R implementation of the P-model and its corollary predictions (Prentice et al., 2014; Han et al., 2017).
rpmodel( tc, vpd, co2, fapar, ppfd, patm = NA, elv = NA, kphio = ifelse(do_ftemp_kphio, ifelse(do_soilmstress, 0.087182, 0.081785), 0.049977), beta = 146, soilm = stopifnot(!do_soilmstress), meanalpha = 1, apar_soilm = 0, bpar_soilm = 0.733, c4 = FALSE, method_optci = "prentice14", method_jmaxlim = "wang17", do_ftemp_kphio = TRUE, do_soilmstress = FALSE, returnvar = NULL, verbose = FALSE )
tc | Temperature, relevant for photosynthesis (deg C) |
---|---|
vpd | Vapour pressure deficit (Pa) |
co2 | Atmospheric CO2 concentration (ppm) |
fapar | (Optional) Fraction of absorbed photosynthetically active
radiation (unitless, defaults to |
ppfd | Incident photosynthetic photon flux density
(mol m-2 d-1, defaults to |
patm | Atmospheric pressure (Pa). When provided, overrides
|
elv | Elevation above sea-level (m.a.s.l.). Is used only for
calculating atmospheric pressure (using standard atmosphere (101325 Pa),
corrected for elevation (argument |
kphio | Apparent quantum yield efficiency (unitless). Defaults to
0.081785 for |
beta | Unit cost ratio. Defaults to 146.0 (see Stocker et al., 2019). |
soilm | (Optional, used only if |
meanalpha | (Optional, used only if |
apar_soilm | (Optional, used only if |
bpar_soilm | (Optional, used only if |
c4 | (Optional) A logical value specifying whether the C3 or C4
photosynthetic pathway is followed.Defaults to |
method_optci | (Optional) A character string specifying which method is
to be used for calculating optimal ci:ca. Defaults to |
method_jmaxlim | (Optional) A character string specifying which method
is to be used for factoring in Jmax limitation. Defaults to |
do_ftemp_kphio | (Optional) A logical specifying whether
temperature-dependence of quantum yield efficiency after Bernacchi et al.,
2003 is to be accounted for. Defaults to |
do_soilmstress | (Optional) A logical specifying whether an empirical
soil moisture stress factor is to be applied to down-scale light use
efficiency (and only light use efficiency). Defaults to |
returnvar | (Optional) A character string of vector of character strings specifying which variables are to be returned (see return below). |
verbose | Logical, defines whether verbose messages are printed.
Defaults to |
A named list of numeric values (including temperature and pressure dependent parameters of the photosynthesis model, P-model predictions, including all its corollary). This includes :
ca
: Ambient CO2 expressed as partial pressure (Pa)
gammastar
: Photorespiratory compensation point \(\Gamma*\),
(Pa), see gammastar.
kmm
: Michaelis-Menten coefficient \(K\) for photosynthesis
(Pa), see kmm.
ns_star
: Change in the viscosity of water, relative to its
value at 25 deg C (unitless).
$$\eta* = \eta(T) / \eta(25 deg C)$$
This is used to scale the unit cost of transpiration.
Calculated following Huber et al. (2009).
chi
: Optimal ratio of leaf internal to ambient CO2 (unitless).
Derived following Prentice et al.(2014) as:
$$
\chi = \Gamma* / ca + (1- \Gamma* / ca) \xi / (\xi + \sqrt D )
$$
with
$$
\xi = \sqrt (\beta (K+ \Gamma*) / (1.6 \eta*))
$$
\(\beta\) is given by argument beta
, \(K\) is
kmm
(see kmm), \(\Gamma*\) is
gammastar
(see gammastar). \(\eta*\) is ns_star
.
\(D\) is the vapour pressure deficit (argument vpd
), \(ca\) is
the ambient CO2 partial pressure in Pa (ca
).
ci
: Leaf-internal CO2 partial pressure (Pa), calculated as \((\chi ca)\).
lue
: Light use efficiency (g C / mol photons), calculated as
$$
LUE = \phi(T) \phi0 m' Mc
$$
where \(\phi(T)\) is the temperature-dependent quantum yield efficiency modifier
(ftemp_kphio) if do_ftemp_kphio==TRUE
, and 1 otherwise. \(\phi 0\)
is given by argument kphio
.
\(m'=m\) if method_jmaxlim=="none"
, otherwise
$$
m' = m \sqrt( 1 - (c/m)^(2/3) )
$$
with \(c=0.41\) (Wang et al., 2017) if method_jmaxlim=="wang17"
. \(Mc\) is
the molecular mass of C (12.0107 g mol-1). \(m\) is given returned variable mj
.
