Radiation-use efficiency is sensitive to variations in atmospheric $CO_2$ concentrations and equations have been incorporated into SWAT+ to modify the default radiation-use efficiency values in the plant database for climate change studies. The relationship used to adjust the radiation-use efficiency for effects of elevated $CO_2$ is (Stockle et al., 1992):

$RUE=\frac{100*CO_2}{CO_2+exp(r_1-r_2*CO_2)}$ 5:2.1.4

where $RUE$ is the radiation-use efficiency of the plant (kg/ha⋅(MJ/m$^2$)$^{-1}$ or 10$^{-1}$ g/MJ), $CO_2$ is the concentration of carbon dioxide in the atmosphere (ppmv), and $r_1$ and $r_2$ are shape coefficients.

The shape coefficients are calculated by solving equation 5:2.1.4 using two known points ($RUE_{amb}$, $CO_{2amb}$) and (, ):

5:2.1.5

5:2.1.6

where is the first shape coefficient, is the second shape coefficient, is the ambient atmospheric concentration (ppmv), is the radiation-use efficiency of the plant at ambient atmospheric concentration (kg/ha⋅(MJ/m) or 10 g/MJ), is an elevated atmospheric concentration (ppmv), is the radiation-use efficiency of the plant at the elevated atmospheric concentration, , (kg/ha⋅(MJ/m) or 10 g/MJ). Equation 5:2.1.4 was developed when the ambient atmospheric concentration was 330 ppmv and is valid for carbon dioxide concentrations in the range 330-660 ppmv. Even though the ambient atmospheric concentration of carbon dioxide is now higher than 330 ppmv, this value is still used in the calculation. If the concentration used in the simulation is less than 330 ppmv, the model defines RUE = .

Stockle and Kiniry (1990) have shown that a plant’s radiation-use efficiency is affected by vapor pressure deficit. For a plant, a threshold vapor pressure deficit is defined at which the plant’s radiation-use efficiency begins to drop in response to the vapor pressure deficit. The adjusted radiation-use efficiency is calculated:

if 5:2.1.7

if 5:2.1.8

where is the radiation-use efficiency adjusted for vapor pressure deficit (kg/ha⋅(MJ/m) or 10 g/MJ), is the radiation-use efficiency for the plant at a vapor pressure deficit of 1 kPa (kg/ha⋅(MJ/m) or 10 g/MJ), is the rate of decline in radiation-use efficiency per unit increase in vapor pressure deficit (kg/ha⋅(MJ/m)⋅kPa or (10 g/MJ)⋅kPa), is the vapor pressure deficit (kPa), and is the threshold vapor pressure deficit above which a plant will exhibit reduced radiation-use efficiency (kPa). The radiation-use efficiency value reported for the plant in the plant growth database, , or adjusted for elevated carbon dioxide levels (equation 5:2.1.4) is the value used for . The threshold vapor pressure deficit for reduced radiation-use efficiency is assumed to be 1.0 kPa for all plants ().

The radiation-use efficiency is never allowed to fall below 27% of . This minimum value was based on field observations (Kiniry, personal communication, 2001).

Once the potential water uptake has been modified for soil water conditions, the actual amount of water uptake from the soil layer is calculated:

$w_{actualup,ly}=min\lfloor w''_{up,ly},(SW_{ly}-WP_{ly})\rfloor$ 5:2.2.7

where $w_{actualup,ly}$ is the actual water uptake for layer $ly$ (mm H$_2$O), $SW_{ly}$ is the amount of water in the soil layer on a given day (mm H$_2$O), and $WP_{ly}$ is the water content of layer $ly$ at wilting point (mm H$_2$O). The total water uptake for the day is calculated:

$w_{actualup}=\sum^n_{ly=1} w_{actualup,ly}$ 5:2.2.8

where is the total plant water uptake for the day (mm HO), is the actual water uptake for layer (mm HO), and n is the number of layers in the soil profile. The total plant water uptake for the day calculated with equation 5:2.2.8 is also the actual amount of transpiration that occurs on the day.

