4 and Figure 1 illustrate the potential equation requires one to specify the boundaries of fates of NEP, plus any organic C imported into the the ecosystem and a specific interval over system, as either (a) storage in the ecosystem, for which the change in C storage is evaluated. The instance as an increment of the organic C pool in terms in Eq. 1 are usually expressed as mass of C , , or sediments; or (b) export from per unit area for that specified time interval, or the ecosystem, for instance as dissolved organic C, sometimes as instantaneous rates (i.e., mass area)1 particulate C, or as harvest removal of organic time)1). material; or (c) nonbiological oxidation by fire or If NEP is defined as: UV oxidation. Thus, NEP represents the C poten- tially available for storage within a system, but not NEP ¼ GPP À R ð2Þ e all of the NEP is necessarily stored. Similarly, some C that is stored within an ecosystem may have been then imported into the ecosystem rather than fixed there, and would not contribute to the system’s D 3 NEP ¼ Corg þ E þ Oxnb À I ð Þ NEP. A few simple examples will help to clarify these and points. First, consider a forest ecosystem, with the lower bound of the ecosystem set at the bottom of NEP þ I ¼ DCorg þ E þ Oxnb ð4Þ the rooting zone. If some of the C that is fixed by photosynthesis in the forest leaches below the In aquatic ecosystems, NEP is usually measured rooting zone as (DOC), directly from the mass balance of (O2)or that C is not stored in the ecosystem but represents carbon dioxide (CO2) (for example, Cole and others part of the NEP because it is fixed by the 2000; Hanson and others 2003). Total respiration is but not respired within the ecosystem. Likewise, if obtained from the gas balance at (when GPP this forest grows for 50 years and then is harvested, is zero) and then GPP is calculated from Eq. 2. This with some of the woody material removed from the gas balance approach is also the basis for estimating ecosystem, the C in the harvested wood should be NEP in some terrestrial using eddy covari- included in the calculation of NEP even though it ance techniques, or globally using large-scale was exported from the system. This is exactly par- atmospheric data with some additional isotopic allel to the fact that herbivory and detrital losses are measurements (for example, Luz and others 1999). included in the calculation of NPP. In terrestrial studies, however, NEP is often calcu- A somewhat more problematic issue is the cal- lated by summing the fates of NEP in the ecosys- culation of NEP if the forest were burned rather tem, usually just using the DCorg term and ignoring than harvested. Some of the C lost during the fire the other terms on the right-hand side of Eq. 3. would be exported from the system as organic C in Net Ecosystem Production and Carbon Storage 3

soot and smoke, and some of it would be nonbio- become a very large sink for atmospheric CO2 in logically oxidized to CO2. Both of these losses the industrial era, storing some 2 Gt C/y at present. should be included if NEP is calculated by summing This storage is almost entirely due to the diffusion its potential fates, as in Eq. 3. For example, if a of atmospheric CO2 into water that in the 2 forest accumulated 100 g C/m /y over 50 years, preindustrial era, when CO2 levels were lower, had and in year 50 a fire consumed all of the accumu- been in equilibrium with the atmosphere. lated 5,000 g C/m2, the NEP over the 50-year is a very minor factor in this flux (for example, period would still be 5,000 g C/m2 even though Siegenthaler and Sarmiento 1993; Broecker and there would be no C accumulation in the system. others 1979). In this case, the fate of the NEP is in the export and Randerson and others (2002) suggest that the nonbiological oxidation terms of Eq. 3. definition of NEP be changed to equate NEP with Consider also a surrounded by a the rate of C accumulation in the ecosystem. They forest. If from the forest blow into the lake, argue that we need a term for C accumulation, and then sink to the bottom of the lake and accumulate that the term ‘‘NEP’’ should be appropriated for in the sediments, does this C storage increase the that use. We maintain that NEP has a different NEP of the lake? The answer is no, because the meaning, and that the change in the organic C pool leaves contribute equally to the import (I) and C per unit time should simply be called the organic accumulation (DCorg) terms in Eq. 3, so the NEP is carbon accumulation rate. It would be positive for not affected. If the leaves were partially decom- an increment in C and negative for a decrement. posed before storage in the sediments, the CO2 loss Net ecosystem production is a useful term for a would increase ecosystem respiration and thus concept that is different from C accumulation – it is cause the lake’s NEP to be more negative. This type the amount of organic C fixed in an ecosystem that of C subsidy from terrestrial ecosystems is the rea- is not respired there and is therefore available for son that lakes can simultaneously have positive C accumulation, export, or nonbiological oxidation. accumulation and negative NEP. Net ecosystem production may be a good approxi- Measuring NEP can be challenging, especially in mation of the organic C accumulation rate within terrestrial ecosystems. Estimates of terrestrial eco- the system if inputs and outputs of organic C are system production and respiration (if NEP is cal- negligible, but it is incorrect to assume that NEP culated from Eq. 2) and of and vegetation C and organic C accumulation are always equivalent. accumulation (if NEP is calculated from Eq. 3) Many studies in the literature have used NEP and typically have high uncertainities. Some studies organic C accumulation interchangeably, with the compare both methods of calculation (for example, implicit assumption that import, export, and non- Hamilton and others 2002). In recent years, eddy biological oxidation of organic C are negligible. covariance techniques have been used frequently However, in many cases, these terms cannot be to estimate NEP. By measuring the vertical net flux ignored. In fact, at continental to global spatial of CO2 above the ecosystem, eddy covariance scales, the major fate of terrestrial NEP is export as measurements estimate the total CO2 exchange, riverine DOC (Schlesinger 1997). often called the ‘‘net ecosystem exchange’’ (NEE). Schulze and others (2000) proposed the term Net ecosystem exchange is equal to the NEP plus ‘‘net production’’ to integrate the terrestrial sources and sinks for CO2 that do not involve C balance over larger spatial and temporal scales, conversion to or from organic C (although the NEE but we agree with Randerson and others (2002) is by convention opposite in sign, so that fluxes into view that net biome production is simply organic the ecosystem are negative): C accumulation at a larger scale and that it would be an unnecessary addition to an already confus- ÀNEE = NEP + inorganic sinks for CO 2 ð5Þ ing list of terms. À inorganic sources of CO2 We recommend retaining the original definition of NEP as GPP minus Re because (a) it represents a useful concept, (b) it is congruous with the ac- Examples of inorganic sources and sinks are cepted definition of autotrophic and heterotrophic reactions, precipitation or dissolution of systems, and (c) its similarity to NPP provides a carbonates, and atmosphere–water equilibrations. cohesive set of conceptual constructs for ecosystem Although they are likely to be minor terms in the . Net ecosystem production should not be CO exchange of a forest (but see Raymond and 2 equated with organic C accumulation in theory or Cole 2003), these inorganic sources and sinks can in practice unless the other terms in Eq. 3 can be be very important in the . The ocean has shown to be negligible. 4 G. M. Lovett and others

REFERENCES Lichter J. 1998. and forest development on coastal Lake Michigan sand dunes. Ecol Monogr 68:487–510. Broecker WS, Takahashi T, Simpson HJ, Peng TH. 1979. Fate of Luz B, Barkan E, Bender ML, Thiemens MH, Boering KA. 1999. carbon dioxide and the global carbon budget. Sci- Triple-isotope composition of atmospheric oxygen as a tracer ence 206:409–18. of biosphere . 400:547–50. Caspersen JP, Pacala SW, Jenkins JC, Hurtt GC, Moorcroft PR, Randerson JT, Chapin FS, Harden JW, Neff JC, Harmon ME. Birdsey RA. 2000. Contributions of land-use history to carbon 2002. Net ecosystem production: a comprehensive measure of accumulation in US . Science 290:1148–51. net carbon accumulation by ecosystems. Ecol Appl 12:937–47. Chapin FS. III, Matson PA, Mooney HA. 2002. Principles of Raymond PA, Cole JJ. 2003. Increase in the export of alkalinity terrestrial New York: Springer. form ’s largest river. Science 302:88–91. Cole JJ, Pace ML, Carpenter SR, Kitchell JF. 2000. Persistence of Schlesinger WH. 1997. : An Analysis of Global net heterotrophy in lakes during nutrient addition and Change. 2nd ed. Academic Press, San Diego, p. 159. web manipulations. Limnol Oceangr 45(8):1718–30. Schulze ED, Wirth C, Heimann M. 2000. Managing forest after Hamilton JG, DeLucia EH, George K, Naidu SL, Finzi AC, Kyoto. Science 289:2058–2059. Schlesinger WH. 2002. Forest carbon balance under elevated Siegenthaler U, Sarmiento JL. 1993. Atmospheric carbon dioxide CO2. Oecologia 131:250–60. and the ocean. Nature 365(9):119–25. Hanson PC, Bade DL, Carpenter SR, and Kratz TK. 2003. Lake Woodwell GM, Whittaker RH. 1968. Primary production in : relationships with dissolved organic carbon and terrestrial ecosystems. Am Zoologist 8:19–30. . Limnol Oceanogr 48:1112–9.