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Books
Responses of Northern U.S. Forests
to Environmental Change
ISBN 0-387-98900-5
Chapter 5: Interacting Effects of Multiple Stresses
on Growth and Physiological Processes in Northern Forest Trees
Judson G. Isebrands, Richard E. Dickson, Joanne
Rebbeck, and David F. Karnosky
Two pollutants that have substantial impacts on plant growth and
are increasing in the atmosphere as a consequence of human activity
are CO2 and O3. Plant responses to CO2 and O3 are complex, and become
even more difficult to interpret when other known stressors are
considered, including nitrogen limitation, temperature and moisture
extremes, and pests. In general, increasing atmospheric CO2 increases
photosynthetic rates, height growth, and biomass production, while
increasing atmospheric O3 decreases photosynthetic rates and biomass
production, and increases leaf senescence. The amount and sometimes
the direction of change depends on internal plant factors such as
age and genotype. Higher CO2 concentrations may compensate for some
other environmental stresses. For example, most studies show that
CO2 enrichment increases growth even though light and/or nutrients
are limiting. It is becoming clear that both increasing CO2 and
O3 impact fundamental plant processes which in turn affect susceptibility
to plant-feeding insects.
Studies on trembling aspen show that O3 usually decreases growth
although the effect varies significantly with genotype. Root growth
appears particularly sensitive to O3. In contrast, substantial increases
in relative below-ground C allocation were found in response to
elevated CO2. Experiments with both elevated CO2 and O3 suggest
that elevated CO2 does not compensate for reduced growth caused
by elevated O3. When N limits growth, there appears to be no response
to elevated CO2. Because CO2 and O3 change the chemical composition
of the foliage, resistance to insect attack and nutritional value
of foliage are altered. Elevated O3 appears to increase insect growth
and elevated CO2 decreases insect growth. Under field conditions,
these changes in insect physiology may offset increases or decreases
in biomass production that are associated with a changing atmosphere.
Consistent growth responses of yellow poplar to O3 have not been
reported even though the species shows visual foliar symptoms of
exposure. One study reported an increase in biomass production during
the first year of exposure in open-top chambers, and a decrease
after two seasons. Elevated CO2 appears to increase yellow poplar
growth regardless of level of exposure to O3. In general, research
on yellow poplar suggests that under field conditions, this species
will increase biomass production even when nutrients and moisture
are limited and in the presence of O3.
In summary, there appear to be major differences in the response
of species and genotypes to multiple interacting stresses. The mechanisms
causing the responses are largely unknown. Experimental methods
have a major impact on how results from these studies should be
interpreted. Significant chamber effects are common, limiting extrapolation
of many experimental results to field conditions. Open-air exposure
experiments and field physiological studies will help alleviate
this concern over time, as will improvements of physiological models.
Below: Changes in total
stem dry weight of aspen clones exposed to elevated O3
and O3 + CO2 for 3 years in open-top chambers.
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