Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests.

Three young northern temperate forest communities in the north-central United States were exposed to factorial combinations of elevated carbon dioxide (CO2 ) and tropospheric ozone (O3 ) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO2 enhanced ecosystem C content by 11%, whereas elevated O3 decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO2 and O3 . Treatment effects on ecosystem C content resulted primarily from changes in the near-surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r(2) = 0.96). Elevated CO2 enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m(-2) ) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O3 lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (∆NPP/∆N) decreased through time with further canopy development, the O3 effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O3 and less soil C from 0.1 to 0.2 m in depth under elevated CO2 . Overall, these results suggest that elevated CO2 may create a sustained increase in NPP, whereas the long-term effect of elevated O3 on NPP will be smaller than expected. However, changes in soil C are not well-understood and limit our ability to predict changes in ecosystem C content.

Partitioning of 14C-labeled photosynthate to allelochemicals and primary metabolites in source and sink leaves of aspen: evidence for secondary metabolite turnover.

Theories on allelochemical concentrations in plants are often based upon the relative carbon costs and benefits of multiple metabolic fractions. Tests of these theories often rely on measuring metabolite concentrations, but frequently overlook priorities in carbon partitioning. We conducted a pulse-labeling experiment to follow the partitioning of 14CO2-labeled photosynthate into ten metabolic pools representing growth and maintenance (amino acids, organic acids, lipids plus pigments, protein, residue), defense (phenolic glycosides, methanol:water and acetone-soluble tannins/phenolics), and transport and storage (sugars and starch) in source and importing sink leaves of quaking aspen (Populus tremuloides). The peak period of 14C incorporation into sink leaves occurred at 24 h. Within 48 h of labeling, the specific radioactivity (dpm/mg dry leaf weight) of phenolic glycosides declined by over one-third in source and sink leaves. In addition, the specific radioactivity in the tannin/phenolic fraction decreased by 53% and 28% in source and sink leaves, respectively. On a percent recovery basis, sink leaves partitioned 1.7 times as much labeled photosynthate into phenolic glycosides as source leaves at peak 14C incorporation. In contrast, source leaves partitioned 1.8 times as much 14C-labeled photosynthate into tannins/phenolics as importing sink leaves. At the end of the 7-day chase period, sink leaves retained 18%, 52%, and 30% of imported 14C photosynthate, and labeled source leaves retained 15%, 66%, and 19% of in situ photosynthate in metabolic fractions representing transport and storage, growth and maintenance, and defense, respectively. Analyses of the phenolic fractions showed that total phenolics were twice as great and condensed tannins were 1.7 times greater in sink than in source leaves. The concentration of total phenolics and condensed tannins did not change in source and sink leaves during the 7-day chase period.

Distribution and metabolism of current photosynthate by single-flush northern red oak seedlings.

Carbon distribution and metabolism by northern red oak seedlings (Quercus rubra L.) were followed for 72 h after a 30-min photosynthetic exposure to (14)CO(2). Approximately 50% of the assimilated carbon was lost during this time, presumably through respiration. Most of the (14)C recovered in the plant remained in the source leaf. Most of the exported (14)C was found in stems and especially roots. Position of the source leaf did not affect distribution of translocated (14)C; however, upper source leaves retained slightly more (14)C than lower source leaves. Most (14)C in all tissues was found initially in sugars. Labeled carbon in this fraction decreased rapidly and increased in other chemical fractions, particularly residue (structural carbohydrates and lignin). More (14)C was incorporated into residue than into any other chemical fraction, indicating continued growth by each of the tissues during the lag stage of seedling development. Labeled carbon increased in proteins for 6 to 12 h after treatment, then remained essentially constant for the remainder of the transport period, indicating both rapid incorporation of the pulsed (14)C into, and slow turnover of, the labeled protein components. In source leaves, (14)C incorporated into starch in the light was lost during the dark period, a typical diurnal storage pattern. In stems, (14)C in starch increased for 12 h, then decreased during the dark period, and then increased during the rest of the transport period. This pattern indicates that stems have both diurnal and long-term storage pools of starch. In roots, (14)C in starch increased rapidly for 12 h, then remained constant for the rest of the transport period, indicating primarily long-term storage in this tissue. Peak (14)C in amino acids in taproot preceded that in stem tissue, indicating recycling of (14)C from the roots to the shoot in amino acids.

Altered performance of forest pests under atmospheres enriched by CO2 and O3.

Human activity causes increasing background concentrations of the greenhouse gases CO2 and O3. Increased levels of CO2 can be found in all terrestrial ecosystems. Damaging O3 concentrations currently occur over 29% of the world's temperate and subpolar forests but are predicted to affect fully 60% by 2100 (ref. 3). Although individual effects of CO2 and O3 on vegetation have been widely investigated, very little is known about their interaction, and long-term studies on mature trees and higher trophic levels are extremely rare. Here we present evidence from the most widely distributed North American tree species, Populus tremuloides, showing that CO2 and O3, singly and in combination, affected productivity, physical and chemical leaf defences and, because of changes in plant quality, insect and disease populations. Our data show that feedbacks to plant growth from changes induced by CO2 and O3 in plant quality and pest performance are likely. Assessments of global change effects on forest ecosystems must therefore consider the interacting effects of CO2 and O3 on plant performance, as well as the implications of increased pest activity.