LandscapeDNDC  1.36.0
PlaMox - Plant Growth Model

User guide

PlaMox simulates the carbon and nitrogen cycle of crops and grass species. Processes are described in a universal way and plants are primarily distinguished by species-specific parameters that can be accessed and calibrated externally. However, for a growing number of specific species (e.g., rice, maize, ...), there exist specific functionalities, which are continuously developed.

Model structure

PlaMox includes a submodel for photosynthesis calculation based on von Caemmerer and Farquahr 1981. Since the photosynthesis submodel requires a subdaily time step, PlaMox can also only be used with a subdaily time resolution. The recommendation is 24 time steps per day. PlaMox requires further models for:

  • watercycle (e.g., transpiration)
  • soilchemistry (e.g., nitrogen uptake)
  • microclimate (e.g., radiation distribution within the canopy)

Parametrization

The following lists include species parameters that might be calibrated in order to represent a specific plant. See the description of respective sections for more details on parameter behaviour.

Photosynthesis:

  • \( C4\_TYPE \)
  • \( VCMAX25 \) (rubisco activity)
  • \( SLAMAX \) (leaf area)
  • \( SLADECLINE \) (decline of specific leaf area with plant development)

Nitrogen related parameters:

  • \( NC\_FRUIT\_MAX \) (optimum nitrogen content of the fruit)
  • \( NC\_FRUIT\_MIN \) (minimum nitrogen content of the fruit)
  • \( NC\_FINEROOTS\_MAX \) (optimum nitrogen content of fine roots)
  • \( NC\_FINEROOTS\_MIN \) (minimum nitrogen content of fine roots)
  • \( NC\_FOLIAGE\_MAX \) (optimum nitrogen content of foliage)
  • \( NC\_FOLIAGE\_MIN \) (minimum nitrogen content of foliage)
  • \( NC\_STRUCTURAL\_TISSUE\_MAX \) (optimum nitrogen content of structural tissue)
  • \( NC\_STRUCTURAL\_TISSUE\_MIN \) (minimum nitrogen content of structural tissue)

Allocation related parameters:

  • \( FRACTION\_ROOT \) (assimilated carbon fraction allocated to roots)
  • \( FRACTION\_FRUIT \) (assimilated carbon fraction allocated to the fruit)
  • \( FRACTION\_FOLIAGE \) (assimilated carbon fraction allocated to foliage)
  • \( MFOLOPT \) (optimum foliage biomass)

Plant development related parameters:

  • \( GDD\_BASE\_TEMPERATURE \)
  • \( GDD\_MAX\_TEMPERATURE \)
  • \( GDD\_STEM\_ELONGATION \)
  • \( GDD\_FLOWERING \)
  • \( GDD\_GRAIN\_FILLING \)
  • \( GDD\_MATURITY \)

Vernalization related parameters:

  • \( CHILL\_TEMP\_MAX \)
  • \( CHILL\_UNITS \)

Cutting

  • \( SHOOT\_STIMULATION\_REPROD \) (changes root/shoot ratio before first cutting)

Stress

  • \( H2OREF\_A \) (determines drought resistance)

Nitrogen uptake

  • \( TLIMIT \) (minimum temperature required for nitrogen uptake)
  • \( K\_MM\_NITROGEN\_UPTAKE \) (root affinity to soil nitrogen)

Nitrogen fixation

  • \( INI\_N\_FIX \) (fraction of total nitrogen that might be fixed)
  • \( NFIX\_RATE \) (maximum daily rate of nitrogen fixation)

Respiration

  • \( MAINTENANCE\_TEMP\_REF \) (reference temperature for maintenance respiration)
  • \( MC\_LEAF \) (maintenance respiration coefficient for leafs)
  • \( MC\_ROOT \) (maintenance respiration coefficient for roots)
  • \( MC\_STEM \) (maintenance respiration coefficient for stems)
  • \( MC\_STORAGE \) (maintenance respiration coefficient for fruit/storage organs)
  • \( FYIELD \) (growth respiration efficiency)

