Atmospheric CO2 enrichment increases growth and photosynthesis of Pinus taeda: a 4 year experiment in the field

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Abstract
Forest trees are major components of the terrestrial biome and their response to rising atmospheric CO2 plays a prominent role in the global carbon cycle. In this study, loblolly pine seedlings were planted in the field in recently disturbed soil of high fertility, and CO2 partial pressures were maintained at ambient CO2 (Amb) and elevated CO2 (Amb + 30 Pa) for 4 years. The objective of the study was to measure seasonal and long‐term responses in growth and photosynthesis of loblolly pine exposed to elevated CO2 under ambient field conditions of precipitation, light, temperature and nutrient availability. Loblolly pine trees grown in elevated CO2 produced 90% more biomass after four growing seasons than did trees grown in ambient CO2. This large increase in final biomass was primarily due to a 217% increase in leaf area in the first growing season which resulted in much higher relative growth rates for trees grown in elevated CO2. Although there was not a sustained effect of elevated CO2 on relative growth rate after the first growing season, absolute production of biomass continued to increase each year in trees grown in elevated CO2 as a consequence of the compound interest effect of increased leaf area on the production of more new leaf area and more biomass. Allometric analyses of biomass allocation patterns demonstrated size‐dependent shifts in allocation, but no direct effects of elevated CO2 on partitioning of biomass. Leaf photosynthetic rates were always higher in trees grown in elevated CO2, but these differences were greater in the summer (60–130% increase) than in the winter (14–44% increase), reflecting strong seasonal effects of temperature on photosynthesis. Our results suggest that seasonal variation in the relative photosynthetic response to elevated CO2 will occur in natural ecosystems, but total non‐structural carbohydrate (TNC) levels in leaves indicate that this variation may not always be related to sink activity. Despite indications of canopy‐level adjustments in carbon assimilation, enhanced levels of leaf photosynthesis coupled with increased total leaf area indicate that net carbon assimilation for the whole tree was greater for trees grown under elevated CO2 compared with ambient CO2. If the large growth enhancement observed in loblolly pine were maintained after canopy closure, then these trees could be a large sink for fossil carbon emitted to the atmosphere and produce a negative feedback on atmospheric CO2.