ALBERT

Notes: We studied net ecosystem CO2 exchange (NEE) dynamics in a high-elevation, subalpine forest in Colorado, USA, over a two-year period. Annual carbon sequestration for the forest was 6.71 mol C m−2 (80.5 g C m−2) for the year between November 1, 1998 and October 31, 1999, and 4.80 mol C m−2 (57.6 g C m−2) for the year between November 1, 1999 and October 31, 2000. Despite its evergreen nature, the forest did not exhibit net CO2 uptake during the winter, even during periods of favourable weather. The largest fraction of annual carbon sequestration occurred in the early growing-season; during the first 30 days of both years. Reductions in the rate of carbon sequestration after the first 30 days were due to higher ecosystem respiration rates when mid-summer moisture was adequate (as in the first year of the study) or lower mid-day photosynthesis rates when mid-summer moisture was not adequate (as in the second year of the study). The lower annual rate of carbon sequestration during the second year of the study was due to lower rates of CO2 uptake during both the first 30 days of the growing season and the mid-summer months. The reduction in CO2 uptake during the first 30 days of the second year was due to an earlier-than-normal spring warm-up, which caused snow melt during a period when air temperatures were lower and atmospheric vapour pressure deficits were higher, compared to the first 30 days of the first year. The reduction in CO2 uptake during the mid-summer of the second year was due to an extended drought, which was accompanied by reduced latent heat exchange and increased sensible heat exchange. Day-to-day variation in the daily integrated NEE during the summers of both years was high, and was correlated with frequent convective storm clouds and concomitant variation in the photosynthetic photon flux density (PPFD). Carbon sequestration rates were highest when some cloud cover was present, which tended to diffuse the photosynthetic photon flux, compared to periods with completely clear weather.The results of this study are in contrast to those of other studies that have reported increased annual NEE during years with earlier-than-normal spring warming. In the current study, the lower annual NEE during 2000, the year with the earlier spring warm-up, was due to (1) coupling of the highest seasonal rates of carbon sequestration to the spring climate, rather than the summer climate as in other forest ecosystems that have been studied, and (2) delivery of snow melt water to the soil when the spring climate was cooler and the atmosphere drier than in years with a later spring warm-up. Furthermore, the strong influence of mid-summer precipitation on CO2 uptake rates make it clear that water supplied by the spring snow melt is a seasonally limited resource, and summer rains are critical for sustaining high rates of annual carbon sequestration.

Notes: Plant responses to elevated atmospheric CO2 have been characterized generally by stomatal closure and enhanced growth rates. These responses are being increasingly incorporated into global climate models that quantify interactions between the biosphere and atmosphere, altering climate predictions from simpler physically based models. However, current information on CO2 responses has been gathered primarily from studies of crop and temperate forest species. In order to apply responses of vegetation to global predictions, CO2 responses in other commonly occurring biomes must be studied. A Free Air CO2 Enrichment (FACE) study is currently underway to examine plant responses to high CO2 in a natural, undisturbed Mojave Desert ecosystem in Nevada, USA. Here we present findings from this study, and its companion glasshouse experiment, demonstrating that field-grown Ephedra nevadensis and glasshouse-grown Larrea tridentata responded to high CO2 with reductions in the ratio of transpirational surface area to sapwood area (LSR) of 33% and 60%, respectively. Thus, leaf-specific hydraulic conductivity increased and stomatal conductance remained constant or was increased under elevated CO2. Field-grown Larrea did not show a reduced LSR under high CO2, and stomatal conductance was reduced in the high CO2 treatment, although the effect was apparent only under conditions of unusually high soil moisture. Both findings suggest that the common paradigm of 20–50% reductions in stomatal conductance under high CO2 may not be applicable to arid ecosystems under most conditions.

Notes: Stomatal function mediates physiological trade-offs associated with maintaining a favourable H2O balance in leaf tissues while acquiring CO2 as a photosynthetic substrate. The C3 and C4 species appear to have different patterns of stomatal response to changing light conditions, and variation in this behaviour may have played a role in the functional diversification of the different photosynthetic pathways. In the current study, we used gain analysis theory to characterize the stomatal conductance response to light intensity in nine different C3, C4 and C3-C4 intermediate species Flaveria species. The response of stomatal conductance (gs) to a change in light intensity represents both a direct (related to a change in incident light intensity, I) and indirect (related to a change in intercellular CO2 concentration, Ci) response. The slope of the line relating the change in gs to Ci was steeper in C4 species, compared with C3 species, with C3-C4 species having an intermediate response. This response reflects the greater relative contribution of the indirect versus direct component of the gs versus I response in the C4 species. The C3-C4 species, Flaveria floridana, exhibited a C4-like response whereas the C3-C4 species, Flaveria sonorensis and Flaveria chloraefolia, exhibited C3-like responses, similar to their hypothesized position along the evolutionary trajectory of the development of C4 photosynthesis. There was a positive correlation between the relative contribution of the indirect component of the gs versus I response and water use efficiency when evaluated across all species. Assuming that the C3-C4 intermediate species reflect an evolutionary progression from fully expressed C3 ancestors, the results of the current study demonstrate an increase in the contribution of the indirect component of the gs versus I response as taxa evolve toward the C4 extreme. The greater relative contribution of the indirect component of the stomatal response occurs through both increases in the indirect stomatal components and through decreases in the direct. Increases in the magnitude of the indirect component may be related to the maintenance of higher water use efficiencies in the intermediate evolutionary stages, before the appearance of fully integrated C4 photosynthesis.