Root exudation

  • \( DOC\_RESP\_RATIO \) (ratio of doc exudation in relation to root respiration)

Senescence

  • \( SENESCENCE\_DROUGHT \) (coefficient of senescence related to drought)
  • \( SENESCENCE\_FROST \) (coefficient of senescence related to frost)
  • \( SENESCENCE\_HEAT \) (coefficient of senescence related to heat)
  • \( SENESCENCE\_AGE \) (coefficient of senescence related to age)
  • \( FRET\_N \)

Fineroots turnover

  • \( TOFRTBAS \)

water demand

  • \( WUECMAX \) (water use efficiency)

Structure

  • \( EXP\_ROOT\_DISTRIBUTION \) (coefficient for exponential root distribution)
  • \( RS\_CONDUCT \) (conductiviy of root aerenchyma)
  • \( HEIGHT\_MAX \) (maximum plant height)

Model options

Available options: Default options are marked with bold letters.

  • Unlimited nitrogen availability (npot: no / yes)
  • Transpiration (transpirationmethod: wateruseefficiency / stomatalconductance)
  • Droughtstress (droughtstressmethod: uniform / rootweighted)
  • Considered species (plantfamilies: crops grass)

Management

Considered field mangement includes:

  • Planting
  • Harvest
  • Grazing
  • Cutting

Planting event

For planting events, the following event inputs are considered:

  • Plant type and name
  • Initial biomass

All other quantities are determined by the model:

  • Annual nitrogen fixation is assumed to be zero at planting
  • Plant development index and growing degree days are set to 0
  • Tissue nitrogen concentration is set to optimum

The N-contents of fine roots, foliage, and structural tissue are set to optimum (parameters NC_FINEROOTS_MAX, NC_FOLIAGE_MAX, and NC_STRUCTURAL_TISSUE_MAX). Mass is only distributed to fine roots and foliage by FRACTION_ROOT and 1 - FRACTION_ROOT.

Harvest event

For harvest events, the following event inputs are considered:

  • Export root wood
  • Remains
  • Stubble height

The term wood with regard to the export of roots is neglected and all root parts are considered. The fraction given as remains determines the fraction of straw that remains on the field. If there is not remains fraction given, the amount of straw can be determined via stubble height. If neither remains nor stubble height are given all aboveground biomass is removed from the field.

Grazing

After grazing the development index of the plant is set to 0.

Cutting

After cutting the development index of the plant is set to 0.

Phenology

Phenology of plant growth depends on the plant development stage \( DVS \), which is defined between 0 (germination) and 1 (maturity).

Growing degree days

Plant development depends on accumulated growing degree days \( AGDD \), which is the sum of growing degree days over the complete vegetation period:

\[ AGDD = \sum GDD \]

Growing degree days depend on daily mean temperature and a species-specific base temperature:

\[ GDD = (T_{avg} - GDD\_BASE\_TEMPERATURE) \; f_{chill} \]

The factor \(f_{chill} \) retards plant development due to insufficient vernalization (see: vernalization).

Plant development \( \frac{d DVS}{dt} \) is given by:

\[ \frac{d DVS}{dt} = \frac{GDD}{GDD\_MATURITY} \]

In addition to \( DVS \), there exists a mortality state index \( MOS \) that is calculated in the same way as \( DVS \) but interpreted differently and not reset after grazing and cutting events.

Emergence

Emergence is regulate by accumulated growing degree days \( AGDD \), drought stress and snow. Three conditions must be satisfied for emergence:

\[ AGDD > GDD\_EMERGENCE \\ f_h2o > H2OREF\_FLUSHING \\ snow < 0.01 [m] \]

In case GDD_EMERGENCE is not defined, the plant development index must be greater 5%

Vernalization

Vernalization is only implemented for crops. The following species-specific parameters determine vernalization:

  • \( CHILL\_UNITS \)
  • \( CHILL\_TEMP\_MAX \)
  • \( GDD\_FLOWERING \)

The state of chilling if calculated following [27] (see: Vernalization).

