A.H.C.M. Schapendonk

Effects of elevated CO2 concentration on photosynthetic acclimation and productivity of two potato cultivars grown in open-top chambers

In two subsequent years, an early maturing potato cultivar with low leaf area index (LAI) and a late culti- var with high LAI were grown at concentrations of 350 and 700 µL CO2 L -1 in open-top chambers. The average increase of tuber dry matter yield by elevated CO2 was 27% in 1995 and 49% in 1996. During the first weeks after planting, elevated CO2 stimulated the light-saturated rate of photosynthesis (Amax) of both cultivars by 80%. However, Amax under elevated CO2 declined to the level of the low-CO2 treatment in the course of the growing season. In 1995 this convergence due to acclimation of photosynthesis was completed within 6 weeks, but in 1996, accli- mation proceeded until the end of the growing season. Photosynthetic acclimation was accompanied by a reduced Rubisco content, and was correlated more closely with accumulation of sucrose than of starch. From fluorescence measurements it was concluded that, in contrast to the carboxylation efficiency, the efficiency of photosynthetic reactions centers was not affected by acclimation to elevated CO2. The faster photosynthetic acclimation in 1995 coincided with overall lower values of Amax, crop growth rate and growth response to elevated CO2. It is shown that the indeterminate growth pattern of potato with its large sink capacity does not preclude acclimation. The effect of acclimation on yield was quantified by computer simulations. The simulated results indicated that photosynthetic acclimation reduced the positive effect of elevated CO2 on tuber yield by 50%.

On the relative magnitudes of photosynthesis, respiration, growth and carbon storage in vegetation

BACKGROUND AND AIMS The carbon balance of vegetation is dominated by the two large fluxes of photosynthesis (P) and respiration (R). Mechanistic models have attempted to simulate the two fluxes separately, each with their own set of internal and external controls. This has led to model predictions where environmental change causes R to exceed P, with consequent dieback of vegetation. However, empirical evidence suggests that the R : P ratio is constrained to a narrow range of about 0.4-0.5. Physiological explanations for the narrow range are not conclusive. The aim of this work is to introduce a novel perspective by theoretical study of the quantitative relationship between the four carbon fluxes of P, R, growth and storage (or its inverse, remobilization). METHODS Starting from the law of conservation of mass - in this case carbon - equations are derived for the relative magnitudes of all carbon fluxes, which depend on only two parameters: the R : P ratio and the relative rate of storage of carbon in remobilizable reserves. The equations are used to explain observed flux ratios and to analyse incomplete data sets of carbon fluxes. KEY RESULTS The storage rate is shown to be a freely varying parameter, whereas R : P is narrowly constrained. This explains the constancy of the ratio reported in the literature. With the information thus gained, a data set of R and P in grassland was analysed, and flux estimates could be derived for the periods after cuts in which plant growth is dominated by remobilization before photosynthesis takes over. CONCLUSIONS It is concluded that the relative magnitudes of photosynthesis, respiration, growth and substrate storage are indeed tightly constrained, but because of mass conservation rather than for physiological reasons. This facilitates analysis of incomplete data sets. Mechanistic models, as the embodiment of physiological mechanisms, need to show consistency with the constraints.

Radiation and nitrogen use at the leaf and canopy level by wheat and oilseed rape during the critical period for grain number definition

During the critical period for grain number definition, the amount of biomass produced per unit absorbed radiation is more sensitive to nitrogen (N) supply in oilseed rape than in wheat, and reaches a higher value at high N. This response was investigated by combining experimental and modelling work. Oilseed rape and wheat were grown at three levels of N supply, combined with two levels of plant density at high N supply. Canopy photosynthesis and daytime radiation use efficiency (RUEA) were calculated with a model based on observed N-dependent leaf photosynthesis and observed canopy vertical distribution of light and leaf N. In oilseed rape, RUEA was higher than in wheat and, in contrast to wheat, the sensitivity to canopy leaf N content increased from the start to the end of the critical period. These results were partly explained by the higher leaf photosynthesis in oilseed rape vs wheat. In addition, oilseed rape leaves were increasingly shaded by the inflorescence. Thus, RUEA increased because more leaves were operating at non-saturating light levels. In both species, the vertical distribution of leaf N was close to that optimising canopy photosynthesis. The results are discussed in relation to possibilities for improvement of N productivity in these crops.

Nitrogen shortage in a tomato crop; Scaling up from effects of electron transport rate to plant productivity

Optimization of nitrogen supply in a tomato plant is stepwise assessed, from chloroplast characteristics to whole crop performance. Experiments are reported in which important key processes are quantified in relation to the nitrogen content of leaves. Interactions of N effects with leaf aging and canopy light distribution are analyzed. A simulation model that integrates this knowledge is constructed. The results of three nitrogen partitioning scenarios for Rubisco suggest that optimization of the distribution of Rubisco between leaf layers is less important for plant productivity than within-leaf optimization of the photosynthetic apparatus

Temperature sensitivity of photosynthesis in Lolium perenne swards: a comparison of two methods for deriving photosynthetic parameters from in vivo measurements

The seasonal variation in photosynthetic rate of grass swards is partly the result of changes in the environment and partly the result of changes in the photosynthetic capacity of the sward itself. We evaluated two types of photosynthesis equations regarding their capacity to analyse seasonal and short-term temperature effects on photosynthesis of ryegrass (Lolium perenne L.). Intact cores of a field-grown ryegrass sward were taken to the laboratory 10 d after cutting for measurement of photosynthesis under controlled conditions. This was done during two four-week periods, in summer and autumn. Net photosynthetic rate (PN) of the sward was lower in autumn than in summer. Both a simple negatively exponential photosynthesis irradiance-response curve and the Farquhar equations for photosynthesis were applied to the in vivo canopy measurements. Application of the irradiance-response curve showed that irradiance-saturated gross photosynthetic rate increased linearly with increasing temperature and was higher in summer than in autumn. The initial radiation use efficiency did not differ between the seasons but decreased with the temperature rise. This explains the observation that total canopy photosynthetic rate decreased after short-term temperature increases in both seasons. The parameters in Farquhar equations that represent the temperature sensitivity of the maximum electron transport rate and of the Michaelis-Menten constants for CO2 and O2 fixation could not be quantified satisfactorily. Parameterisation of the Farquhar equations was hampered by a lack of robust information on many biochemical parameters, and the use of simple empirical response-functions may be preferable in the case of in vivo canopy measurements on grass swards.

Exploring the optimum nitrogen partitioning to predict the acclimation of C3 leaf photosynthesis to varying growth conditions

An analytical solution to the optimum partitioning of leaf nitrogen among photosynthetic proteins is developed, providing a new simple avenue of modelling leaf photosynthetic acclimation to environmental conditions during growth.

Climate change and agricultural productivity

Abstract Research performed at the AB-DLO and the Free University was intended to quantify the effects of a doubled CO 2 concentration on some key agronomic species and grasslands. A set of physiological and morphological processes was studied and related to above-and below ground cycling of carbon. The research was based on experiments and simulation studies at the level of plant, crop and soil in laboratory facilities and semi-field conditions with controlled CO 2 supply. Agricultural crops were grown in “Open Top Chambers” or greenhouses and grasses in transparent tunnels made of Lexan. Soil processes and root respiration were studied in the Wageningen Rhizolab. Photosynthesis and assimilate partitioning were measured simultaneously in the photosynthesis laboratory. Simulation models appeared to be useful tools to quantify the consequences of elevated CO 2 and climate change on the productivity of grasses and crops with different growth strategies.