L. Balaguer et al., PRODUCTION AND UTILIZATION OF ASSIMILATES IN WHEAT (TRITICUM-AESTIVUML) LEAVES EXPOSED TO ELEVATED O-3 AND OR CO2/, New phytologist, 129(4), 1995, pp. 557-568
This study examined the effects of elevated ozone (O-3) and/or carbon
dioxide (CO2) on the diel allocation of photosynthetically fixed carbo
n in fully expanded leaves of young (growth stages 4-5) spring wheat (
Triticum aestivum L. cv. Hanno). Plants were grown in controlled envir
onment chambers and exposed to two O-3 regimes ['non-polluted' air (CF
), < 5 nmol mol(-1); 'polluted' air, CF + 75 nmol mol(-1) 7 h d(-1)] a
nd two CO2 treatments ('ambient', 354 mu mol mol(-1); 'elevated', 700
mu mol mol(-1)) over a 30 d period. Neutral sugars (predominantly sucr
ose) were found to be the most abundant form of carbohydrate accumulat
ed by leaves during the day, but significant quantities of starch and
high degree of polymerization (d.p.) fructans were also present. Eleva
ted concentrations of O-3 and/or CO2 were found to have marked effects
on diel patterns of export, storage and respiration, whilst the propo
rtions of fixed carbon allocated to each of these processes were broad
ly similar. O-3 depressed the rate of net CO2 assimilation (-20%) and
reduced stomatal conductance (-19%). This was reflected in a reduced a
mount of carbohydrate accumulated in, and exported by, source tissue d
uring the day. Effects of O-3 on the rate of CO2 fixation were aggrava
ted by an increased demand for carbon by dark respiratory processes. I
n contrast, doubling the atmospheric concentration of CO2 enhanced the
rate of net CO2 assimilation (+ 47%) and reduced the proportion of fi
xed carbon retained in the leaf blade, increasing the rate of export.
The favourable carbon balance of CO2 enriched leaves was further enhan
ced by a decrease in the cost of maintenance respiration, whilst simul
taneous measurements of CO2 efflux and O-2 uptake at night suggested a
shift in the substrates metabolized at high CO2. Effects of elevated
CO2 and O-3 on the carbon balance of individual leaf blades over a sin
gle 24 h light/dark cycle were entirely consistent with the cumulative
effects of the gases on plant growth over a 30 d period. O-3 reduced
the rate of plant growth (-10%), but there were differential effects o
f O-3 on the growth of root and shoot which exacerbated the decrease i
n assimilate availability induced by O-3. In contrast the favourable e
ffects of CO2 enrichment on the carbon balance of individual source le
aves was reflected in the enhanced accumulation of dry matter in exist
ing sinks, and the initiation of new sinks (i.e. increased tillering).
In the combined treatment (elevated CO2 + O-3), O-3 counteracted the
favourable effects of CO2 enrichment on the carbon balance of individu
al leaves, and the combined effects of the individual gases on the die
l partitioning of photosynthetically fixed carbon in fully expanded le
af blades was reflected in a decreased rate of plant growth at elevate
d CO2, a situation further exacerbated by O-3-induced shifts in the re
lative partitioning of carbon between root and shoot. There was no evi
dence that CO2 enrichment afforded additional protection against O-3 d
amage: the extent of the O-3-induced reduction in photosynthesis, carb
ohydrate availability and growth observed at elevated CO2 was similar
to that induced by O-3 in ambient air, despite additive effects of the
gases on stomatal conductance that would reduce the effective dose of
O-3 by approximate to 30%. The wider ecological significance of inter
actions between elevated CO2 and O-3 is discussed in the light of othe
r recent findings.