Allocation

Allocation of assimilated carbon and nitrogen is determined by the plant development stage \( DVS \). PlaMox distinguishes the following compartments:

  • Fruit / Reserves
  • Roots
  • Stem
  • Leaves

The plant growth is reduced by a deficient N-content of the leaves, which limits photosynthesis. Then, also the C to N ratio increases.

Crops


The fruit fraction \( \theta_{fruit} \) is given by:

\[ \theta_{fruit} = \left\{\begin{array}{cc} 0 & AGDD \le GDD\_GRAIN\_FILLING \\ FRACTION\_FRUIT \cdot \frac{AGDD - GDD\_GRAIN\_FILLING}{GDD\_MATURITY - GDD\_GRAIN\_FILLING} & AGDD > GDD\_GRAIN\_FILLING \end{array} \right. \label{eq2} \]

If there was heatstress during the flowering phase, the fruit fraction is reduced by the factor influence_heat_reduction_grainfilling as calculated in Heat Stress Limitation

  • The plant allocation into the fruit is reduced if TMINCRIT() and M_FRUIT_OPT() are set. If temperatures are higher than TMINCRIT(), the grain development capacity (m_fruit_max) is reduced.

m_fruit_max = M_FRUIT_OPT() * (1- influence_heat_reduction_grainfilling)

if no C is being allocated to the fruit

The root fraction \( \theta_{root} \) is constant over time before grain filling starts and then decreases to \( FRACTION\_ROOT \).

\[ \theta_{root} = (1-\theta_{fruit})/(1-FRACTION\_FRUIT) \cdot FRACTION\_ROOT \]

For some species families specific calculations exist:

  • Rice

    \[ \theta_{root} = FRACTION\_ROOT - 0.5 \cdot DVS \cdot FRACTION\_ROOT \]

    According to [64], root fraction of rice declines from about 20% at seedling stage to 10% at maturity, which can be reflected by setting the species-specific parameter \( FRACTION\_ROOT = 0.2 \).
  • Corn

    \[ \theta_{root} = \frac{1-\theta_{fruit}}{1 + \frac{1}{RS}} \]

    with the root-shoot ratio \( RS \) given by :

    \[ RS = 0.45 \cdot \left (0.15 + 0.5 \cdot e^{-3 \cdot DVS} \right) \]

  • Rapeseed

    \[ \theta_{root} = \frac{1-\theta_{fruit}}{1 + \frac{1}{RS}} \]

    with the root-shoot ratio \( RS \) given by :

    \[ RS = 0.3 - 0.22 \cdot DVS \]

  • Milt
  • Sorg
    • Wheat

      \[ target_{root} = 1 - DVS \cdot 0.5 + DVS \cdot FRACTION\_ROOT \]

      \[ value_{root} = \frac{fineroots}{total\_biomass} \]

      If the target biomass is larger than the current value,

      \[ \theta_{root} = target_{root} \]

Stem and leaf fraction are given by:

  • If the number of growing degree days is larger than GDD_STEM_ELONGATION = 0 (for crops), no C is allocated to stems, but only to leaves.
  • Otherwise, and in case of drought, no C is allocated to leaves. Instead it goes into stems.
  • Otherwise, and without drought, it is

    \[ \theta_{stem} = (1 - \theta_{fruit} - \theta_{root}) \cdot (1-FALEAF)\\ \theta_{leaf} = 1 - \theta_{fruit} - \theta_{root} - \theta_{stem} \]

    where \(FALEAF\) is the fraction of straw (leaves + stems) forming leaves.

