LandscapeDNDC  1.36.0
PSIM - Physiological Simulation Model

The PSIM runs in sub-daily (hourly) time steps and gets assimilated carbon based on a Farquhar approach [18], calculated in the photofarquhar.cpp, which is considered along with the ideas of Ball et al. [5] regarding consideration of stomatal conductance (ld_berryball.cpp).

Processes are generally calculated specifically for species (as cohorts containing trees of equal dimensions), canopy- (2-40) and soil layers (individually set in the site properties). In addition, long-living foliage is considered in different age classes.

In the module itself, respiration is considered to originate from either biomass growth, nitrate conversion, or maintenance of different compartments (foliage, fine roots, structural reserves, living wood) and depends on temperature and nitrogen concentrations [9]. Allocation from net photosynthesis (or nitrogen uptake) into the respective compartments is realized according to its sink strength that is determined from targed allometric relations and dynamic gain and loss processes [23]. Therefore, the model assumes a certain longevity of each compartment, resulting in an empirically determined senescence.

Specifically, the main physiological processes of carbon and nitrogen in the plant are:

  • hydraulic conductance (PSIM_HydraulicConductance)
  • growth of new foliage from buds (PSIM_BudBurst)
  • senescence (PSIM_Senescence)
  • carbon allocation and growth (PSIM_CarAllocation)
  • nitrogen allocation (PSIM_NitAllocation)
  • respiration (PSIM_Respiration)
  • nitrogen uptake (PSIM_NitrogenUptake) Which all are described further down.

In addition, a change in enzymatic capacity is derived indicating a seasonality and stress-dependence of photosynthetic enzyme activity (PSIM_PhotosynthesisRates).

Besides from the photosynthesis input (see PhotoFarquhar), the module depends on variables describing:

  • microclimate, i.e. daily average canopy temperature for budburst, soil temperature for root growth limitations, and temperature in all canopy- and soil layers for mainanence respiration
  • air and soil chemistry, i.em. nitrogen concentrations in the air (no2, nh3) and the soil (no3, nh4) in in all canopy- and soil layers for nitrogen uptake
  • vegetation structure, i.e. distribution of foliage and fine roots throughout canopy and soil layers. In addition, tree dimensions are needed for sapwood allocation.

Modes

The calculations are reduced to the seasonality, flushing and senescence modules if the model runs in the MoBiLE_IsOneLeaf mode where plant growth is not of interest.

Author
  • Ruediger Grote

Model options

Available options (default options are marked with bold letters):

  • Branch fraction ("branchfraction" = diameter / volume)
    Branch fraction is either estimated from parameterized tree diameter relationship (option: diameter) or from crown volume (option: volume).
  • Crown length ("crownlength" = diameter / height)
    Crown length is either estimated from a parameterized relation to tree height (option: height, see: veglibs_crown-length_parameter) or based on crown diameter (option: diameter).
  • Competition ("competition" = true / false)
    The Competition effect for stand density on height/diameter ratios is either neglected (option: false) or considered (option: true).
  • Forest structure ("foreststructure" = true / false )
    Structural growth either neglected (option: false) or considered (option: true). If structural growth is not included, seasonal woody biomass growth is reset in order to avoid structural growth. Removed woody biomass is added to the litter pools (see: PSIM_ResetBiomassFromLastYear)
  • Transpiration (transpirationmethod: wateruseefficiency / potentialtranspiration)
    Transpiration is either calculated using fixed relations to photosynthesis (WUECMAX, WUECMIN), or the transpiration that is defined by stomatal conductance, depending on vapor pressure deficit and plant hydraulic conductance (with GSMIN as minimum conductance).
  • Stomatal conductance (stomatalconductance: leuning_1995 / ballberry_1987 / medlyn_2011(a,b) / eller_2020)
    Various stomatal conductance routines are available with and without consideration of soil water supply. While Leuning et al. [39] with linear reduction of conductance with relative available soil water is the standard option [35], Eller et al. [15] is the only option that considers plant water potential instead soil water availability.

