Topic 2

Plant and Canopy Ecophysiology:

Photosynthesis and Transpiration
‣ Readings: Bonan Ch.15, 16, 17.2−17.4
Limitations of Biogeographical Approach
‣ What are the limitations of using a biogeographical approach to
predict future vegetation patterns in response to climate
change? There are many…
‣ One is that it assumes a unidirectional (and causal) relationship
between climate and vegetation, but in reality the relationship is
bidirectional - climate also responds to vegetation changes.
‣ Climate and vegetation essentially coevolve with each other.
‣ So how do plants affect climate?

Climate change Climate change Climate change

Vegetation change Vegetation change Vegetation change

 Photosynthesis
‣ Photosynthesis: CO2 + H2O + hν → CH2O + O2
‣ Oxidation no. of C changes from +4 to 0, i.e., C is reduced.
Atmospheric CO2 is “fixed” by autotrophs, i.e., photosynthetic
land plants and marine phytoplankton (roughly half by each).
Ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO)
is the primary enzyme involved.
‣ Carbon is converted to a variety of
products (carbohydrates, lipids,
proteins, etc.). “CH2O” correctly
accounts for the stoichiometry and
oxidation no. of the composite of
these products on average.
‣ Energy from sunlight is
converted and stored in reduced
carbon compounds (mostly as
carbohydrates and lipids).
[Mayer]
 Photosynthesis in Land Plants
Photosynthesis: absorbs atmospheric
CO2 and combines it with H2O to make
food → cooling effect

Stoma
Stoma

CO2 + H2O + hν
→ CH2O + O2

‣ http://newswatch.nationalgeographic.com/2013/
08/14/mesmerizing-gifs-of-breathing-earth [Mayer]
Photosynthesis: Light Reaction
In plants,
photosynthesis
takes place in
specialized
organelles called
chloroplasts.
Light-dependent reaction occurs on
the thylakoid membrane. Photons from
sunlight are absorbed mostly by a
pigment called chlorophyll. Accessory
pigments such as carotene help
absorb a wider spectrum of sunlight. A
cluster of pigment molecules together
form a photosystem (PS).
Plants can only make use of radiation
of wavelengths between 0.4–0.7 μm,
known as photosynthetically active
radiation (PAR).
 Visible vs. Near-Infrared Radiation
‣ Green leaves typically absorb >85% of visible solar radiation (i.e.,
PAR) for photosynthesis, but <50% of near-infrared to avoid
overheating. Other surfaces have smaller spectral differences.
‣ Normalized difference vegetation index (NDVI), ranging from −1
to 1, is developed as a measure of “greenness”:
rnir = reflectance in near-IR
rred = reflectance in red wavelengths

Annual average NDVI for

1982−1993 (×0.1)
[Bonan Fig. 24.5]
 Photosynthesis: Light Reaction
Photosystem II
(PSII/P680) is excited
and loses electrons
upon receiving photons
from sunlight. Lost
electrons are replaced
by splitting H2O,
producing O2 and
releasing H+ into the
thylakoid lumen.
The excited electrons go through an electron transport chain (ETC) from
PSII and PSI (P700), and along the way energy is released to pump H+
from the stroma into thylakoid lumen, building a proton gradient across
the thylakoid membrane. The electrons received by PSI are excited again
by the absorption of photons and finally used to reduce NADP+ to
NADPH. As protons flow down the gradient back to the stroma via the
ATP synthase complex, electrochemical energy is released to synthesize
ATP in a process known as photophosphorylation.
Photosynthesis: Light Reaction
Overall:
A total of 8−9 photons must be
absorbed to yield 2 NADPH and
3 ATP to reduce 1 CO2 molecule
in the Calvin Cycle.

[Bonan]
 Photosynthesis: Dark Reaction
RuBP C3 plants Enzyme Rubisco combines
CO2 and RuBP to form two
molecules of 3-PGA, which
is then reduced to G3P (or
GAP).
3-PGA Some (1/6) of G3P is used to
produce carbohydrates and
other metabolic products; the
remainder (5/6) is used to
regenerate RuBP.