Grass


The reserve/fruit fraction ( \( \theta_{fruit}\)) increases linearly with the plant development in accordance to \( FRACTION\_FRUIT\):

\[ \theta_{fruit} = DVS \cdot FRACTION\_FRUIT \]

A cutting event influences the root/shoot ratio by a factor \( \gamma_{roots}\) (here determined by the fraction of roots \( \theta_{roots}\)):

  • Before the first cutting of the year:
    \( \gamma_{roots} = \frac{1.0}{1.0 \; + \; SHOOT\_STIMULATION\_REPROD} \), with \(SHOOT\_STIMULATION\_REPROD\) = 0

  • After the first cutting of the year:
    \( \gamma_{roots} = 1.0 \)

The root fraction is given by:

\[ \theta_{roots} = (1.0 - \theta_{fruit}) \frac{\gamma_{roots} \; FRACTION\_ROOT}{1 - FRACTION\_FRUIT - (1- \gamma_{roots}) \; FRACTION\_ROOT} \]

As a default or after the first cutting of the year it is

\[ \theta_{roots} = \frac{1.0 - \theta_{fruit}}{1 - FRACTION\_FRUIT} \cdot FRACTION\_ROOT \, . \]

If the current root mass is higher than predicted by the allocation factor, the root allocation factor is decreased exponentially.

The straw fraction is given by

\[ FRACTION\_STRAW = 1 - \theta_{roots} - \theta_{fruit} \, . \]

The current foliage to straw ratio is given by

\[ faleaf = \frac{m_{fol}}{m_{fol}+m_{stem}} \, . \]

If the foliage biomass, \( m_{fol}\), is 0, \( faleaf = 0 \).

Redistribution

If some reserves exist in the stem and after grain filling, some biomass from the stem goes into the grain (reproductive tissue).

Photosynthesis

Actual photosynthesis is calculated by the external model PhotoFarquhar (Berry Ball). This requires the canopy height specific information of:

  • Rubisco activity
  • Electron transport
  • Photorespiration

The latter two quantities are calculated depending on the rusbisco activity and with the species specific parameters QJVC (Maximum electron transport rate and RubP saturated rate of carboxylation) and QRD25, respectively.

The rubisco activity depends on the species specific parameter VCMAX25 (Maximum RubP saturated rate of carboxylation at 25oC for sun leaves). Further, the following properties are factored in:

  • Severe drought stress reduces enzyme activity
  • Plant age reduces enzyme activity
  • Temperature: Heat and frost stress
  • Nitrogen availability

Nitrogen uptake

Nitrogen availability can be dependent on location specific N-distribution. Only the share \( \phi_L \) of total N that is either located close to the plant or that is homogenously distributed is available.

Daily nitrogen demand is calculated by:
n_opt() \( - \) total_nitrogen()

Nitrogen uptake is calculated for every layer individually. Only layers containing roots are considered.

Temperature dependency of N uptake is given by:

\[ \phi_T = \begin{cases} & 0, \;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\ T < 0.8 \cdot TLIMIT \\ & \frac{t - 0.8 \cdot TLIMIT}{TLIMIT - 0.8 \cdot TLIMIT}, \ 0.8 \cdot TLIMIT < T < TLIMIT \\ & 1, \;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\;\ T > TLIMIT \\ \end{cases} \]

Nitrogen uptake of \( N_x \) is given by;

\[ \frac{dN_x}{dt} = US\_NH4 \cdot N_x \cdot m_{roots} \cdot \phi_L \cdot \phi_T \ \frac{N_x}{N_x + K\_MM\_NITROGEN\_UPTAKE} \]

Nitrogen fixation

Two different approaches are considered:

  • Nitrogen fixation based on potential nitrogen fixation rate under consideration of water availability, temperature and the nodule surface. This approach is chosen as soon as: \( NFIX\_RATE > 0.0\).
  • Nitrogen fixation based on total fixable nitrogen (TFN) amount and plant development. This approach is chosen as soon as: \( NFIX\_RATE = 0.0\) and \( INI\_N\_FIX > 0.0\).