5:2.2.9

where is the actual amount of transpiration on a given day (mm HO) and is the total plant water uptake for the day (mm HO).

Table 5:2-2: SWAT+ input variables that pertain to plant water uptake.

Plant maturity is reached when the fraction of potential heat units accumulated, $fr_{PHU}$, is equal to 1.00. Once maturity is reached, the plant ceases to transpire and take up water and nutrients. Simulated plant biomass remains stable until the plant is harvested or killed via a management operation.

Table 5:2-1: SWAT+ input variables that pertain to optimal plant growth.

If nitrate levels in the root zone are insufficient to meet the demand of a legume, SWAT+ allows the plant to obtain additional nitrogen through nitrogen fixation. Nitrogen fixation is calculated as a function of soil water, soil nitrate content and growth stage of the plant.

$N_{fix}=N_{demand}*f_{gr}*min(f_{sw},f_{no3},1)$ 5:2.3.9

where $N_{fix}$ is the amount of nitrogen added to the plant biomass by fixation (kg N/ha), $N_{demand}$ is the plant nitrogen demand not met by uptake from the soil (kg N/ha), $f_{gr}$ is the growth stage factor (0.0-1.0), $f_{sw}$ is the soil water factor (0.0-1.0), and $f_{no3}$ is the soil nitrate factor (0.0-1.0). The maximum amount of nitrogen that can be fixed by the plant on a given day is $N_{demand}$.

Growth stage exerts the greatest impact on the ability of the plant to fix nitrogen. The growth stage factor is calculated:

when 5:2.3.10

when 5:2.3.11

when 5:2.3.12

when 5:2.3.13

when 5:2.3.14

where is the growth stage factor and is the fraction of potential heat units accumulated for the plant on a given day in the growing season. The growth stage factor is designed to reflect the buildup and decline of nitrogen fixing bacteria in the plant roots during the growing season.

The soil nitrate factor inhibits nitrogen fixation as the presence of nitrate in the soil goes up. The soil nitrate factor is calculated:

when 5:2.3.15

when 5:2.3.16

when 5:2.3.17

where is the soil nitrate factor and is the nitrate content of the soil profile (kg NO-N/ha).

The soil water factor inhibits nitrogen fixation as the soil dries out. The soil water factor is calculated:

5:2.3.18

where is the soil water factor, is the amount of water in soil profile (mm HO), and is the water content of soil profile at field capacity (mm HO).

Plant nitrogen uptake is controlled by the plant nitrogen equation. The plant nitrogen equation calculates the fraction of nitrogen in the plant biomass as a function of growth stage given optimal growing conditions.

$fr_N=(fr_{N,1}-fr_{N,3})*[1-\frac{fr_{PHU}}{fr_{PHU}+exp(n_1-n_2*fr_{PHU})}]+fr_{N,3}$ 5:2.3.1

where $fr_N$ is the fraction of nitrogen in the plant biomass on a given day, $fr_{N,1}$ is the normal fraction of nitrogen in the plant biomass at emergence, $fr_{N,3}$ is the normal fraction of nitrogen in the plant biomass at maturity, $fr_{PHU}$ is the fraction of potential heat units accumulated for the plant on a given day in the growing season, and $n_1$ and $n_2$ are shape coefficients.

The shape coefficients are calculated by solving equation 5:2.3.1 using two known points ($fr_{N,2}$, $fr_{PHU,50\%}$ ) and ($fr_{N,3}$, $fr_{PHU,100\%}$):

5:2.3.2

5:2.3.3

where is the first shape coefficient, is the second shape coefficient, is the normal fraction of nitrogen in the plant biomass at emergence, is the normal fraction of nitrogen in the plant biomass at 50% maturity, is the normal fraction of nitrogen in the plant biomass at maturity, is the normal fraction of nitrogen in the plant biomass near maturity, is the fraction of potential heat units accumulated for the plant at 50% maturity (=0.5), and is the fraction of potential heat units accumulated for the plant at maturity (=1.0). The normal fraction of nitrogen in the plant biomass near maturity () is used in equation 5:2.3.3 to ensure that the denominator term does not equal 1. The model assumes

To determine the mass of nitrogen that should be stored in the plant biomass on a given day, the nitrogen fraction is multiplied by the total plant biomass:

5:2.3.4

where is the optimal mass of nitrogen stored in plant material for the current growth stage (kg N/ha), is the optimal fraction of nitrogen in the plant biomass for the current growth stage, and is the total plant biomass on a given day (kg ha).