Hydraulic conductance

The hydraulic approach calculates canopy water potential based on soil water potential, which then affects stomal conductance following the 'Stomata on Xylem' (SOX) model (see: Farquhar model): First, xylem water potential is derived from the soil conditions, weighting soil layer importance by fine root abundance and considering a threshold soil water potential at which plant tissues are decoupled from the soil:

\[ psi\_{sr} = Sum[(min(CSR\_{REF}; psi_{sl} \cdot fFrt_{sl})] \]

with

  • CSR_REF: Threshold soil water potential at which roots and soil get decoupled (MPa)
  • psi: Soil water potential (MPa) (see assign_soillayer_van_genuchten_parameter)
  • fFrt: Biomass fraction of fine roots in a particular soil layer
  • sl: Indicator for a specific soil layer

Canopy water potential is then calculated from the gradient between the water potential in the xylem and from evaporative demand, also including the gravitational force.

\[ psi\_{can} = (psi\_{sr} + psidecline\_{cum}) - (\frac {transp}{CWP\_{REF} \cdot kxyl} ) - dpsi \]

with

  • psidecline_cum: previous day cumulative water deficit within the tree
  • kxyl: previous day root-to-canopy (plant hydraulic) conductance (in mol MPa-1 s-1 m-2leaf)
  • transp: current day transpiration (mol m-2leaf s-1)
  • CWP_REF: reference (=maximum) value for the (leaf area-normalized) specific xylem conductance (mol MPa-1 m-2leaf s-1)
  • dpsi: water potential decline due to plant height (MPa)

The water deficit is build from the water that is transpired but was not available from the rooting zone. It is supposed to reflect the water resources within a tree, i.e. the stem water content. Surplus on water supply will refill this storage while additional deficits will empty it further.

The total plant, or xylem, conductance is calculated based on empirical relations with species-specific parameters (PSI_REF, PSI_EXP).

\[ kxyl = 1.0 - (1.0 - exp(-((\frac {psi\_{can}} {PSI\_{REF}})^{PSI\_{EXP}} ))) \]

Note
The CSR_REF parameter defines a limit for water uptake that is in principle the same as the wilting point and might replace this site property.

Phenology

Leaf flushing is calculated using the growing degree days approach, which is using the temperature sum since this years 1st January, considering chilling requirements and drought stress limitations.

Therefore, temperature sum (cumulative growing degree days, gdd),is first calculated from daily temperature sum ([38]):

\[ gdd\_cum = temp \cdot \frac{dayl}{12.0} \]

with

  • temp: daily average leaf surface temperature (oC)
  • dayl: daylength (h)

When cumulative gdd reaches a threshhold value (gdd_thresh) that depends on the degree chilling requirements are met, flushing is initiated. This value is calculated as:

\[ gdd\_thresh = 100.0 + GDDSTART \cdot exp( -0.0075 \cdot chilldays) \]

with

  • GDDSTART: temperature sum value for flushing without considering chilling, ON TOP of the minimum GDD which is assumed to be 100.

Chilling days (chilldays) are calculated from average temperature and amplitude [43] :

\[ chilldays = 2.0 \cdot arcos( 1.0 - ( max( 0.0, - \frac {temp\_avg - 0.5 \cdot temp\_ampl} {0.5 \cdot temp\_ampl} )) \cdot \frac {diy}{2.0 \cdot PI} \]

with

  • temp_avg: average annual temperature (oC)
  • temp_ampl: annual temperature amplitude (oC)
  • diy: number of days in year (-)
  • PI: constant (3.1416)

After temperature requirements are met, flushing goes on provided water supply is above a species-specific treshold value (H2OREF_FLUSHING) and there is no water deficit in the stem.

Senescence

Dry matter senescence loss is calculated assuming fixed tissue longevity separately for foliage, fine roots, and living (sap-)wood. Fine root and sapwood senescence is decreased by a species-specific fraction each day. Senescence of buds is directly related to foliage growth, which is driven by a similar function than foliage senescence (see Carbon allocation and growth).

Foliage senescence (sFol) is empirically determined (for evergreen species in every age class, na) based on a mortality factor ranging between 1 and 0:

\[ sFol = mFol\_{na} \cdot (1.0 - \frac {1.0 - dvsmort} {1.0 - dvsmort\_{old}} ) \]

\[ dvsmort = exp( -1.0 \cdot \frac { foliage\_{age} - DLEAFSHED)^{ 2 } } { (0.5 \cdot NDMORTA)^ { 2 } \cdot log(2.0) } ) \]

with:

  • mFol: foliage biomass (kgDW m-2)
  • foliage_age: age of foliage (age class) (days)
  • DLEAFSHED: parameter describing the day where all foliage (of one age class) is shed (days)
  • NDMORTA : parameter indicating the duration of senescence (days)

The nitrogen loss is defined by the nitrogen concentration in the tissue considering a species-specific retranslocation rate that is the same for all tissues. The maximum retranslocation rate is reduced if the overall nitrogen demand is smaller than the maximum amount of nitrogen that could be retranslocated.

Under some circumstances, stress induced senescence (i.e. xylem and foliage loss due to extensive drought stress) is also considered. This part is under development.