Approach \( NFIX\_RATE > 0.0\)

  • Plant N demand:
    • \( \text{N}_{\text{demand}} = \text{N}_{\text{optimum}} - \text{N}_{\text{plant}} \)
      See also
      n_opt()
  • Water availability [50]
    • \( f_{w} = \) NitrogenFixation::get_fact_water()
  • Temperature
    • \( f_{t} = \) NitrogenFixation::get_fact_temperature()
  • Nitrogen availability
    • Hurley Pasture Model (Thornley, 1998)
    • \( f_{n} = \frac{1}{1 + \frac{\text{Fine root nitrogen concentration}}{0.01}} \)
  • Nodule surface: (Weisz et al., 1985)
    • Nodule surface area represented by vegetative plant biomass
  • Actual nitrogen fixation rate:
    • If the plant N demand is \( > 0 \):
      • \( f = \text{Potential N fixation rate} * \text{Nodule surface area} * \text{Water availability} * \text{Temperature} * \text{Nitrogen availability} \)
      • the carbon costs of the nitrogen fixation are restricted by the fine root biomass and are determined:
        • MIN(Actual fixation rate \( * \) NFIX_CEFF, 0.99 \( * \) Fine root biomass carbon content)

Approach \( INI\_N\_FIX > 0.0\)

  • Plant N demand:
    • \( \text{N}_{\text{demand}} = \text{N}_{\text{optimum}} - \text{N}_{\text{plant}} \)
      See also
      n_opt()
  • Total nitrogen demand:
    • \( f_{n} = \) Foliage biomass under optimal, closed canopy conditions (parameter MFOLOPT) \( * \) optimum nitrogen concentration of foliage (parameter NC_FOLIAGE_MAX)
  • Potential N fixation rate:
    • \( f_{pot} = \) MIN(plant development stage \( * \) parameter INI_N_FIX (0-10) \( * \) total foliage nitrogen demand \( - \) the yearly nitrogen fixation, \( \text{N}_{\text{demand}} \))
  • Fine root carbon costs:
    • \( f_{pot} * \) carbon use efficiency for nitrogen fixation (parameter NFIX_CEFF)
  • Biological nitrogen fixation (BNF) happens if the potential N fixation rate \( > 0 \) AND fine root carbon costs (total transport and uptake respiration) \( < \) total fine root carbon

Respiration

Maintenance respiration is calculated after [52] :

  • Maintenance respiration coefficient of leaves, roots, stems and storage organs (parameter MC_LEAF, MC_ROOT, MC_STEM and MC_STORAGE (the latter three are by default chosen differently to [52] ) are used.
  • If the plant is in chilling stadium (get_chill_factor()) there is no maintenance respiration.

  • Maintenance respiration is based on the cost of metabolic activity (lower temperature, higher age of plant (latter not for grass)):
    • Temperature scale factor:
      • \( f_{ts} = \) get_frost_factor() * (2.0^{((T - Reference temperature for maintenance respiration)/10)} * 1/24)
    • Temperature chill factor:
      • \( f_{tc} = f_{ts} * \text{Carbon Content} * \) get_chill_factor()
    • Reduction with age:
      Maintenance respiration is reduced to 50% of daily photosynthesis as long as plant has only little foliage biomass
      (guarantees accrue of plant growth)
      • \( f_{a} = \) MIN((1.0 - dvsFlush * 0.5), get_age_factor()) for crops
      • \( f_{a} = 1 \) for grass species
    • Respiration is calculated for all compartments seperately and then added up to the complete maintenance respiration:
      • \( r_{fol} = MC\_LEAF \cdot \text{Foliage Biomass} * f_{a} * f_{tc} \)
      • \( r_{root} = MC\_ROOT \cdot \text{Fine Root Biomass} * f_{a} * f_{tc} \)
      • \( r_{stem} = MC\_STEM \cdot \text{Sapwood Biomass} * f_{a} * f_{tc} \)
      • \( r_{storage} = MC\_STORAGE \cdot \text{Bud Biomass} * f_{tc} \)

Growth respiration:

  • The fraction of growth respiration (respired C) relative to gross assimilation (assimilated C) \(FYIELD\) is used to calculate the respired C from the uptaken carbon. The assimilated part goes into biomass, the respired C is used energetically for building this biomass.

    \[ C_{resp} = \frac{\text{FYIELD}}{1.0 - \text{FYIELD}} * carbonuptake \]

  • The separate growth respiration rates for foliage, sapwood, storage organs (buds), and fine roots are calculated by multiplying \( C_{resp} \) with the (static?) fractions of foliage, sapwood, buds, or fine roots.
  • A maximum of 90% of the biomass of every compartment is allowed for growth respiration.
    Respiration of storage organs is only added if the plant species is a tuber plant (parameter TUBER), which means it has belowground storage organs, otherwise this factor is 0.