G3P / GAP Overall:

ATP and NADPH are used
Calvin
to convert CO2 to reduced
Cycle
organic compounds (e.g.,
glucose):
Other metabolic nCO2 + 2nH2O + hν
processes
→ (CH2O)n + nO2 + nH2O
Photosynthesis: Dark Reaction

Overall:
 Photorespiration and C4 Plants
C4 plants

low CO2:O2

Rubisco can also

catalyze the oxidation
of RuBP by O2 (photorespiration),
greatly reducing net carbon uptake.
C4 plants have a spatial separation
to create a high CO2:O2 ratio at the
site of Calvin Cycle. In the mesophyll
cells, CO2 is assimilated into a 4-C
compound first, which is then
transformed and transported to the high CO2:O2
bundle sheath cells, releasing CO2
there for Calvin Cycle CO2 fixation.
 C3 vs. C4 Plants
C4 pathway greatly reduces C4 plants
photorespiration, increases net
photosynthetic rate and improves low CO2:O2
water-use efficiency, especially at
high light levels and warm
temperatures.
C4 plants (usually grasses) live well in
a relatively warm climate with periodic
droughts, e.g., tropical savanna.

high CO2:O2
 Crassulacean Acid Metabolism
‣ In desert environments, some plants utilize the Crassulacean Acid
Metabolism (CAM) photosynthetic pathway, which represents a
temporal separation between initial CO2 assimilation and Calvin Cycle.

[Garcia et
al., 2014]
Photosynthetic Characteristics of Different Plants
Environmental Controls of Photosynthesis
‣ Photosynthesis rate Jack pine [Bonan Fig. 16.3]
(A) can be limited by
and depends on a
variety of factors, e.g.:

Ti = leaf intercellular
temperature
ci = leaf intercellular CO2
concentration
PPFD = photosynthetic
photon flux density

‣ Dependence on water
is generally coupled to
stomatal conductance
and evapotranspiration.
 Limitations for Photosynthesis
RuBP-limited: RuBP Rubisco-limited:
• Regeneration of • CO2 availability
RuBP • Kinetic properties
• Controlled by and amount of
availability of Rubisco
photons to
produce ATP 3-PGA
and NADPH

G3P / GAP
Product-limited:
• Utilization of G3P in
carbohydrate ATP and NADPH
synthesis, which are produced by the
releases phosphate electron transport
Other metabolic
for further ATP processes chain in the light-
synthesis dependent reaction
 Modeling Photosynthesis
‣ Farquhar et al. [1980] pioneered a simplified mathematical model for
photosynthesis. But he also said, “Unfortunately, the complexity of
photosynthesis means that analytical descriptions can only be
achieved at the expense of gross simplification… (these models)
can be useful aids to understanding, and for prediction, but are also
potential hazards when the simplifications involved are forgotten.”
‣ Rubisco-limited rate (Ac, μmol CO2 m−2 s−1) is derived from the
Michaelis-Menton model for fast-equilibrium enzyme kinetics:

k1 k3
E + S ⇌ ES → E + P
k2
• Vcmax = maximum rate of carboxylation (μmol m−2 s−1) M-M constant: Km = k2/k1
• ci = intercellular CO2 concentration (μmol mol−1) Lower Km, higher enzyme
• Γ∗ = CO2 compensation point where photosynthesis (E) affinity for substrate (S)
balances leaf respiration (μmol mol )−1
and faster formation of
• Kc = M-M constant for carboxylation (μmol mol ) −1
product (P)
• Ko = M-M constant for oxygenation (mmol mol ) −1

• oi = O2 concentration = 209 mmol mol−1 (Monson & Baldocchi Ch.3, 5)

 Modeling Photosynthesis
‣ RuBP-limited rate (Aj, μmol CO2 m−2 s−1) is dependent on the electron
transport rate (J, μmol m−2 s−1). First, we calculate the light utilized in
electron transport by photosystem II (IPSII, μmol photon m−2 s−1):

• I = photosynthetically active radiation (PAR) incident

on a leaf (μmol photon m−2 s−1)
• αl = leaf absorptance (fraction) ≈ 0.8
• ΦPSII = quantum yield of PS II (mol mol-1) ≈ 0.85
‣ Then we solve an empirically derived quadratic function and choose the
smaller of the two roots for J:
• Jmax = maximum potential electron
transport rate (μmol m−2 s−1)
• ΘJ = curvature parameter ≈ 0.7−0.9
‣ Finally, based on stoichiometry of carboxylation, oxygenation and
various products and reactants in the Calvin Cycle, the RuBP-limited rate
is approximated to be:
 Modeling Photosynthesis
‣ Product-limited rate (Ap, μmol CO2 m−2 s−1) is simply:
Tp = rate of G3P (triose phosphate) utilization (μmol m−2 s−1)

