A mechanistic evaluation of photosynthetic acclimation at elevated CO2

Plants grown at elevated pCO₂ often fail to sustain the initial stimulation of net CO₂ uptake rate (A). This reduced, acclimated, stimulation of A often occurs concomitantly with a reduction in the maximum carboxylation velocity (V_c,max) of Rubisco. To investigate this relationship we used the Farquhar model of C₃ photosynthesis to predict the minimum V_c,max capable of supporting the acclimated stimulation in A observed at elevated pCO₂. For a wide range of species grown at elevated pCO₂ under contrasting conditions we found a strong correlation between observed and predicted values of V_c,max. This exercise mechanistically and quantitatively demonstrated that the observed acclimated stimulation of A and the simultaneous decrease in V_c,max observed at elevated pCO₂ is mechanistically consistent. With the exception of plants grown at a high elevated pCO₂ (> 90 Pa), which show evidence of an excess investment in Rubisco, the failure to maintain the initial stimulation of A is almost entirely attributable to the decrease in V_c,max and investment in Rubisco is coupled to requirements.     

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

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.     

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO2 and/or O3

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

Effects of free-air CO2 enrichment (FACE) on CH4 emission from a rice paddy field

Methane (CH₄) is a particularly potent greenhouse gas with a radiative forcing 23 times that of CO₂ on a per mass basis. Flooded rice paddies are a major source of CH₄ emissions to the Earth's atmosphere. A free‐air CO₂ enrichment (FACE) experiment was conducted to evaluate changes in crop productivity and the crop ecosystem under enriched CO₂ conditions during three rice growth seasons from 1998 to 2000 in a rice paddy at Shizukuishi, Iwate, Japan. To understand the influence of elevated atmospheric CO₂ concentrations on CH₄ emission, we measured methane flux from FACE rice fields and rice fields with ambient levels of CO₂ during the 1999 and 2000 growing seasons. Methane production and oxidation potentials of soil samples collected when the rice was at the tillering and flowering stages in 2000 were measured in the laboratory by the anaerobic incubation and alternative propylene substrates methods, respectively. The average tiller number and root dry biomass were clearly larger in the plots with elevated CO₂ during all rice growth stages. No difference in methane oxidation potential between FACE and ambient treatments was found, but the methane production potential of soils during the flowering stage was significantly greater under FACE than under ambient conditions. When free‐air CO₂ was enriched to 550 ppmv, the CH₄ emissions from the rice paddy field increased significantly, by 38% in 1999 and 51% in 2000. The increased CH₄ emissions were attributed to accelerated CH₄ production potential as a result of more root exudates and root autolysis products and to increased plant‐mediated CH₄ emissions because of the larger rice tiller numbers under FACE conditions.     

Asymmetric responses of adaxial and abaxial stomata to elevated CO2: impacts on the control of gas exchange by leaves

The response of adaxial and abaxial stomatal conductance in Rumex obtusifolius to growth at elevated atmospheric concentrations of CO₂ (250 μmol mol^?1 above ambient) was investigated over two growing seasons. The conductance of both the adaxial and abaxial leaf surfaces was found to be reduced by elevated concentrations of CO₂. Elevated CO₂ caused a much greater reduction in conductance for the adaxial surface than for the abaxial surface. The absence of effects upon stomatal density indicated that the reductions were probably the result of changes in stomatal aperture. Partitioning of gas exchange between the leaf surfaces revealed that increased concentrations of CO₂ caused increased rates of photosynthesis only via the abaxial surface. Additionally, leaf thickness was found to increase during growth at elevated concentrations of CO₂. The tendency for these amphistomatous leaves to develop a distribution of conductance approaching that of hypostomatous leaves clearly reduced their maximum photosynthetic potential. This conclusion was supported by measurements of stomatal limitation, which showed greater values for the adaxial surfaces, and greater values at elevated CO₂. This reduction in photosynthesis may in part be caused by higher diffusive limitations imposed because of increased leaf thickness. In an uncoupled canopy, asymmetrical stomatal responses of the kind identified here may appreciably reduce transpiration. Species which show symmetrical responses are less likely to show reduced transpirational rates, and a redistribution of water loss between species may occur. The implications of asymmetrical stomatal responses for photosynthesis and canopy transpiration are discussed.     