Root exudation

  • Exudation is modelled as a loss of fine root biomass. The exudation losses are calculated using the parameter DOC_RESP_RATIO, which gives the ratio between root exudates and losses from root respiration. The total exudation losses are calculated by
    \( C_{loss} = DOC\_RESP\_RATIO * \text{Belowground Respiration} \) .
  • A maximum of 10% of living root biomass is allowed to be used for exudation:
    \( C_{maxloss} = 0.1 * \text{Fine Root Biomass} * \text{Carbon Content} \)

Senescence

Senescence calculates fluxes from living to dead plant tissue separately for above- and belowground plant parts.

Aboveground senescence

List of senescence processes affecting aboveground tissue:

  • Drought
  • Frost
  • Heat
  • Age

Drought stress

\[ \Phi_d = SENESCENCE\_DROUGHT \cdot \phi_d \]

The drought stress factor \( \phi_d \) is given by: Linear relationship

Frost stress

Frost stress for \( T < 0 \) is given by:

\[ \Phi_f = \begin{cases} & 0, T >= 0 \\ & SENESCENCE\_FROST \cdot \frac{T}{-20}, -20 < T < 0 \\ & SENESCENCE\_FROST, T <= -20 \\ \end{cases} \]

Heat stress

\[ \Phi_h = SENESCENCE\_HEAT \cdot (1 - \phi_{h}) \]

The heat stress factor \( \phi_h \) is given by: Heat factor

Senescence due to age

Grass:

\[ \Phi_{a,leaf} = SENESCENCE\_AGE \cdot DVS \\ \Phi_{a,stem} = SENESCENCE\_AGE \cdot DVS \]

Crops:

\[ \Phi_{a,leaf} = SENESCENCE\_AGE \cdot \frac{GDD - GDD\_GRAIN\_FILLING}{GDD\_MATURITY - GDD\_GRAIN\_FILLING} \\ \Phi_{a,stem} = 0.0 \]

Belowground senescence

List of senescence processes affecting belowground tissue:

  • Drought (considering every soil layer)
  • Frost (considering every soil layer)
  • Age (for non-grass), temperature (for grass), (single value for the whole root system)

Drought stress

\[ \Phi_d = SENESCENCE\_DROUGHT \cdot \phi_d \]

Frost stress

Frost stress for \( T < 0 \) is given by:

\[ \Phi_f = \begin{cases} & 0, T >= 0 \\ & SENESCENCE\_FROST \cdot \frac{T}{-20}, -20 < T < 0 \\ & SENESCENCE\_FROST, T <= -20 \\ \end{cases} \]

Age or temperature stress

Age or temperature stress

  • Age for non-grass: Similar to age for above ground biomass, the parameter is TOFRTBAS
  • T for grass, the parameter is TOFRTBAS

Transpiration

Calculates potential transpiration on
a) water use efficiency and carbon uptake:

b) stomatal conductance and vapour pressure deficit:

If an hourly timestep is chosen, this is done hourly. However, only the accumulated potential transpiration is stored.

Root Structure

Distribution

Roots are represented in a one-dimensional way by the fine root mass distribution and the total fine root mass.

Currently available distribution functions for vertical root distribution:

The environmental function is used for \( ROOTS\_ENVIRONMENTAL = true\). If \( ROOTS\_ENVIRONMENTAL = false\), the exponential function is used for \( EXP\_ROOT\_DISTRIBUTION > 0\). If \( ROOTS\_ENVIRONMENTAL = false\) and \( EXP\_ROOT\_DISTRIBUTION <= 0\), the sigmoid function is used.