‣ Net photosynthetic rate (An, μmol CO2 m−2 s−1) is:

Rd = leaf mitochondrial respiration rate

[Bonan, Fig. 16.4]

 Modeling Photosynthesis
‣ Vcmax, Jmax, Tp and Rd all vary with leaf temperature.
‣ Vcmax and Jmax are proportional to amount of Rubisco and chlorophyll,
respectively, both dependent on foliage nitrogen content.
‣ Observations from a wide range of plants show an optimal allocation
of nitrogen to balance enzymatic (Rubisco) and light-harvesting
(chlorophyll) capabilities.
‣ Jmax, Tp and Rd can all be scaled to
Vcmax, e.g., Jmax = 1.67Vcmax at 25°C, but
Vcmax itself is poorly constrained.

[Bonan, Fig. 16.5]

 Exercise 2.1
‣ Calculate the light-saturated photosynthesis rate for a
C3 plant with Vcmax = 60 μmol m−2 s−1 and Jmax = 100
μmol m−2 s−1. Kc = 404.9 μmol mol−1, Ko = 278.4 mmol
mol−1, and Γ∗ = 42.75 μmol mol−1 for a leaf temperature
Tl = 25°C. Use ci = 0.7ca, oi = 209 mmol mol−1, and ca =
380 μmol mol−1. ΘJ = 0.9, ΦPSII = 0.85, and αl = 0.8.
Photosynthesis and Plant-Atmosphere Fluxes
‣ Photosynthesis is the central
process tightly linked to the
exchange of heat, water, CO2
and other gases between
plants and the atmosphere.
‣ These exchange processes
are generally represented as
fluxes.
 Flux as a Unifying Concept
‣ “Flux” is a unifying concept for surface-atmosphere exchange.
‣ Flux is formally termed flux density, which is flow (of mass,
heat or momentum) per unit area per unit time, and is
equivalent to velocity multiplied by density (of mass, heat or
momentum).
‣ Net flux occurs in the direction opposite to the gradient of its
density. This is best exemplified by the Fick’s first law of
diffusion:
F = mass flux [kg m−2 s−1] ρ = mass density [kg m−3]
D = diffusivity [m2 s−1] x = arbitrary position [m]

Arbitrary area δA

Particles flow from

y high to low density
or concentration.
x
Generalized Flux for Surface-Atmosphere Exchange
‣ Fick’s first law can be generalized for various kinds of biogeochemical
and biogeophysical fluxes involving the atmosphere, in terms of a
vertical difference in a related quantity modified by a conductance:

F = flux of X into the atmosphere

φs = a quantity related to X evaluated at
reference height φa the surface
in atmosphere
F
φa = a quantity related to X evaluated at
r some reference height of the atmosphere
surface (land φs g = total conductance
or ocean) c = dimensionality “constant”

‣ F > 0: net flux from surface to atmosphere

‣ Sensible heat flux: φ = temperature; latent heat flux: φ = water
vapor pressure or specific humidity; chemical flux: φ = concentration
‣ Conductance (g) is a function of various environmental factors
(meteorology, vegetation, land surface characteristics, etc.).
‣ “Constant” c just makes sure the units on both sides match.
‣ Resistance (r) is the reciprocal of conductance: r = 1/g
Resistance Model for Land-Atmosphere Exchange
r = ra + rb + rc
ra = aerodynamic resistance
(turbulent diffusivity, wind,
stability, roughness, …)
rb = boundary-layer
resistance (viscosity,
molecular diffusivity, …)
[Bonan Fig. 15.1]

Vegetated surface
/ canopy

rc = surface/canopy
resistance (surface
characteristics)
[Brasseur & Jacob 2014]
 Heat and Gas Exchange in Leaves
Stomata: pores or
openings on leaves
that allow the
exchange of gases
between plants and
the atmosphere

← O3, SO2
In: CO2, O3, SO2, etc.
Out: H2O, BVOCs
(biogenic volatile
[Sellers et al. 1997] organic compounds)
‣ Stomata open to allow CO2 uptake during
photosynthesis, but close to prevent excess water loss
during transpiration. Stomatal conductance (gs) is a
measure of how open the stomata are, the regulation of
which reflects a compromise between two conflicting
goals: to maximum photosynthetic CO2 gain and to
minimize transpirational water loss.
 Heat and Gas Exchange in Leaves
Stomata: pores or
openings on leaves
that allow the
exchange of gases
between plants and
the atmosphere