Originally, SWAT+ calculated the plant nitrogen demand for a given day by taking the difference between the nitrogen content of the plant biomass expected for the plant’s growth stage and the actual nitrogen content . This method was found to calculate an excessive nitrogen demand immediately after a cutting (i.e. harvest operation). The equation used to calculate plant nitrogen demand is now

5:2.3.5

where is the potential nitrogen uptake (kg N/ha), is the optimal mass of nitrogen stored in plant material for the current growth stage (kg N/ha), is the actual mass of nitrogen stored in plant material (kg N/ha), is the normal fraction of nitrogen in the plant biomass at maturity, and is the potential increase in total plant biomass on a given day (kg/ha).

The depth distribution of nitrogen uptake is calculated with the function:

5:2.3.6

where is the potential nitrogen uptake from the soil surface to depth (kg N/ha), is the potential nitrogen uptake (kg N/ha), is the nitrogen uptake distribution parameter, is the depth from the soil surface (mm), and is the depth of root development in the soil (mm). Note that equation 5:2.3.6 is similar in form to the depth distribution for water uptake described by equation 5:2.2.1. The potential nitrogen uptake for a soil layer is calculated by solving equation 5:2.3.6 for the depth at the upper and lower boundaries of the soil layer and taking the difference.

5:2.3.7

where is the potential nitrogen uptake for layer (kg N/ha), is the potential nitrogen uptake from the soil surface to the lower boundary of the soil layer (kg N/ha), and is the potential nitrogen uptake from the soil surface to the upper boundary of the soil layer (kg N/ha).

Root density is greatest near the surface, and nitrogen uptake in the upper portion of the soil will be greater than in the lower portion. The depth distribution of nitrogen uptake is controlled by , the nitrogen uptake distribution parameter, a variable users are allowed to adjust. Figure 5:2-4 illustrates nitrogen uptake as a function of depth for four different uptake distribution parameter values.

Nitrogen removed from the soil by plants is taken from the nitrate pool. The importance of the nitrogen uptake distribution parameter lies in its control over the maximum amount of nitrate removed from the upper layers. Because the top 10 mm of the soil profile interacts with surface runoff, the nitrogen uptake distribution parameter will influence the amount of nitrate available for transport in surface runoff. The model allows lower layers in the root zone to fully compensate for lack of nitrate in the upper layers, so there should not be significant changes in nitrogen stress with variation in the value used for .

The actual amount if nitrogen removed from a soil layer is calculated:

5:2.3.8

where is the actual nitrogen uptake for layer (kg N/ha), is the potential nitrogen uptake for layer (kg N/ha), is the nitrogen uptake demand not met by overlying soil layers (kg N/ha), and is the nitrate content of soil layer (kg NO-N/ha).

Plant phosphorus uptake is controlled by the plant phosphorus equation. The plant phosphorus equation calculates the fraction of phosphorus in the plant biomass as a function of growth stage given optimal growing conditions.

5:2.3.19

where is the fraction of phosphorus in the plant biomass on a given day, is the normal fraction of phosphorus in the plant biomass at emergence, is the normal fraction of phosphorus in the plant biomass at maturity, is the fraction of potential heat units accumulated for the plant on a given day in the growing season, and and are shape coefficients.

The shape coefficients are calculated by solving equation 5:2.3.19 using two known points () and ():

For each day of simulation, potential plant growth, i.e. plant growth under ideal growing conditions, is calculated. Ideal growing conditions consist of adequate water and nutrient supply and a favorable climate. Differences in growth between plant species are defined by the parameters contained in the plant growth database.

