Plant Diseases

Impacts of climate change on specific host-pathogen systems are variable (Coakley et al. 1999; Chakraborty et al. 2000). In addition, O3 effects on plants lead to altered disease susceptibility, but the effect is inconsistent. In wheat, leaf rust disease was strongly inhibited by O3, but largely unaffected by elevated CO2, both in the presence and absence of O3 stress (Von Tiedemann and Firsching 2000). O3 damage to leaves was strongly affected by CO2 concentration and infection. On infected plants, O3 lesions appeared 2-4 weeks earlier and were up to fourfold more severe compared to non-infected plants. Elevated CO2 did not delay the onset of lesions, but it significantly reduced the severity of leaf damage. The relative increases in growth and yield induced by CO2 were much larger on O3-stressed than on non-stressed plants. Both, O3 and fungal infection reduced biomass formation and grain yield. However, adverse effects of leaf rust infection were more severe. Elevated

CO2 compensated for the negative effects of O3, but not for the detrimental effects of fungal infection. Therefore, the impact of O3 in the field should not be estimated without considering the predisposing effects deriving from fungal infections and the compensating effects deriving from elevated CO2. The interaction between O3 and pathogens may be determined primarily by the timing of O3 exposure relative to the presence of inoculation. The outcome of plant-pathogen interactions may strongly vary with timing, stage of plant development, predisposing factors, and environmental conditions (Fuhrer 2003).

10.11 Conclusions, Policy Implications and Research Needs

This review has revealed that there is currently much uncertainty in predicting the effects of O3 in the changing climate of the 21st century. When the changes in mean global O3 concentration with changing profile (decreased peak with increasing background concentrations) are also brought into the predictions for the 21st century, the scope for generalising responses is rather limited. Furthermore, feedback mechanisms will influence the magnitude of climate change and therefore the effects on crops. For example, a reduction in the stomatal O3 flux in a future climate might lead to an increase in ambient O3 concentrations due to a reduction in stomatal deposition to vegetation and therefore result in enhanced radiative forcing. O3-induced productivity losses would continue to affect the global carbon cycle by reduced sequestration (Sitch et al. 2007).

The flux-based approach for the risk assessment of O3 effects on vegetation offers a possibility to include the modifying influence of the environment on plant responses to O3. As such, it becomes possible to predict with higher certainty how plants will respond to the cocktail of environmental conditions which is of particular relevance for evaluation of the impacts of global climate change and for the modelling of pollutant effects across continents. However, there are also disadvantages that need to be considered. These include the higher requirements of measured and modelled meteorological data to drive the models, the more complicated calculations to make risk assessments, and the challenge of communicating results which are based on more complex models than those purely using pollutant concentrations. These disadvantages are, however, inevitable to permit the generalisation of results, for instance, between years with different weather conditions, sites with different climate, estimations of future impacts of climate change and the quantification of economic consequences of O3 pollution (Pleijel 2005).

10.11.1 Policy Implications

When considering the policy implications of the effects of O3 in a changing climate, the following sources of uncertainty need to be taken into account:

• Plant responses to climate change (including elevated O3 concentrations) are species-specific and sometimes even cultivar-specific, and are influenced by other factors, such as local climate, soil characteristics and management.

• Vegetation responses to changes in single drivers of climate change cannot simply be scaled up to predict responses to changes in multiple drivers.

• Results from field release (FACE) systems provide the most reliable information on plant responses to combinations of drivers. Although FACE studies generally substantiate predictions based on chamber studies, some inconsistencies between the results of chamber and FACE studies have been reported. Results from chamber-based experiments might over- or underestimate crop responses to environmental changes.

10.11.2 Conclusions

Taking into account the uncertainties described above and the current information available the following conclusions on the plant responses to O3 in a changing climate can be tentatively drawn:

• There is a tendency for O3 effects to be less pronounced at elevated CO2. This effect has been detected in several crop species. Primarily, CO2-induced reduction in the stomatal uptake of O3 is believed to be responsible for this beneficial effect.

• Global warming may reduce the ameliorating effect of elevated CO2 on the plant response to O3. Under conditions of no water stress, at a temperature below the optimum for the species concerned, warming is likely to increase O3 uptake, while a rise in elevated CO2 is likely to decrease uptake.

• The combined drivers of climate change will influence the flux of O3 into leaves and will thus influence the magnitude of the effects for a given atmospheric concentration. The magnitude and direction of impact on flux will depend on the combined response to temperature, CO2 concentration, vapour pressure deficit, soil moisture content, O3 concentration and plant development stage at any moment in time.

• There is no conclusive evidence that elevated CO2 will increase the rate of O3 detoxification within plants. Responses appear to be species- specific. More studies show no effect than others showing an increase in detoxification of O3. The generalized conclusion is that the beneficial effects of CO2 in reducing O3 effects are mediated by stomatal closure rather than detoxification.

• Responses of insects and pathogens to the combined effects of elevated CO2 and O3 are species-specific. By altering leaf chemistry, both O3 and CO2 can enhance insect and pathogen attack in some species.

• Elevated O3 at relatively low concentrations can significantly reduce the growth enhancement by elevated CO2 and therefore reduce C sequestration. This may mean that worldwide vegetation growth stimulations will not be as great as previously predicted from elevated CO2 studies. Thus, it is important to bring an understanding of O3 as a moderator of CO2 responses in global models of terrestrial net primary productivity and C sequestration. The situation is even more complicated when plant-herbivore interactions or feedbacks operating through the soil are considered.

10.11.3 Research Recommendations

The following research recommendations are being made to improve scientific knowledge on the impacts of ozone on crops in a changing climate:

• The influence of climate change should be taken into account when predicting the future effects of O3 on vegetation and vice versa.

• Further improvements linking the actual O3 uptake and detoxification processes, to carbon assimilation and allocation will be needed to provide an even more reliable mechanistic basis for future risk assessments (Ashmore 2005; Mussel-man et al. 2006). Furthermore, the flux approach needs to be scaled to the canopy level to be generally applicable (Fiscus et al. 2005)

• In the longer-term, it may be necessary to develop alternative modelling procedures since the current method is based on the stomatal responses to climatic and plant factors considered in isolation rather than in combination.

• There is a clear need for multi-factorial experiments to provide information for O3-effect modelling. Because of the high cost involved with FACE systems, this may only be possible using enclosed or semi-enclosed chamber systems, especially where warming and CO2 are considered as factors.

• There is a need for a wider and longer-term use of FACE for crops to expand the range of species for which data exists.

• Much more work is needed before unified mechanistic models of O3 response can be developed that apply to all main agricultural and horticultural crops.

Acknowledgments This work was supported in part by the UK Department for Environment,

Food and Rural Affairs (Defra) under contract EPG 1/3/205 and AQ03509 with the Centre for

Ecology and Hydrology. The authors also wish to thank Dr Lisa Emberson and Dr Hakan Pleijel for supplying some of the figures used in this chapter.

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