← O3, SO2
In: CO2, O3, SO2, etc.
Out: H2O, BVOCs
(biogenic volatile
[Sellers et al. 1997] organic compounds)

‣ Plant-atmosphere exchange of heat, water vapor, CO2 and other

gases can be modeled as a diffusion process modulated by both
boundary-layer conductance (gb) and stomatal conductance (gs).
‣ Sensible heat exchange (heat transfer by conduction and
convection) is controlled by gb alone. Water vapor (i.e., latent heat)
and CO2 exchange is controlled by both gb and gs.
 Leaf Boundary Layer and Stomata
Stomata

Sensible heat flux is closely

related to the leaf-air temperature
difference.

‣ It is common to treat the conductances for heat (e.g., gbh) and those for
water vapor (e.g., gbw) to be the same.
‣ Thus, in what follows, when unspecified the conductances are meant to be
for water vapor. For other gases, appropriate scaling factors are needed.
Heat and Gas Exchange in Leaves

‣ Sensible heat flux occurs on both side of the leaf; the two
resistances (rb) act in parallel.
‣ Stomata are typically located on the lower leaf surface
(hypostomatous). Some stomata are on both sides of the leaf
(amphistomatous), and their resistances (rs) act in parallel.
‣ For H2O and CO2 exchange, rs and rb act in series.
 Boundary-Layer Conductance
‣ Boundary-layer conductance (mol H2O m−2 s−1) can be
parameterized per unit leaf area (one-sided) as a function of leaf
size (l, m) and wind speed (u, m s−1):

[Bonan Fig. 15.2] ‣ Large gb indicates effective

heat exchange between the
leaf and the air.
‣ Larger leaf has a thicker
boundary layer and lower gb;
leaf conditions are decoupled
from air.
‣ Higher wind depletes the leaf
boundary layer, increasing gb;
leaf conditions are strongly
coupled to air.
 Leaf Size and Shape
‣ It is suggested that leaf size is adapted to maximize water-use
efficiency (photosynthetic carbon gain per unit transpirational
water loss) and reflects a compromise between leaf energy
exchange, temperature and photosynthesis.
 Evapotranspiration
‣ Evapotranspiration (ET) is the sum of surface evaporation
(from soils or water bodies) and plant transpiration (transport of
water from plant roots to leaves and release from stomata).
‣ ET depends on available energy (solar
radiation, temperature), air humidity,
wind speed, soil type and moisture, and
plant physiology.

Absorb
latent heat

Liquid Water
Release
water vapor
latent heat
‣ Wet surfaces provide energy to evaporate
water, thus are cooled → cooling effect
‣ When water vapor condenses to form clouds,
it releases latent heat and warms upper air.
 Transpiration in Land Plants
‣ Transpiration accounts for 80–90% of global terrestrial
evapotranspiration. A single well-watered tree can transpire
100–150 L of water every day.
‣ In the soil-plant-atmosphere continuum, water flows down a
gradient of water potential (negative suction).

Stoma
 Transpiration in Land Plants
‣ ET is usually high in the day (highest soil-plant-atmosphere
gradient in water potential), and low at night (~0 gradient
before dawn) when stomata close and plant water uptake
replenishes water depleted during the day.
‣ Predawn foliage water pure water
potential is thus a good
indicator of soil moisture.
‣ As plants extract water
from the soil, some
critical water content is
reached at which further
decrease in soil water
increases plant stress.

[Bonan Fig. 10.4]

Transpiration as a Diffusive Process
‣ Transpiration rate (E) (mol H2O m−2 s−1) ea
can be represented as a diffusive process:

es
• gb and gs = boundary-layer and
stomatal conductance for water vapor
(mol H2O m−2 s−1)
• ea, es and ei = ambient, leaf surface
and intercellular water vapor pressure
(Pa)
• P = surface air pressure
• gl = (1/gb + 1/gs)−1 = total leaf ei
conductance for water vapor
‣ Generally, the intercellular space in leaves is assumed to be
saturated with moisture. Therefore:
which is the saturation vapor pressure according to
Clausius-Clapeyron equation at the leaf temperature Tl
Diffusive Limitations on CO2 Supply
‣ Leaf net photosynthesis (μmol CO2 m−2 s−1)
can be represented as a diffusive process: ca

