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Whereas the metabolism of photorespiration has been the sub- MATERIALS AND METHODS
ject of numerous studies, relatively few data are available con-
cerning the associated gas fluxes in organs or whole plants, al- The experiments were conducted on whole wheat plants (Triti-
though such gas-exchange measurements were at the origin of the cum aestivum L var Champlein) grown from the 8th day after
discovery of, first, the Warburg effect, then the loss of CO2 (16) sowing in a "C23A mini-chamber" (volume, 6 to 18 liters). PAR
and the uptake of 02 in light (8, 22), which are the main expres- during growth was 175 w/m2 (610 ,IE/m2 s-1), day/night temper-
atures were 20 ± lC/15 ± IC, and the CO2 level was kept at 330
sions of photorespiration. Interest in the gas exchanges ebbed after
these discoveries, as the complexity of the underlying mechanisms tdl F'. The root compartment was separated from the aerial part.
Techniques were described in detail by Andre et al. (1, 2).
was uncovered through biochemistry, challenging the value of gas The plants were grown for 40 to 70 days. As the plants had not
flux measurements and showing the difficulty of their interpreta- undergone winter frosts, vegetative growth still continued after 70
tion. days without flowering. Apparent photosynthesis was around 100
Nonisotopic methods do not allbw a measurement of photores- ml/h at the end of the experiments, still increasing by 2 to 4%/
piration during photosynthesis; moreover, they are not consistent. day; it was limited mainly by the reciprocal shading and the
For example, the Warburg effect depends on CO2 concentration disturbance of the ventilation by the leaves.
but not on light intensity (6, 34), whereas the evolution of CO2 in The roots were placed in a beaker containing 2.3 liters nutrient
C02-free air and the CO2 postillumination burst (14, 37) increase solution at pH 6.5. This solution was changed every day and
with light intensity but are independent of CO2 (7). All results analyzed for nutrient uptake. The volume of solution was enough
agree on the stimulating action of 02 on photorespiration. to provide a roughly constant concentration of elements, except
Isotopic methods also have their flaws. As in the preceding case, NH4', which was exhausted after a few hours of photosynthesis.
several phenomena may be involved. Methods with 14C are used The experiments were done in the same growth chamber.
to measure true photosynthesis (CO2 uptake) (25) or CO2 evolution
(41), but contradictions have arisen (13). Unfortunately, the results Abbreviations: P, apparent photosynthesis or net CO2 assimilation; P',
'
are underestimated because of internal recycling phenomena (10, or net 02 evolution; E, 02 evolution; U, 02 uptake; PS, net
apparent
32) which can only be estimated. However, tests with 14C have photosynthesis in standard conditions; R, dark respiration; For clarity the
been used to estimate the photorespiratory loss of CO2 at 10 to gas fluxes are schematized in Figure 1.
1032
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Copyright © 1980 American Society of Plant Biologists. All rights reserved.
Plant Physiol. Vol. 66, 1980 02 EXCHANGES IN WHEAT 1033
0 100 PS~
ale(
0
0 100 200 300 400 500
9I1-1.' CO2
b 2% 62
E
C,)
0.
100 4R
C.) 10
LuA
R -
0 100 200 300 400
Light intensity (W.M-2)
FIG. 3. Effect of light intensity on the gas exchanges of a wheat plant
at 330 ill -'; 1 w m-2 = 3.48 ,uE m-2 s-'.
40 50 60 70
light intensities greater than 12.5 w m-2 (43.5 ,tE m-2 s-') and PLANT AuE (DAYS)
evolved afterwards following a classical curve. The ratio of P to U
was highest at 175 w m-2. This means that the reducing power FIG. 4. Evolution with plant age of the gas exchanges of wheat. As the
was used most efficiently at the standard light intensity, which was plot is semilogarithmic, the linear part of the curves (left) corresponds to
about half-saturating. the exponential growth period; the flat part of the curves (right) corre-
At higher intensities, P approached saturation but U continued sponds to a linear growth period.
growing linearly. Canvin et al. (I 1) have shown that U could even
exceed P at high light intensities in the C3 plant Hirschfeldia
incana Lowe. This indicates that the reaction with 02 has a low
affinity but a high maximum rate (higher than that of photosyn- 7.5
thesis), which allows an efficient elimination of excess reducing
power. Nevertheless, we can see that E does not increase linearly
with light intensity; this could be due to a beginning of saturation C)0.
of the photosystems or of the 02 uptake reaction.
0 5
AGE OF PLANT