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where
q
ci
and
q
ce
are the speciic CO
2
concentrations
1
inside the substomatal cavity
and just above the stomate, respectively, and
r
s,c
is the stomatal resistance for CO
2
transport. Note that we take
A
n
positive when the assimilation is positive. In that case
the CO
2
lux
F
c
is negative as it is directed towards the leaf.
2
Despite the similarity in pathways, the resistances for CO
2
transport differ from
those for water vapour transport. The transport at this scale is due to molecular diffu-
sion and the respective diffusion coeficients
D
v
and
D
c
are contained in the resistances:
r
s
=
l
/
D
v
(where
l
is the diffusion path length). According to Graham's law, the molec-
ular diffusion coeficient is inversely proportional to the molecular mass of the gas
under consideration (Willmer and Fricker,
1996
). Thus
DMMD
1
16
=
/
≈
D
c
v
c
v
v
.
and the relationship between the stomatal resistances for CO
2
and water vapour
becomes
r
=1.
r
. Consequently, Eq. (
6.23
) can be written as
3
:
s,c
s
qq
r
qq
r
−
A
=
ρ
ce
ci
n
s,c
(6.24)
−
g
−
( )
=
ρ
ce
ci
=
s
ρ
qq
ce
ci
16
.
16
.
s
For a steady-state situation, the net CO
2
lux entering the leaf is equal to that entering
the mesophyll cells. This means that, referring to
Figure 6.12
, the lux can also be
written as:
qq
−
A
=
ρ
ci
cc
(6.25)
n
r
m
With the use of this expression we can look at the difference between C
3
and C
4
plants. Because C
4
plants use a more eficient enzyme in the irst ixation step they
can have a lower intracell CO
2
concentration
q
cc
than C
3
plants: 4·10
-6
kg kg
-1
rather
than 70·10
-6
kg kg
-1
. At the same time the mesophyll resistance
r
m
is also lower in C
4
plants than in C
3
plants: 60 s m
-1
rather than 140 s m
-1
(Ronda et al.,
2001
). Thus for
a given net uptake of CO
2
, C
4
plants can maintain a lower internal CO
2
concentration
A
because
q
=
n
rq
+
. A lower
q
ci
in turn means that for a given net CO
2
uptake
ci
mcc
ρ
1
Note that in literature CO
2
concentrations are often given as densities (
ρρ
c
=
q
) or as volume fractions
c
M
M
(
f
=
q
, where M
CO2
is the molar mass of CO
2
). Volume fractions are usually given in parts per million by
c
c
CO
2
volume (ppmv). See also
Appendix B
.
2
This sign convention is consistent with the notion that transport occurs generally down the gradient, in this case
of CO
2
.
3
Here we ignore the fact that the diffusion of water vapour and CO
2
will interact, leading to a modiication of
the relationship between net assimilation and CO
2
concentration difference (see Von Caemmerer and Farquhar,
1981
).
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