• gb and gs = boundary-layer and cs

stomatal conductance for water vapor
(mol H2O m−2 s−1)
• ca, cs and ci = ambient, leaf surface
and intercellular CO2 concentration
(μmol mol−1)
• Factors 1.4 and 1.6 account for lower
diffusivity of CO2 compared with H2O ci
• gl = (1.4/gb + 1.6/gs)−1 = total leaf
conductance for CO2
‣ Jarvis [1976] developed an empirical model for gs:
I = PAR; T = temperature
D = vapor pressure deficit
ψ = foliage water potential
Environmental Controls of Stomatal Conductance
(tuned to empirical data) [Bonan,
Fig. 16.6]
Photosynthesis-Transpiration Compromise
‣ Stomata are regulated to
maximize CO2 gain via
photosynthesis and minimize
water loss via transpiration.
‣ An and gs vary in near constant
proportion for a given set of
conditions.
‣ Plants achieve this optimization
by adjusting gs to maintain a
nearly constant ci/ca ratio for
given environmental conditions,
thus, assuming gb ≫ gs:

ci/ca ≈ 0.65−0.80 for C3 plants

[Bonan,
ci/ca ≈ 0.40−0.60 for C4 plants
Fig. 16.7]
Photosynthesis-Stomatal Conductance Model
‣ Ball et al. [1987] developed a stomatal conductance model (Ball-
Berry model) that can be directly coupled to the Farquhar
photosynthesis model:
• hs = fractional humidity at leaf surface (unitless)
• cs = leaf surface CO2 concentration (μmol mol−1)
• g0 = minimum conductance ≈ 0.01 mol H2O m−2 s−1
for C3 plants
• g1 = slope of relationship ≈ 9 for C3 plants
‣ Together with Farquhar model and the diffusion equations, An and
gs can be solved iteratively until ci converges:

An ↑ gs ↑ An ↑
An ↑
ca ↑ Overall
gs ↓ An ↓ gs ↓
Water-Use Efficiency and Stomatal Conductance
‣ Original Ball-Berry model accounts for well-watered conditions only. Some
models multiply g1 or An by a soil water stress function to account for the
effect of low soil moisture or low soil water potential.
‣ Water-use efficiency (WUE) is defined as the ratio of carbon gain during
photosynthesis to water loss during transpiration. Assuming gb ≫ gs:
D = (ei − ea)/P = vapor
pressure deficit
e = water vapor pressure
P = air pressure

‣ It is proposed that plants have evolved such that gs adjusts to maintain

WUE constant over some time period. Formally, this means ∂An/∂E =
constant.
‣ Combining the original Ball-Berry model and WUE optimization, Medlyn et
al. [2011] has derived an alternative function for gs:

cs
Free-Air CO2 Enrichment Experiments

Free-air CO2 enrichment (FACE) experiment

‣ Ainsworth and Long [2005]
summarized results from various
FACE experiments: light-saturated
leaf photosynthesis increased by
31% and gs decreased by 20% on
average when exposed to 475−600
ppm elevated CO2.
‣ Photosynthetic response is greater
for C3 plants than C4 plants. [Bonan, Fig. 16.10]
 CO2 and Stomatal Conductance
‣ A 200-ppm (from ~370 to
~570) enhancement in CO2
leads to 10−40% increase in
net primary production.
‣ Corresponding reduction in
stomatal conductance, and
increase in water-use
efficiency. Recall:
[Bonan,
Fig. 20.23]
‣ Franks et al. [2013] suggest that relative changes in
An and gs at elevated CO2 can be approximated by:

• ca0 = 360 ppm and Γ∗ = 40 ppm

• Assumed: An = RuBP-limited rate; g1 and hs unchanged; g0 ≪ gs
 CO2 and Stomatal Conductance
[Bonan, Fig. 16.9]
Over 400 Over 400
million years million years