Gross primary production is stimulated for three Populus species grown under free-air CO2 enrichment from planting through canopy closure

How forests will respond to rising [CO₂] in the long term is uncertain, most studies having involved juvenile trees in chambers prior to canopy closure. Poplar free‐air CO₂ enrichment (Viterbo, Italy) is one of the first experiments to grow a forest from planting through canopy closure to coppice, entirely under open‐air conditions using free‐air CO₂ enrichment technology. Three Populus species: P. alba, P. nigra and P. x euramericana, were grown in three blocks, each containing one control and one treatment plot in which CO₂ was elevated to the expected 2050 concentration of 550 ppm. The objective of this study was to estimate gross primary production (GPP) from recorded leaf photosynthetic properties, leaf area index (LAI) and meteorological conditions over the complete 3‐year rotation cycle. From the meteorological conditions recorded at 30 min intervals and biweekly measurements of LAI, the microclimate of leaves within the plots was estimated with a radiation transfer and energy balance model. This information was in turn used as input into a canopy microclimate model to determine light and temperature of different leaf classes at 30 min intervals which in turn was used with the steady‐state biochemical model of leaf photosynthesis to compute CO₂ uptake by the different leaf classes. The parameters of these models were derived from measurements made at regular intervals throughout the coppice cycle. The photosynthetic rates for different leaf classes were summed to obtain canopy photosynthesis, i.e. GPP. The model was run for each species in each plot, so that differences in GPP between species and treatments could be tested statistically. Significant stimulation of GPP driven by elevated [CO₂] occurred in all 3 years, and was greatest in the first year (223–251%), but markedly lower in the second (19–24%) and third years (5–19%). Increase in GPP in elevated relative to control plots was highest for P. nigra in 1999 and for P. x euramericana in 2000 and 2001, although in 1999 P. alba had a higher GPP than P. x euramericana. Our analysis attributed the decline in stimulation to canopy closure and not photosynthetic acclimation. Over the 3‐year rotation cycle from planting to harvest, the cumulative GPP was 4500, 4960 and 4010 g C m^?2 for P. alba, P. nigra and P. x euramericana, respectively, in current [CO₂] and 5260, 5800 and 5000 g C m^?2 in the elevated [CO₂] treatments. The relative changes were consistent with independent measurements of net primary production, determined independently from biomass increments and turnover.     

Stomatal responses to increased CO2: implications from the plant to the global scale

Increased atmospheric CO₂ often but not always leads to large decreases in leaf conductance. Decreased leaf conductance has important implications for a number of components of CO₂ responses, from the plant to the global scale. All of the factors that are sensitive to a change in soil moisture, either amount or timing, may be affected by increased CO₂. The list of potentially sensitive processes includes soil evaporation, run-off, decomposition, and physiological adjustments of plants, as well as factors such as canopy development and the composition of the plant and microbial communities. Experimental evidence concerning ecosystem-scale consequences of the effects of CO₂ on water use is only beginning to accumulate, but the initial indication is that, in water-limited areas, the effects of CO₂-induced changes in leaf conductance are comparable in importance to those of CO,₂-induced changes in photosynthesis. Above the leaf scale, a number of processes interact to modulate the response of canopy or regional evapotran-spiration to increased CO₂. While some components of these processes tend to amplify the sensitivity of evapo-transpiration to altered leaf conductance, the most likely overall pattern is one in which the responses of canopy and regional evapotranspiration are substantially smaller than the responses of canopy conductance. The effects of increased CO₂ on canopy evapotranspiration are likely to be smallest in aerodynamically smooth canopies with high leaf conductances. Under these circumstances, which are largely restricted to agriculture, decreases in evapotranspiration may be only one-fourth as large as decreases in canopy conductance. Decreased canopy conductances over large regions may lead to altered climate, including increased temperature and decreased precipitation. The simulation experiments to date predict small effects globally, but these could be important regionally, especially in combination with radiative (greenhouse) effects of increased CO₂.     

Changes in net photosynthesis and growth of Pinus eldarica seedlings in response to atmospheric CO2 enrichment

Pinus eldarica L. trees, rooted in the natural soil of an agricultural field at Phoenix, Arizona, were grown from the seedling stage in clear-plastic-wall open-top enclosures maintained at four different atmospheric CO₂ concentrations for 15 months. Light response functions were determined for one tree from each treatment by means of whole-tree net CO₂ exchange measurements at the end of this period, after which rates of carbon assimilation of an ambient-treatment tree were measured across a range of atmospheric CO₂ concentrations. The first of these data sets incorporates the consequences of both the CO₂-induced enhancement of net photosynthesis per unit needle area and the CO₂-induced enhancement of needle area itself (due primarily to the production of more needles), whereas the second data set reflects only the first of these effects. Hence the division of the normalized results of the first data set by the normalized results of the second set yields a representation of the increase in whole-tree net photosynthesis due to enhanced needle production caused by atmospheric CO₂ enrichment. In the solitary trees we studied, the relative contribution of this effect increased rapidly with the CO₂ concentration of the air to increase whole-tree net photosynthesis by nearly 50% at a CO₂ concentration approximately 300 μmol mol⁻¹ above ambient.     

