

Custom Products
EppleyVGPM Net Primary Production (NPP) calculationsThe following text provides a brief description of the EppleyVGPM calculations. Select one of the other tabs above to find similar information regarding the other Custom Products algorithms. The EppleyVGPM The EppleyVGPM is a sort of hybrid model that employs the basic model structure and parameterization of the standard VGPM ( Behrenfeld and Falkowski (1997a) ) but replaces the polynomial description of Pb_opt with the exponential relationship described by Morel (1991) and based on the curvature of the temperaturedependent growth function described by Eppley (1972) (see discussion on Home Page). The exponential Pb_opt function is commonly used in NPP models. Its application in the VGPM structure yield NPP distributions that are highly consistent with other algorithms employing the same exponential "Eppley" curve. The EppleyVGPM is a "chlorophyllbased" algorithm (see discussion on Home Page) and is similar in form to the early models of Ryther and Yentch (1957) and Talling (1957). The foundation of the EppleyVGPM and other chlorophyllbased models is that NPP varies in a predictable manner with chlorophyll concentration (chl): NPP = f(chl) Because NPP is a rate and chlorophyll is a standing stock, derivation of the former from the later requires a "rate" term, specifically a chlorophyllspecific assimilation efficiency for carbon fixation. The description of this rate term is the single most important uncertainty in all chlorophyll based models. The EppleyVGPM employs a variable termed Pb_opt, which is the maximum daily net primary production found within a given water column and expressed in units of mg carbon fixed per mg chlorophyll per hour. NPP at the depth of Pb_opt is thus: NPP = chl * pb_opt * day length where day length is the number of hours of day light at the location of interest and NPP is milligrams of carbon fixed per day per unit volume. For studies of global ocean productivity and understanding its relationship to the ocean carbon cycle and ecosystem functioning, the quantity we are interested in is water column integrated productivity per unit of ocean area. The EppleyVGPM thus needs a function to project surface NPP values through the water column to get production per unit surface area: NPP = chl * pb_opt * day length * volume function Volume functionThe volume function is often an aspect of confusion for people working with the EppleyVGPM or other chlorophyllbased NPP models. Here's how I like to think about it ... Water column NPP is generally regarded as the primary production taking place from the surface to the depth at which 1% of surface light is available. We call this light depth the "euphotic depth" or "z_eu". If you were to consider the hypothetical condition where photosynthetic rates were uniform from the surface to z_eu, then water column production could simply be calculated as: NPP = chl * pb_opt * day length * z_eu or, in words, surface production times the euphotic depth. In the real world, however, photosynthesis through the water column is far from constant. The most important factor driving this vertical variability is light. As sunlight penetrates the water column, some of it is absorbed and scattered backward. Consequently, sunlight decreases rapidly with depth in a near exponential manner. If it is really bright at the surface, photosynthesis will be light saturated and relatively constant in the upper layer, but eventually it will begin to decrease with depth toward z_eu. If surface light levels are low (e.g., cloudy day or high latitude winter), photosynthetic rates will be maximal right at the surface and decrease rapidly throughout the euphotic zone. These effects of light on water column production are accounted for in the VGPM by including a lightdependent term, f(par), in the volume function: volume function = f(par) * z_eu The f(par) term can be thought of as the ratio of realized water column integrated NPP to the maximum potential NPP if photosynthetic rates were maintained at maximum levels (i.e., Pb_opt) throughout the water column. The paramterization of this lightdependent term in the VGPM was determined empirically using thousands of field productivity measurements and is given by: f(par) = 0.66125 * par / ( par + 4.1 ) Similar relationships have been derived empirically and theoretically on multiple occasions, with the most significant difference between expressions reflecting the severity of nearsurface photoinhibition (i.e., decreases in photosynthesis due to damage by excessive light levels) inherent in the field data employed or assumed in the photosynthesisirradiance model chosen. Replacing the "volume function" with the above two equations, yields the basic EppleyVGPM relationship: NPP = chl * pb_opt * day length * [0.66125 * par / ( par + 4.1 )] * z_eu For more details on the volume function, the f(par) relationship, and differences between NPP algorithms, see Platt & Sathyendranath (1993) and Behrenfeld & Falkowski (1997b). Pb_optChlorophyllbased NPP models take many forms. Some models are simple expressions relating surface properties to water column integrated products. Other models are highly sophisticated, describing the spectral attenuation of light through the water column, depthdependent changes in phytoplankton pigment concentration, and timeresolved light absorption by phytoplankton. But, in the end, the two primary factors that control differences and similarities between chlorophyllbased NPP models are the choice of input chlorophyll data and the description of how lightsaturated photosynthetic efficiencies vary in the environment (see Campbell et al. 2002, Carr et al. 2006). All NPP models require this description of physiological variability [whether it is based on daily integrated production measurements (Pb_opt) or "instantaneous" photosynthesisirradiance measurements (Pb_max)] and it is universally the "Achilles tendon" of each algorithm. For the EppleyVGPM, physiological variability is linked to the Pb_opt variable and it is described as a function of sea surface temperature. The EppleyVGPM Pb_opt function is an exponent that has a slope taken from a maximum growth rate model by Eppley (1972) and parameterized to yield a Pb_opt value of 4.6 at 20 degrees C. pb_opt = 1.54 * 10**[(0.0275*sst)  0.07] From a physiological standpoint, the dependence of Pb_opt on temperature is not envisioned as reflecting a direct effect of temperature on carbon fixation efficiencies. Instead, the monotonically increasing values of Pb_opt with temperature in the EppleyVGPM are viewed as reflecting a correlation between sea surface temperature (SST) and photoacclimation. Specifically, warmer waters generally have shallower physiological surface mixing depths and higher incident PAR. These conditions favor highlight acclimated phytoplankton, which requires less chlorophyll to support photosynthetic requirements for growth. This lower chlorophyll concentration give rise to high chlorophyllspecific photosynthetic rates. In other words, increasing Pb_opt with increasing SST. Of course there is also the countering effect that warm ocean areas are also generally accompanied by vanishingly low surface nutrient concentrations. Nutrient stress causes Pb_opt to decrease. The EppleyVGPM does not account for nutrient stress effects by allowing Pb_opt to decrease in warm waters, but does take into account simultaneous influences of highlight photoacclimation as SST increases. Thus, the standard VGPM Pbopt relationship emphasizes the nutrientstress effect at high sea surface temperatures (SSTs), while the EppleyVGPM Pb_opt relationship emphasizes the photoacclimation effect at high SSTs. (see Website Home Page for more discussion) z_euEuphotic depth (z_eu) in the EppleyVGPM is calculated using the Morel and Berthon (1989) Case I model. This model estimates z_eu from surface chlorophyll concentrations and is based on empirical equations to fit field data. In practice, total water column chlorophyll concentration is calculated from satellite surface chlorophyll using a formula that distinguishes between lower and higher chlorophyll waters. Then, given the amount of total chlorophyll, the euphotic depth is estimated, again using separate equations for lower and higher total chlorophyll conditions. See Morel and Berthon (1989) for more details. In essence, light penetration is inversely related to chlorophyll: the more phytoplankton, the shallower the euphotic depth, and vice versa. Variables neededThe EppleyVGPM equation is: NPP = chl * pb_opt * day length * f(par) * z_eu NPP calculations with the EppleyVGPM require the following input data fields:
These ancillary data can be accessed here. For details on EppleyVGPM implementation, please see the code. 