Over 400 Over 400 million

million years years + past 200
Over geological years
timescale, plants
evolved in response
to changing CO2 by
optimizing stomatal
density and size to
maintain a nearly
constant ci/ca ratio Best-fit curve for
[Franks et al., 2013].
 Plant Ecophysiology and Climate
‣ As ambient CO2 increases, An increases but gs decreases.
WUE is usually enhanced.
‣ As plants encounter a severe dry condition (high water vapor
deficit), gs is reduced substantially to save water, and An is also
reduced.
‣ Reduced gs decreases transpiration. Since evapotranspiration
is an important way via which heat and water are exchanged
between the land surface and the atmosphere, reduced
transpiration may warm the surface, dry the atmosphere and
thicken the boundary layer → positive feedback
‣ Via regulating both photosynthesis and transpiration, the
physiology of stomata links the biochemistry of plants to land-
atmosphere biogeophysical and biogeochemical fluxes, with
important ramifications for climate and atmospheric
composition.
 Exercise 2.2
‣ Frank et al. [2013] suggest that the net photosynthesis rate (An) at an
elevated CO2 level (ca, ppm), relative to the baseline photosynthesis rate
(An0) can be approximated by:

where ca0 = 360 ppm is the baseline CO2 level, and Γ∗ = 40 ppm is the
CO2 compensation point where photosynthesis balances respiration.
‣ Assuming that a given plant adjusts its stomatal conductance (gs) so
as to maintain a constant ratio of leaf internal to ambient CO2
concentration of ci/ca = 0.7, and that gas exchange is not limited by
leaf boundary resistance (i.e., gb ≫ gs), estimate the relative change
in gs (i.e., calculate gs/gs0) for a doubling of CO2 level (i.e., ca = 720
ppm). Explain how this would affect evapotranspiration and surface
temperature.
Leaf Area Index and Light Penetration
‣ Leaf area index (LAI, m2 m−2) is leaf area (one-sided) per unit area of
ground. It is typically 4−6 m2 m−2 for a productive forest.
‣ As incident radiation
penetrates through a
canopy, it is
attenuated by the
leaves via absorption
or scattering, and thus
irradiance at height
(z) below the canopy
top is a function of
cumulative LAI.

[Bonan, Fig. 17.1]

 Canopy Photosynthesis
‣ Canopy photosynthesis, i.e., gross primary production (GPP), is
the sum of photosynthesis rates of individual leaves over the entire
canopy. It is the CO2 uptake per unit area of land (instead of leaf).
‣ There are many ways to model GPP. At one extreme, we can
express GPP as empirical functions of photosynthetically active
radiation (PAR), temperature (T), soil moisture (θ), vapor pressure
deficit (D), etc., modulated by a light-use efficiency (ϵ ≈ 1.5 gC
MJ−1), as in the production efficiency model: • F = absorbed PAR
• f = empirical functions
scaled from 0 to 1
‣ At the other extreme, we can explicitly calculate light penetration,
photosynthesis and conductance at each canopy layer, and
integrate them over all layers, as in a multilayer canopy model.
‣ In many climate models, canopy fluxes are usually calculated using
a big-leaf approach that uses scaling to account for within-canopy
light attenuation, sunlit vs. shaded leaves, vertical variation in leaf
nitrogen content, etc.
 Canopy Photosynthesis
‣ Under a big-leaf approach, we can apply proper scaling (to
account for light attenuation, leaf nitrogen content variations,
etc.) to calculate canopy-average parameters from leaf-level
parameters, and then use, e.g., Farquhar-Ball-Berry model to
calculate canopy-average photosynthesis rate and stomatal
conductance per unit area of leaf.
‣ GPP (μmol CO2 m−2 s−1, where m−2 refers to a unit land area) is
then:
GPP = 𝐴! 𝐿
• Al = leaf-level photosynthesis rate averaged over canopy (μmol
CO2 m−2 s−1, where m−2 refers to a unit leaf area)
• L = leaf area index (LAI, m2 m−2)

‣ Similarly, leaf-level mitochondrial respiration Rd can be scaled

up to the canopy level by multiplying Rd with L.
 Canopy Conductance
‣ Canopy conductance (gc) (mol H2O m−2 s−1) is the integration of leaf
boundary-layer (gb) and stomatal (gs) conductances over whole canopy,
scaled up by leaf areas:
For water 𝐿
For heat: 𝑔!" = 2𝑔# 𝐿 𝑔! = $%
vapor: 𝑔# + 𝑔&$%

‣ Together with the

aerodynamic conductance (ga)
that accounts for turbulent
transport above the canopy,
canopy conductance controls
the flux of water vapor (latent
heat) and CO2 to/from the
atmosphere, dominantly
shaping weather, atmospheric
composition, climate and
biogeochemical cycles.