Notes: The ability of seedlings to tolerate temperature extremes is important in determining the distribution of perennial plants in the arid south-western USA, and the manner in which elevated CO2 impacts the ability of plants to tolerate high temperatures is relatively unknown. Whereas the effects of chronic high temperature (30–38°C) and elevated CO2 are comparatively well understood, little research has assessed plant performance in elevated CO2 during extreme (〉 45 °C) temperature events. We exposed three species of Yucca to 360 and 700 μmol CO2 mol–1 for 8 months, then 9 d of high temperature (up to 53 °C) to evaluate the impacts of elevated CO2 on the potential for photosynthetic function during external high temperature. Seedlings of a coastal C3 species (Yucca whipplei), a desert C3 species (Yucca brevifolia), and a desert CAM species (Yucca schidigera), were used to test for differences among functional groups. In general, Yuccas exposed to elevated CO2 showed decreases in carboxylation efficiency as compared with plants grown at ambient before the initiation of high temperature. The coastal species (Y. whipplei) showed significant reductions (33%) in CO2 saturated maximum assimilation rate (Amax), but the desert species (Y. brevifolia and Y. schidigera) showed no such reductions in Amax. Stomatal conductance was lower in elevated CO2 as compared with ambient throughout the temperature event; however, there were species-specific differences over time. Elevated CO2 enhanced photosynthesis in Y. whipplei at high temperatures for a period of 4 d, but not for Y. brevifolia or Y. schidigera. Elevated CO2 offset photoinhibition (measured as Fv/Fm) in Y. whipplei as compared with ambient CO2, depending on exposure time to high temperature. Stable Fv/Fm in Y. whipplei occurred in parallel with increases in the quantum yield of photosystem II (ΦPSII) at high temperatures in elevated CO2. The value of ΦPSII remained constant or decreased with increasing temperature in all other treatment and species combinations. This suggests that the reductions in Fv/Fm resulted from thermal energy dissipation in the pigment bed for Y. brevifolia and Y. schidigera. The greater efficiency of photosystem II in Y. whipplei helped to maintain photosynthetic function at high temperatures in elevated CO2. These patterns are in contrast to the hypothesis that high temperatures in elevated CO2 would increase the potential for photoinhibition. Our results suggest that elevated CO2 may offset high-temperature stress in coastal Yucca, but not in those species native to drier systems. Therefore, in the case of Y. whipplei, elevated CO2 may allow plants to survive extreme temperature events, potentially relaxing the effects of high temperature on the establishment in novel habitats.

Notes: The photosynthetic response of Larrea tridentata Cav., an evergreen Mojave Desert shrub, to elevated atmospheric CO2 and drought was examined to assist in the understanding of how plants from water-limited ecosystems will respond to rising CO2. We hypothesized that photosynthetic down-regulation would disappear during periods of water limitation, and would, therefore, likely be a seasonally transient event. To test this we measured photosynthetic, water relations and fluorescence responses during periods of increased and decreased water availability in two different treatment implementations: (1) from seedlings exposed to 360, 550, and 700 μmol mol–1 CO2 in a glasshouse; and (2) from intact adults exposed to 360 and 550 μmol mol–1 CO2 at the Nevada Desert FACE (Free Air CO2 Enrichment) Facility. FACE and glasshouse well-watered Larrea significantly down-regulated photosynthesis at elevated CO2, reducing maximum photosynthetic rate (Amax), carboxylation efficiency (CE), and Rubisco catalytic sites, whereas droughted Larrea showed a differing response depending on treatment technique. Amax and CE were lower in droughted Larrea compared with well-watered plants, and CO2 had no effect on these reduced photosynthetic parameters. However, Rubisco catalytic sites decreased in droughted Larrea at elevated CO2. Operating Ci increased at elevated CO2 in droughted plants, resulting in greater photosynthetic rates at elevated CO2 as compared with ambient CO2. In well-watered plants, the changes in operating Ci, CE and Amax resulted in similar photosynthetic rates across CO2 treatments. Our results suggest that drought can diminish photosynthetic down-regulation to elevated CO2 in Larrea, resulting in seasonally transient patterns of enhanced carbon gain. These results suggest that water status may ultimately control the photosynthetic response of desert systems to rising CO2.