Calculate CO2 concentration in the chloroplast or mesophyll
apply_gm.Rd
Calculates CO2 concentration in the chloroplast or mesophyll, the CO2 drawdown across the stomata, and the CO2 drawdown across the mesophyll. This function can accomodate alternative column names for the variables taken from the Licor file in case they change at some point in the future. This function also checks the units of each required column and will produce an error if any units are incorrect.
Usage
apply_gm(
licor_exdf,
photosynthesis_type = 'C3',
calculate_drawdown = TRUE,
a_column_name = 'A',
ca_column_name = 'Ca',
ci_column_name = 'Ci',
gmc_column_name = 'gmc',
total_pressure_column_name = 'total_pressure'
)
Arguments
- licor_exdf
An
exdf
object representing data from a Licor gas exchange measurement system.- photosynthesis_type
A string indicating the type of photosynthesis being considered (either
'C3'
or'C4'
).- calculate_drawdown
A logical value indicating whether to calculate drawdown values.
- a_column_name
The name of the column in
licor_exdf
that contains the net assimilation inmicromol m^(-2) s^(-1)
.- ca_column_name
The name of the column in
licor_exdf
that contains the ambient CO2 concentration in the chamber inmicromol mol^(-1)
.- ci_column_name
The name of the column in
licor_exdf
that contains the intercellular CO2 concentration inmicromol mol^(-1)
.- gmc_column_name
The name of the column in
licor_exdf
that contains the mesophyll conductance to CO2 inmol m^(-2) s^(-1) bar^(-1)
.- total_pressure_column_name
The name of the column in
licor_exdf
that contains the total pressure inbar
.
Details
For a C3 plant, the mesophyll conductance to CO2 (gmc
) is said to be
the conductance satisfying the following one-dimensional flux-conductance
equation:
(1) An = gmc * (PCi - PCc)
where An
is the net CO2 assimilation rate, PCi
is the partial
pressure of CO2 in the intercellular spaces, and PCc
is the partial
pressure of CO2 in the chloroplast. A key underlying assumption for this
equation is that the flow of CO2 has reached a steady state; in this case, the
flow across the stomata is equal to the flow across the mesophyll.
This equation can be rearranged to calculate PCc
:
(2) PCc = PCi - An / gmc
This version of the equation can be found in many places, for example, as Equation 4 in Sharkey et al. "Fitting photosynthetic carbon dioxide response curves for C3 leaves" Plant, Cell & Environment 30, 1035–1040 (2007) [doi:10.1111/j.1365-3040.2007.01710.x ].
It is common to express the partial pressures in microbar
and the
assimilation rate in micromol m^(-2) s^(-1)
; in this case, the units of
mesophyll conductance become mol m^(-2) s^(-1) bar^(-1)
.
Licor measurement systems provide CO2 levels as relative concentrations with
units of parts per million (ppm
), or equivalently,
micromol mol^(-1)
. Concentrations and partial pressures are related by
the total gas pressure according to:
(3) partial_pressure = total_pressure * relative_concentration
Thus, it is also possible to calculate the CO2 concentration in the
choloroplast (Cc
) using the following equation:
(4) Cc = Ci - An / (gmc * P)
where Ci
is the intercellular CO2 concentration and P
is the
total pressure. In this function, Equation (4) is used to calculate Cc
,
where the total pressure is given by the sum of the atmospheric pressure and
the chamber overpressure.
When a plant is photosynthesizing, it draws CO2 into its chloroplasts, and
this flow is driven by a concentration gradient. In other words, as CO2 flows
from the ambient air across the stomata to the intercellular spaces and then
across the mesophyll into the chloroplast, there is a decrease in CO2
concentration at each step. Sometimes it is useful to calculate these changes,
which are usually referred to as "CO2 drawdown" values. So, in addition to
Ci
, this function (optionally) calculates the drawdown of CO2 across
the stomata (drawndown_cs = Ca - Ci
) and the drawdown of CO2 across the
mesophyll (drawdown_cm = Ci - Cc
).
_Note_: mesophyll conductance is not specified in typical Licor files, so it
usually must be added using set_variable
before calling
apply_gm
.
For a C4 plant, mesophyll conductance instead refers to the conductance
associated with the flow of CO2 from the intercellular spaces into the
mesophyll (rather than into the chloroplast). In this case, the equations
above just require a small modification where Pcc
and Cc
are
replaced by PCm
and Cm
, the partial pressure and concentration
of CO2 in the mesophyll.
Value
An exdf
object based on licor_exdf
that includes the following
additional columns, calculated as described above: Pci
and Ci
(for C3 plants) or PCm
and Cm
(for C4 plants),
drawndown_s
, and drawdown_cm
. The category for each of these new
columns is apply_gm
to indicate that they were created using this
function.
Examples
# Read an example Licor file included in the PhotoGEA package
licor_file <- read_gasex_file(
PhotoGEA_example_file_path('ball_berry_1.xlsx')
)
# Calculate the total pressure in the Licor chamber
licor_file <- calculate_total_pressure(licor_file)
# Set the mesophyll conductance to 1 mol / m^2 / s / bar
licor_file <-
set_variable(licor_file, 'gmc', 'mol m^(-2) s^(-1) bar^(-1)', value = 1.0)
# Calculate Cc and drawdowns
licor_file <- apply_gm(licor_file)
licor_file$units$Cc # View the units of the new `Cc` column
#> [1] "micromol mol^(-1)"
licor_file$categories$Cc # View the category of the new `Cc` column
#> [1] "apply_gm"
licor_file[, 'Cc'] # View the values of the new `Cc` column
#> [1] 216.5128 257.7110 231.1340 208.2051 261.8719 286.6935 346.7131 266.8860
#> [9] 276.4449 273.5312 316.1661 287.6186 307.7915 340.4040 236.9847 283.9512
#> [17] 293.8102 284.9079 322.5466 242.3728 323.0196 206.8106 219.7376 236.6592
#> [25] 270.8341 306.0797 299.8268 247.3581