Carbon allocation and growth

Carbon provided by photosynthesis (minus growth respiration) is distributed into any compartment that didn't comply with parameterized allometric relations which define the optimum biomass. Allocation strength is linearly related to the difference between actual and optimum biomass [23] .

\[ dc_{C} = cPool \cdot afc_{C} - res_{C}Old \]

\[ afc_{C} = \frac {dem_{C} }{cPool } \]

\[ dem_{C} = \frac { max(0.0, m_{C}Opt - m_{C}) \cdot CCDM }{1.0 - FYIELD} + res_{C}Old \]

with

  • C: indicator for plant compartments foliage, fine roots, sapwood, buds (= structural reserves), and the free available carbon (nonstructural reserves) that are represented in percent of all other living tissues (fac)
  • cPool: available carbon for allocation originating from photosynthesis minus growth respiration
  • afc: allocation factor (0-1)
  • res: (maintenance) respiration from the compartments in the last timesteps (C m-2)
  • dem: carbon demand (C m-2)
  • m: biomass (DW m-2)
  • m_Opt: optimum biomass of a compartment (DW m-2)
  • CCDM: constant transforming biomass to carbon values (carbon content)
  • FYIELD: species-specific parameter for growth efficiency (growth respiration fraction)

The foliage compartment is primarily increased by the depletion of reserves (bud compartment), and therefore has no 'demand' to its own. The transfer and thus the increase of foliage biomass is triggering the demand of buds, sapwood, and fine roots via allometric relations:

\[ m_{Frt}Opt = QRF \cdot mFol; \]

\[ m_{Sap}Opt = qsfm \cdot mFol; \]

\[ m_{Bud}Opt = MFOLOPT; \]

\[ m_{Fac}Opt = sum( m_{C} ) \cdot (FACMAX - fac); \]

with

  • frt, sap, bud, fac: descriptors for plant compartments finroots, sapwood, buds (= constitutive reserves)
  • qsfm: desired relation between sapwood and foliage biomass
  • QRF: species-specific parameter describing the desired relation between fine roots and foliage biomass
  • MFOLOPT: species-specific parameter for foliage biomass per area (kgDW m-2ground)
  • FACMAX: species-specific parameter for maximum percentage of free available carbon

qsfm is derived from a desired relation between sapwood area and foliage area (Huber value, QSFA) and the dimension of the tree (assuming species-specific taper functions and fixed fractions for branchiness and coarse roots.

If the supply is larger than all demands, the surplus is distributed between buds, foliage, and fine roots in case of herbaceous plants, wood and buds in case of determined growth (FREEGROWTH = false) or into foliage growth if species are growing leaves continously (FREEGROWTH = true).

Free available carbon can be depleted only by respiration demands higher than supply rates.

Carbon exudation is assumed to be species specific fraction of total fine root growth (see: PSIM_SoilCarbonRelease).

Nitrogen allocation

The nitrogen provided from uptake and retranslocation is distributed according to biomass growth and optimum tissue concentrations [23].

Respiration

Respiration consists of growth (rGro) and maintenance respiration, with the latter split up into respiration that is related to nutrient uptake and transport (rTra), and into remaining (residual) respiration (rRes). I Growth respiration is a fraction of growth is assumed to apply equally for every tissue

\[ rGro = sum( dc_{C} \cdot (1.0 - FYIELD) ) \]

with

  • C: indicator for plant compartments foliage, fine roots, sapwood, buds (= structural reserves)
  • FYIELD: species-specific parameter expressing fraction of growth parameters

II Transport respiration summarizes carbon costs due to uptake of nitrogen and other nutrients (rUpt), phloem transport of carbon into the roots (rPhl), and the reduction of oxygenized nitrogen compounds (rNit). Uptake of nitrogen compounds is explicitly modeleled, while additional costs from uptake and incorporation of other nutrients are estimated from biomass growth, assuming a that all requirements are met.

\[ rTra = rUpt + rPhl + rNit \]

\[ rUpt = PAMM \cdot (uptNH4 + uptNH3) + PNIT \cdot (uptNO3 + uptNOx) + PMIN \cdot growth \cdot \frac{FMIN}{CCDM} \]

\[ rPhl = PPHLOE \cdot (dcFrt + dcSap + exsuLoss) \]

\[ rNit = PREDFRT \cdot uptNO3 \cdot FRFRT + PREDSAP \cdot uptNO3 \cdot (1.0 - FRFRT) + PREDSAP \cdot uptNOx \]

with

  • uptNH4, uptNH3: taken up reduced nitrogen from the air (NH4) and the soil (NH3)
  • uptNOx, uptNO3: taken up oxygenized nitrogen from the air (NOx) and the soil (NO3)
  • dcFrt, dcSap: carbon increase of fineroots and sapwood (C)
  • exsuLoss: carbon loss by exudation out of fine roots into the soil (C)
  • PAMM, PNIT, PMIN: parameters for uptake cost of ammonia, nitrate, and other mineral components (0.17, 0.34, 0.06)
  • FMIN: parameter for fraction of mineral components other than nitrogen to total plant biomass (0.05)
  • CCDM: constant indicating the carbon content of plant dry matter (0.45)
  • PPHLOE: parameter for phloem loading costs to support growth in sapwood, fine roots as well a exudates (0.06)
  • PREDFRT, PREDSAP: parameters for reduction costs of taken up oxygenized nitrogen forms in fine roots and sapwood, respectively (1.72, 0.855)
  • FRFRT: parameter indicating the fraction of nitrate that is reduced in the roots (0.5)