Future atmospheric CO2 leads to delayed autumnal senescence

Growing seasons are getting longer, a phenomenon partially explained by increasing global temperatures. Recent reports suggest that a strong correlation exists between warming and advances in spring phenology but that a weaker correlation is evident between warming and autumnal events implying that other factors may be influencing the timing of autumnal phenology. Using freely rooted, field‐grown Populus in two Free Air CO₂ Enrichment Experiments (AspenFACE and PopFACE), we present evidence from two continents and over 2 years that increasing atmospheric CO₂ acts directly to delay autumnal leaf coloration and leaf fall. In an atmosphere enriched in CO₂ (by ～45% of the current atmospheric concentration to 550 ppm) the end of season decline in canopy normalized difference vegetation index (NDVI) – a commonly used global index for vegetation greenness – was significantly delayed, indicating a greener autumnal canopy, relative to that in ambient CO₂. This was supported by a significant delay in the decline of autumnal canopy leaf area index in elevated as compared with ambient CO₂, and a significantly smaller decline in end of season leaf chlorophyll content. Leaf level photosynthetic activity and carbon uptake in elevated CO₂ during the senescence period was also enhanced compared with ambient CO₂. The findings reveal a direct effect of rising atmospheric CO₂, independent of temperature in delaying autumnal senescence for Populus, an important deciduous forest tree with implications for forest productivity and adaptation to a future high CO₂ world.     

A free-air enrichment system for exposing tall forest vegetation to elevated atmospheric CO2

A free-air CO₂ enrichment (FACE) system was designed to permit the experimental exposure of tall vegetation such as stands of forest trees to elevated atmospheric CO₂ concentrations ([CO₂]_a) without enclosures that alter tree microenvironment. We describe a prototype FACE system currently in operation in forest plots in a maturing loblolly pine (Pinus taeda L.) stand in North Carolina, USA. The system uses feedback control technology to control [CO₂] in a 26 m diameter forest plot that is over 10 m tall, while monitoring the 3D plot volume to characterize the whole-stand CO₂ regime achieved during enrichment. In the second summer season of operation of the FACE system, atmospheric CO₂ enrichment was conducted in the forest during all daylight hours for 96.7% of the scheduled running time from 23 May to 14 October with a preset target [CO₂] of 550 μmol mol^–1, ≈ 200 μmol mol^–1 above ambient [CO₂]. The system provided spatial and temporal control of [CO₂] similar to that reported for open-top chambers over trees, but without enclosing the vegetation. The daily average daytime [CO₂] within the upper forest canopy at the centre of the FACE plot was 552 ± 9 μmol mol^–1 (mean ± SD). The FACE system maintained 1-minute average [CO₂] to within ± 110 μmol mol^–1 of the target [CO₂] for 92% of the operating time. Deviations of [CO₂] outside of this range were short-lived (most lasting < 60 s) and rare, with fewer than 4 excursion events of a minute or longer per day. Acceptable spatial control of [CO₂] by the system was achieved, with over 90% of the entire canopy volume within ± 10% of the target [CO₂] over the exposure season. CO₂ consumption by the FACE system was much higher than for open-top chambers on an absolute basis, but similar to that of open-top chambers and branch bag chambers on a per unit volume basis. CO₂ consumption by the FACE system was strongly related to windspeed, averaging 50 g CO₂ m^–3 h^–1 for the stand for an average windspeed of 1.5 m s^–1 during summer. The [CO₂] control results show that the free-air approach is a tractable way to study long-term and short-term alterations in trace gases, even within entire tall forest ecosystems. The FACE approach permits the study of a wide range of forest stand and ecosystem processes under manipulated [CO₂]_a that were previously impossible or intractable to study in true forest ecosystems.     