Plant growth is modeled by simulating leaf area development, light interception and conversion of intercepted light into biomass assuming a plant species-specific radiation-use efficiency.

The amount of daily solar radiation intercepted by the leaf area of the plant is calculated using Beer’s law (Monsi and Saeki, 1953):

$H_{phosyn}=0.5*H_{day}*(1-exp(-k_{\Box}*LAI))$ 5:2.1.1

where $H_{phosyn}$ is the amount of intercepted photosynthetically active radiation on a given day (MJ m$^{-2}$), $H_{day}$ is the incident total solar (MJ m$^{-2}$), $0.5*H_{day}$ is the incident photosynthetically active radiation (MJ m$^{-2}$), $k_{\Box}$ is the light extinction coefficient, and $LAI$ is the leaf area index.

Photosynthetically active radiation is radiation with a wavelength between 400 and 700 mm (McCree, 1972). Direct solar beam radiation contains roughly 45% photosynthetically active radiation while diffuse radiation contains around 60% photosynthetically active radiation (Monteith, 1972; Ross, 1975). The fraction of photosynthetically active radiation will vary from day to day with variation in overcast conditions but studies in Europe and Israel indicate that 50% is a representative mean value (Monteith, 1972; Szeicz, 1974; Stanhill and Fuchs, 1977).

Radiation-use efficiency is the amount of dry biomass produced per unit intercepted solar radiation. The radiation-use efficiency is defined in the plant growth database and is assumed to be independent of the plant’s growth stage. The maximum increase in biomass on a given day that will result from the intercepted photosynthetically active radiation is estimated (Monteith, 1977):

5:2.1.2

where is the potential increase in total plant biomass on a given day (kg/ha), is the radiation-use efficiency of the plant (kg/ha⋅(MJ/m) or 10 g/MJ), and is the amount of intercepted photosynthetically active radiation on a given day (MJ m). Equation 5:2.1.2 assumes that the photosynthetic rate of a canopy is a linear function of radiant energy.

The total biomass on a given day, , is calculated as:

5:2.1.3

where is the total plant biomass on a given day (kg ha), and is the increase in total plant biomass on day (kg/ha).

With annuals and perennials, the plants are able to reach full maturity within a single calendar year. With trees, a number of years are needed for a plant to transition from a seedling to a sapling to a fully-developed tree. The parameters in the plant growth database related to radiation-use efficiency represent the annual growth for a fully-developed tree. The heat units to maturity input in the management file is also used to simulate growth within a single year, defining for trees and perennials the period within a year bounded by the development of buds at the beginning of the annual growing season and the maturation of plant seeds at the end of the growing season.

To simulate the smaller amount of biomass accumulation seen in seedlings/saplings, tree growth within a single year is limited to a fixed amount determined by the age of the tree relative to the number of years for the tree species to reach full development. Parameters in the plant growth database define the total number of years for trees to reach full development as well as the biomass of a fully-developed tree. Until the trees in an HRU reach full development, the amount of biomass they can accumulate in a single year is limited to:

$bio_{annual}=1000*(\frac{yr_{cur}}{yr_{fulldev}})*bio_{fulldev}$ 5:2.1.9

where $bio_{annual}$ is the amount of biomass a tree can accumulate in a single year (kg/ha), is the current age of the tree (years), is the number of years for the tree species to reach full development (years), is the biomass of a fully developed tree stand for the specific tree species (metric tons/ha), and 1000 is a conversion factor.

Once the total growth in biomass in a year, , reaches the annual limit, , no more growth occurs until the next year when a new annual limit is calculated. When a tree stand has reached its biomass limit in a year, the increase in plant biomass for a day,, is set to 0.

SWAT+ monitors plant uptake of nitrogen and phosphorus.

The change in canopy height and leaf area for annuals and perennials through the growing season as modeled by SWAT+ is illustrated using parameters for Alamo Switchgrass in Figures 5:2-1 and 5:2-2.