III Residual respiration according to temperature and nitrogen content [9]

\[ rRes = sum( km \cdot fsub \cdot n_{C} ) \]

\[ km = KM20 \cdot (temp_{C} - TRMIN) ^ { 2 } \cdot (TRMAX - temp_{C}) \cdot \frac{1.0}{ ( TROPT - TRMIN) ^ { 2 } \cdot ( TRMAX - TROPT) } \]

\[ fsub = \frac {ffac \cdot CCDM}{KMMM + ffac \cdot CCDM} \]

with

  • C: indicator for plant compartments foliage, fine roots, sapwood, buds (= structural reserves)
  • n: nitrogen within a plant compartment
  • fsub: reduction factor due to depleted non-structural reserves
  • km: temperature modification factor
  • temp: tissue temperature
  • ffac: degree to which the virtual pool of non-structural carbon reserves is filled up (0-1)
  • KMMM: Michaelis-Menton constant for the importance of relative available non-structural carbon on respiration

Nitrogen uptake

Uptake of ammonia and nitrate from the soil and canopy is calculated according to supply, soil water availability and fine root density. Total nitrogen uptake is limited to the current whole plant demand. Nitrogen uptake/emission from the canopy depends on NOx air concentration.

Note
Uptake through the canopy may be small but could be significant relative to NOx emissions from the soil. Parameterization should be species specific but necessary information is yet not available. Instead, rough estimates from a tropical landscape are used [51].
Nitrogen fixation is considered similar to agricultural plants, using a species-specific parameter (INI_N_FIX) that indicates that a fraction of nitrogen demand is covered by fixation at no cost (see PSIM_NitrogenFixation).

Daily enzyme activity

\[ v25_{E} = V25_{E} \cdot fdorm \cdot fnit \cdot fwat \]

with

  • v25: value for velocity at 25 oC (umol m-2 s-1)
  • E: indicator for different enzymes (carboxylation reactivity, electron transport activity, dark respiration)
  • V25: parameterized standard values for enzyme velocities at 25 oC (umol m-2 s-1)
  • fdorm, fnit, and fwat: multiplicative reduction factors for phenology, nitrogen- and water supply, respectively

The reduction factor for phenology is differently calculated for deciduous species, where enzyme activity is assumed to develop in parallel to flushing and senescence, and for evergreen species, where the activity is determined from temperature development according to Maekelae et al [48] :

\[ fdorm = C1 \cdot (tFol24 + \frac{24.0}{TAU} \cdot ( temp - tFol24) - PSNTFROST) \]

with

  • temp: (leaf surface) temperature (oC)
  • tFol24: temperature average of the previousl 24 hours (oC)
  • C1, TAU: empirical model constants (0.0367, 330.0, respectively) [48]
  • PSNTFROST: species-specific temperature at which photosynthesis enzymes start to decline (oC)

The reduction factor for nitrogen supply is defined as the degree to which the optimum (equal to maximum) foliage nitrogen concentration has been established:

\[ fnit = \frac {ncFol - NCFOL\_{MIN} }{NCFOL\_{OPT} - NCFOL\_{MIN} } \]

with

  • ncFol: foliage nitrogen concentration (gN gDM-1)
  • NC_FOL (MIN, OPT): species-specific minimum and optimum foliage nitrogen concentrations (gN gDM-1)

The reduction factor for water supply is based on the same function as has been used for determining the loss of hydraulic conductance according to [57]. The recovery of the water damaged enzyme activity is done with a limited rate of 5% (currently fixed).

\[ fwat = \frac {1.0 + exp(A\_{REF} \cdot A\_{EXP}) } {1.0 + exp((A\_{REF} - psi\_{mean}) \cdot A\_{EXP}) } \]

with

  • psi_mean: mean canopy water potential (MPa)
  • A_REF, A_EXP: species-specific empirical parameters that describe the sensitivity (EXP) and a reference water potential (REF) of the drought impact on enzyme activity
Note
vcAct25 is not changed in dependence on season alone as is e.g. indicated in [31].