If do_soilmstress==TRUE
, \(LUE\) is multiplied with a soil moisture stress factor,
calculated with soilmstress.
mj
: Factor in the light-limited assimilation rate function, given by
$$
m = (ci - \Gamma*) / (ci + 2 \Gamma*)
$$
where \(\Gamma*\) is given by gammastar
.
mc
: Factor in the Rubisco-limited assimilation rate function, given by
$$
mc = (ci - \Gamma*) / (ci + K)
$$
where \(K\) is given by kmm
.
gpp
: Gross primary production (g C m-2), calculated as
$$
GPP = Iabs LUE
$$
where \(Iabs\) is given by fapar*ppfd
(arguments), and is
NA
if fapar==NA
or ppfd==NA
. Note that gpp
scales with
absorbed light. Thus, its units depend on the units in which ppfd
is given.
iwue
: Intrinsic water use efficiency (iWUE, Pa), calculated as
$$
iWUE = ca (1-\chi)/(1.6)
$$
gs
: Stomatal conductance (gs, in mol C m-2 Pa-1), calculated as
$$
gs = A / (ca (1-\chi))
$$
where \(A\) is gpp
\(/Mc\).
vcmax
: Maximum carboxylation capacity \(Vcmax\) (mol C m-2) at growth temperature (argument
tc
), calculated as
$$
Vcmax = \phi(T) \phi0 Iabs n
$$
where \(n\) is given by \(n=m'/mc\).
vcmax25
: Maximum carboxylation capacity \(Vcmax\) (mol C m-2) normalised to 25 deg C
following a modified Arrhenius equation, calculated as \(Vcmax25 = Vcmax / fv\),
where \(fv\) is the instantaneous temperature response by Vcmax and is implemented
by function ftemp_inst_vcmax.
jmax
: The maximum rate of RuBP regeneration () at growth temperature (argument
tc
), calculated using
$$
A_J = A_C
$$
rd
: Dark respiration \(Rd\) (mol C m-2), calculated as
$$
Rd = b0 Vcmax (fr / fv)
$$
where \(b0\) is a constant and set to 0.015 (Atkin et al., 2015), \(fv\) is the
instantaneous temperature response by Vcmax and is implemented by function
ftemp_inst_vcmax, and \(fr\) is the instantaneous temperature response
of dark respiration following Heskel et al. (2016) and is implemented by function
ftemp_inst_rd.
Additional variables are contained in the returned list if argument method_jmaxlim=="smith19"
omega
: Term corresponding to \(\omega\), defined by Eq. 16 in
Smith et al. (2019), and Eq. E19 in Stocker et al. (2019).
omega_star
: Term corresponding to \(\omega^\ast\), defined by
Eq. 18 in Smith et al. (2019), and Eq. E21 in Stocker et al. (2019).
Bernacchi, C. J., Pimentel, C., and Long, S. P.: In vivo temperature response func-tions of parameters required to model RuBP-limited photosynthesis, Plant Cell Environ., 26, 1419–1430, 2003
Heskel, M., O’Sullivan, O., Reich, P., Tjoelker, M., Weerasinghe, L., Penillard, A.,Egerton, J., Creek, D., Bloomfield, K., Xiang, J., Sinca, F., Stangl, Z., Martinez-De La Torre, A., Griffin, K., Huntingford, C., Hurry, V., Meir, P., Turnbull, M.,and Atkin, O.: Convergence in the temperature response of leaf respiration across biomes and plant functional types, Proceedings of the National Academy of Sciences, 113, 3832–3837, doi:10.1073/pnas.1520282113,2016.
Huber, M. L., Perkins, R. A., Laesecke, A., Friend, D. G., Sengers, J. V., Assael,M. J., Metaxa, I. N., Vogel, E., Mares, R., and Miyagawa, K.: New international formulation for the viscosity of H2O, Journal of Physical and Chemical ReferenceData, 38, 101–125, 2009
Prentice, I. C., Dong, N., Gleason, S. M., Maire, V., and Wright, I. J.: Balancing the costs of carbon gain and water transport: testing a new theoretical frameworkfor plant functional ecology, Ecology Letters, 17, 82–91, 10.1111/ele.12211,http://dx.doi.org/10.1111/ele.12211, 2014.
Wang, H., Prentice, I. C., Keenan, T. F., Davis, T. W., Wright, I. J., Cornwell, W. K.,Evans, B. J., and Peng, C.: Towards a universal model for carbon dioxide uptake by plants, Nat Plants, 3, 734–741, 2017. Atkin, O. K., et al.: Global variability in leaf respiration in relation to climate, plant func-tional types and leaf traits, New Phytologist, 206, 614–636, doi:10.1111/nph.13253, https://nph.onlinelibrary.wiley.com/doi/abs/10.1111/nph.13253.
Smith, N. G., Keenan, T. F., Colin Prentice, I. , Wang, H. , Wright, I. J., Niinemets, U. , Crous, K. Y., Domingues, T. F., Guerrieri, R. , Yoko Ishida, F. , Kattge, J. , Kruger, E. L., Maire, V. , Rogers, A. , Serbin, S. P., Tarvainen, L. , Togashi, H. F., Townsend, P. A., Wang, M. , Weerasinghe, L. K. and Zhou, S. (2019), Global photosynthetic capacity is optimized to the environment. Ecol Lett, 22: 506-517. doi:10.1111/ele.13210
Stocker, B. et al. Geoscientific Model Development Discussions (in prep.)
if (FALSE) { rpmodel( tc = 20, vpd = 1000, co2 = 400, ppfd = 30, elv = 0) }