In the initial period of plant growth, canopy height and leaf area development are controlled by the optimal leaf area development curve:

$fr_{LAImx}=\frac{fr_{PHU}}{fr_{PHU}+exp(\Box_1 - \Box_2 * fr_{PHU})}$ 5:2.1.10

where $fr_{LAImx}$ is the fraction of the plant’s maximum leaf area index corresponding to a given fraction of potential heat units for the plant, $fr_{PHU}$ is the fraction of potential heat units accumulated for the plant on a given day in the growing season, and $\Box_1$and $\Box_2$ are shape coefficients. The fraction of potential heat units accumulated by a given date is calculated:

5:2.1.11

where is the fraction of potential heat units accumulated for the plant on day d in the growing season, is the heat units accumulated on day (heat units), and is the total potential heat units for the plant (heat units).

The shape coefficients are calculated by solving equation 5:2.1.10 using two known points (,) and (,):

5:2.1.12

5:2.1.13

where is the first shape coefficient, is the second shape coefficient, is the fraction of the growing season (i.e. fraction of total potential heat units) corresponding to the 1st point on the optimal leaf area development curve, is the fraction of the maximum plant leaf area index (i.e. fraction of ) corresponding to the 1st point on the optimal leaf area development curve, is the fraction of the growing season corresponding to the 2nd point on the optimal leaf area development curve, and is the fraction of the maximum plant leaf area index corresponding to the 2nd point on the optimal leaf area development curve.

The canopy height on a given day is calculated:

5:2.1.14

where is the canopy height for a given day (m), is the plant’s maximum canopy height (m), and is the fraction of the plant’s maximum leaf area index corresponding to a given fraction of potential heat units for the plant. As can be seen from Figure 5:2-1, once the maximum canopy height is reached, will remain constant until the plant is killed.

For tree stands, the canopy height varies from year to year rather than day to day:

5:2.1.15

where is the canopy height for a given day (m), is the plant’s maximum canopy height (m), is the age of the tree (years), and is the number of years for the tree species to reach full development (years).

The amount of canopy cover is expressed as the leaf area index. For annuals and perennials, the leaf area added on day is calculated:

5:2.1.16

while for trees, the leaf area added on day is calculated:

5:2.1.17

The total leaf area index is calculated:

5:2.1.18

where is the leaf area added on day , and are the leaf area indices for day and respectively, and are the fraction of the plant’s maximum leaf area index calculated with equation 5:2.1.10 for day and , is the maximum leaf area index for the plant, is the age of the tree (years), and is the number of years for the tree species to reach full development (years).

Leaf area index is defined as the area of green leaf per unit area of land (Watson, 1947). As shown in Figure 5:2-2, once the maximum leaf area index is reached, will remain constant until leaf senescence begins to exceed leaf growth. Once leaf senescence becomes the dominant growth process, the leaf area index for annuals and perrenials is calculated:

5:2.1.19

while for trees, the calculation is

5:2.1.20

where is the leaf area index for a given day, is the maximum leaf area index, is the fraction of potential heat units accumulated for the plant on a given day in the growing season, is the fraction of growing season () at which senescence becomes the dominant growth process, is the number of years of development the tree has accrued (years), and is the number of years for the tree species to reach full development (years).

If upper layers in the soil profile do not contain enough water to meet the potential water uptake calculated with equation 5:2.2.2, users may allow lower layers to compensate. The equation used to calculate the adjusted potential water uptake is:

5:2.2.3

where is the adjusted potential water uptake for layer (mm HO), is the potential water uptake for layer calculated with equation 5:2.2.2 (mm HO), is the water uptake demand not met by overlying soil layers (mm HO), and is the plant uptake compensation factor. The plant uptake compensation factor can range from 0.01 to 1.00 and is set by the user. As approaches 1.0, the model allows more of the water uptake demand to be met by lower layers in the soil. As approaches 0.0, the model allows less variation from the depth distribution described by equation 5:2.2.1 to take place.