Environmentally/dynamic determined root growth

See Sink-strength driven root distribution.

Default static root growth

See Empirical root growth distribution

Root conductivity

Gaseous conductivity of roots is expressed by an root aerenchyme transport \( r_{tc}\) coefficient:

\[ r_{tc} = m_r \cdot RS\_CONDUCT \]

Ground coverage

Ground coverage of grass is always assumed to be 100%

Ground coverage of crops is estimated by lai:

\[ gc = \frac{lai}{3}^{0.5} \]

Full cover is reached with a leaf area index of three (FAO).

Specific leaf area weight (sla)

Calculates specific leaf area weight sla kg m-2 in each canopy layer:

  • sla is assumed to be homogeneously distributed throughout the whole canopy.
  • sla decreases with plant development dvs depending on the species parameter SLADECLINE (mostly 0 or 0.5)

    \[ sla = SLAMAX \cdot ( 1 - dvs \cdot SLADECLINE \cdot dvsMort) \]

    For selected species (mungbean, rice, grass), specific formulations exist.

Common

Heat Stress Limitation

If the plant experiences heat stress during the critical time around flowering, the pod set is reduced. The approach followes the ones introduced by Challinor et al. 2005 and Nendel 2011.

The relevant temperature for this heat stress factor is the temperature during the photoactive period ( \( T_{d}\) ), since it affects the time during which flowers are open.

\[ T_d = T_{max} - \frac{T_{max} - T_{min}}{4} \]

(following Mirschel & Wenke (2007)).

Challinor et al. 2005 introduced a variable Temperature threshold \( T_{crit} \) dependent on timing and duration of the heat stress during the flowering period.

The daily influence ( \( heat_{d}\) ) of the heat limitation is calculated dependent on the daily fraction of flowers open:

\[ heat_{daily} = 1 - (\frac{(T_d - T_{crit})}{(T_{zero} - T_{crit})}) * frac\_flower; \]

The daily fraction of flowers newly opened:

\[ frac\_flower = openFlowers_{today} - openFlowers_{yesterday} \]

The open Flowers on a specific day after flowering (daf) (Moriondo et al. 2011) :

\[ openFlowers = \frac{1}{(1 + \frac{1}{0.015 - 1} * \exp{-1.4 * daf})}; \]

The overall influence on the grain reduction is:

\[ influence\_heat\_reduction\_grainfilling = min(heat_{daily}) \]

Heat factor

Heat factor \( \phi_h \) is given by:

\[ \phi_h = 1 - \frac{1}{1 + e^{-2 (T_{leaf} - PSNTMAX)}} \]

Nitrogen deficiency

Nitrogen deficiency factor \( \phi_n \) is given by:

\[ \phi_n = \frac{c_{N,fol}}{c_{N,fol,opt}}^{N\_DEF\_FACTOR} \]

Aging

The age factor is calculated dependend on the growing degree days (GDD) (see vernalization()), the minimum temperature sum for
foliage activity onset (parameter GDDFOLSTART) and temperature degree days for full plant development .

For grass:

  • If \( \text{GDDFOLSTART} > \text{GDD} \):

    \[ f_{a} = 1.0 - \frac{(\text{GDDFOLSTART} - \text{GDD})}{\text{GDDFOLSTART}} \]

  • else: \( f_{a} = 1.0 \).

For all other species:

  • If \( \text{GDD} < (0.9 * GDD\_MATURITY): f_{a} = 1.0 \)
  • else:

    \[ f_{a} = \text{max}\left(0.0, 1.0 - \frac{\text{GDD} - (0.9 * GDD\_MATURITY)}{GDD\_MATURITY - (0.9 * GDD\_MATURITY)} \right) \]

Optimum nitrogen concentration

Grass: foliage nitrogen concentration is constant (from species parameter NC_FOLIAGE_MAX)
Crops: foliage nitrogen concentration is highest at planting (NC_FOLIAGE_MAX) and decreases until harvest to NC_FOLIAGE_MIN.