The emission of isoprene from the leaves of forest trees is a fundamental component of biosphere-atmosphere interactions, controlling many aspects of photochemistry in the lower atmosphere. As almost all commercial agriforest species emit high levels of isoprene, proliferation of agriforest plantations has significant potential to increase regional ozone pollution and enhance the lifetime of methane, an important determinant of global climate. Here we show that growth of an intact Populus deltoides plantation under increased CO2 (800 micromol x mol(-1) and 1,200 micromol x mol(-1)) reduced ecosystem isoprene production by 21% and 41%, while above-ground biomass accumulation was enhanced by 60% and 82%, respectively. Exposure to increased CO2 significantly reduced the cellular content of dimethylallyl diphosphate, the substrate for isoprene synthesis, in both leaves and leaf protoplasts. We identify intracellular metabolic competition for phosphoenolpyruvate as a possible control point in explaining the suppression of isoprene emission under increased CO2. Our results highlight the potential for uncoupling isoprene emission from biomass accumulation in an agriforest species, and show that negative air-quality effects of proliferating agriforests may be offset by increases in CO2.

The partitioning of soil respiration rates into the component processes of rhizospheric respiration (because of live roots and those microorganisms that subsist on root exudations) and heterotrophic respiration (because of decomposer microorganisms that subsist on the oxidation of soil organic matter) is difficult to accomplish through experimental observation. In order to minimize disturbance to the soil and maximize preservation of the natural relationships among roots, rhizospheric microorganisms, and decomposers, we conducted a girdling experiment in a subalpine forest dominated by lodgepole pine trees. In two separate years, we girdled trees in small forest plots (5-7 m in diameter) and trenched around the plots to sever invading roots in order to experimentally stop the transport of photosynthate from needles to roots, and eliminate rhizospheric respiration. Soil respiration rates in plots with trees girdled over 1 year prior to measurement were higher than those in plots with trees girdled 2-3 months prior to measurement. These results suggest that any stimulation of respiration because of the experimental artifact of fine root death and addition of labile carbon to the pool of decomposer substrates is slow, and occurs beyond the first growing season after girdling. Compared with control plots with nongirdled trees, soil respiration rates in plots with girdled trees were reduced by 31-44% at the mid-summer respiratory maximum. An extreme drought during one of the 2 years used for observations caused greater reductions in the heterotrophic component of soil respiration compared with the rhizospheric component. In control plots, we observed a pulse in K 2 SO 4 -extractable carbon during the spring snowmelt period, which was absent in plots with girdled trees. In control plots, soil microbial biomass increased from spring to summer, coincident with a seasonal increase in the rhizospheric component of soil respiration. In plots with girdled trees, the seasonal increase in microbial biomass was lower than in control plots. These results suggest that the observed seasonal increase in rhizospheric respiration rate in control plots was because of an increase in rhizospheric microbial biomass following 'soil priming' by a spring-time pulse in dissolved organic carbon. Winter-time, beneath-snow microbial biomass was relatively high in control plots. Soil sucrose concentrations were approximately eight times higher during winter than during spring or summer, possibly being derived from the mechanical damage of shallow roots that use sucrose as protection against low-temperature extremes. The winter-time sucrose pulse was not observed in plots with girdled trees. The results of this study demonstrate that (1) the rhizospheric component of soil respiration rate at this site is significant in magnitude, (2) the heterotrophic component of soil respiration rate is more susceptible to seasonal drought than the rhizospheric component, and (3) the trees in this ecosystem exert a major control over soil carbon dynamics by 'pr...

The transition between wintertime net carbon loss and springtime net carbon assimilation has an important role in controlling the annual rate of carbon uptake in coniferous forest ecosystems. We studied the contributions of springtime carbon assimilation to the total annual rate of carbon uptake and the processes involved in the winter-to-spring transition across a range of scales from ecosystem CO2 fluxes to chloroplast photochemistry in a coniferous, subalpine forest. We observed numerous initiations and reversals in the recovery of photosynthetic CO2 uptake during the initial phase of springtime recovery in response to the passage of alternating warm- and cold-weather systems. Full recovery of ecosystem carbon uptake, whereby the 24-h cumulative sum of NEE (NEEdaily) was consistently negative, did not occur until 3-4 weeks after the first signs of photosynthetic recovery. A key event that preceded full recovery was the occurrence of isothermality in the vertical profile of snow temperature across the snow pack; thus, providing consistent daytime percolation of melted snow water through the snow pack. Interannual variation in the cumulative annual NEE (NEEannual) was mostly explained by variation in NEE during the snow-melt period (NEEsnow-melt), not variation in NEE during the snow-free part of the growing season (NEEsnow-free). NEEsnow-melt was highest in those years when the snow melt occurred later in the spring, leading us to conclude that in this ecosystem, years with earlier springs are characterized by lower rates of NEEannual, a conclusion that contrasts with those from past studies in deciduous forest ecosystems. Using studies on isolated branches we showed that the recovery of photosynthesis occurred through a series of coordinated physiological and biochemical events. Increasing air temperatures initiated recovery through the upregulation of PSII electron transport caused in part by disengagement of thermal energy dissipation by the carotenoid, zeaxanthin. The availability of liquid water permitted a slightly slower recovery phase involving increased stomatal conductance. The most rate-limiting step in the recovery process was an increase in the capacity for the needles to use intercellular CO2, presumably due to slow recovery of Rubisco activity. Interspecific differences were observed in the timing of photosynthetic recovery for the dominant tree species. The results of our study provide (1) a context for springtime CO2 uptake within the broader perspective of the annual carbon budget in this subalpine forest, and (2) a mechanistic explanation across a range of scales for the coupling between springtime climate and the carbon cycle of high-elevation coniferous forest ecosystems.

Seasonal differences in the capacity of photosynthetic electron transport, leaf pigment composition, xanthophyll cycle characteristics and chlorophyll fluorescence emission were investigated in two biennial mesophytes (Malva neglecta and Verbascum thapsus) that grow in full sunlight, and in leaves/needles of sun and shade populations of several broad‐leafed evergreens and conifers (Vinca minor, Euonymus kiautschovicus, Mahonia repens, Pseudotsuga menziesii [Douglas fir], and Pinus ponderosa). Both mesophytic species maintained or upregulated photosynthetic capacity in the winter and exhibited no upregulation of photoprotection. In contrast, photosynthetic capacity was downregulated in sun leaves/needles of V. minor, Douglas fir, and Ponderosa pine, and even in shade needles of Douglas fir. Interestingly, photosynthetic capacity was upregulated during the winter in shade leaves/needles of V. minor, Ponderosa pine and Euonymus kiautschovicus. Nocturnal retention of zeaxanthin and antheraxanthin, and their sustained engagement in a state primed for energy dissipation, were observed largely in the leaves/needles of sun‐exposed evergreen species during winter. Factors that may contribute to these differing responses to winter stress, including chloroplast redox state, the relative levels of source and sink activity at the whole plant level, and apoplastic versus symplastic phloem loading, are discussed.