Acclimation response of spring wheat in a free-air CO2 enrichment (FACE) atmosphere with variable soil nitrogen regimes. 1. Leaf position and phenology determine acclimation response

We have examined the photosynthetic acclimation of wheat leaves grown at an elevated CO₂ concentration, and ample and limiting N supplies, within a field experiment using free-air CO₂ enrichment (FACE). To understand how leaf age and developmental stage affected any acclimation response, measurements were made on a vertical profile of leaves every week from tillering until maturity. The response of assimilation (A) to internal CO₂ concentration (C_i) was used to estimate the in vivo carboxylation capacity (Vc_max) and maximum rate of ribulose-1,5-bisphosphate limited photosynthesis (A _sat). The total activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), and leaf content of Rubisco and the Light Harvesting Chlorophyll a/b protein associated with Photosystem II (LHC II), were determined. Elevated CO₂ did not alter Vc_max in the flag leaf at either low or high N. In the older shaded leaves lower in the canopy, acclimatory decline in Vc_max and A _sat was observed, and was found to correlate with reduced Rubisco activity and content. The dependency of acclimation on N supply was different at each developmental stage. With adequate N supply, acclimation to elevated CO₂ was also accompanied by an increased LHC II/Rubisco ratio. At low N supply, contents of Rubisco and LHC II were reduced in all leaves, although an increased LHC II/Rubisco ratio under elevated CO₂ was still observed. These results underscore the importance of leaf position, leaf age and crop developmental stage in understanding the acclimation of photosynthesis to elevated CO₂ and nutrient stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Stomatal behaviour, photosynthesis and transpiration under rising CO2

Definitions of the variables used and the units are given in Table 1

The literature reports enormous variation between species in the extent of stomatal responses to rising CO₂. This paper attempts to provide a framework within which some of this diversity can be explained. We describe the role of stomata in the short-term response of leaf gas exchange to increases in ambient CO₂ concentration by developing the recently proposed stomatal model of Jarvis & Davies (1998 ). In this model stomatal conductance is correlated with the functioning of the photosynthetic system so that the effects of increases in CO₂ on stomata are experienced through changes in the rate of photosynthesis in a simple and mechanistically transparent way. This model also allows us to consider the effects of evaporative demand and soil moisture availability on stomatal responses to photosynthesis and therefore provides a means of considering these additional sources of variation. We emphasize that the relationship between the rate of photosynthesis and the internal CO₂ concentration and also drought will have important effects on the relative gains to be achieved under rising CO₂.  

  Table 1 . Abbreviations     

19.

Simple carbon assimilation response functions from atmospheric CO₂, and daily temperature and shortwave radiation     

D.S. WILKS  D.W. WOLFE  S.J. RIHA 《Global Change Biology》1995,1(5):337-346

A global ‘CO₂ fertilizer effect’ multiplier is often used in crop or ecosystem models because of its simplicity. However, this approach does not take into account the interaction between CO₂, temperature and light on assimilation. This omission can lead to significant under- or overestimation of the magnitude of beneficial effects from elevated CO₂, depending on environmental conditions. We use a mechanistic model of the biochemistry of photosynthesis to represent the response of net assimilation to different levels of CO₂, temperature and radiation, on the daily time scale. Instantaneous assimilation rates for an idealized canopy model are integrated through diurnal cycles of environmental variables derived from historical climate data at three locations in North America. The calculated CO₂ fertilizer effect is greatest at high light and warm temperatures. The results are summarized by assimilation response surfaces specified by the CO₂ concentration, the canopy leaf area index, and by daily values of temperature and radiation available from climatic records. These summary functions are suitable for incorporation into crop or ecosystem models for predicting carbon assimilation or biomass production on a daily time step. An example application of the function reveals that for a relatively cool, high latitude location, the beneficial effects from a CO₂ doubling would be negligible during the early spring, even assuming a + 4°C global warming scenario. In contrast, the beneficial effects from increasing CO₂ at a relatively warm, lower latitude location are greatest in the spring, but decline in late summer because of excessively warm temperatures with a + 4°C global warming.     

20.

Elevated atmospheric CO₂ alters stomatal responses to variable sunlight in a C₄ grass   总被引：1，自引：1，他引：0  

A. K. KNAPP  J. T. FAHNESTOCK  C. E. OWENSBY 《Plant, cell & environment》1994,17(2):189-195

Native tallgrass prairie in NE Kansas was exposed to elevated (twice ambient) or ambient atmospheric CO₂ levels in open-top chambers. Within chambers or in adjacent unchambered plots, the dominant C₄ grass, Andropogon gerardii, was subjected to fluctuations in sunlight similar to that produced by clouds or within canopy shading (full sun > 1500 μmol m⁻² s⁻¹ versus 350 μmol m⁻² s⁻¹ shade) and responses in gas exchange were measured. These field experiments demonstrated that stomatal conductance in A. gerardii achieved new steady state levels more rapidly after abrupt changes in sunlight at elevated CO₂ when compared to plants at ambient CO₂. This was due primarily to the 50% reduction in stomatal conductance at elevated CO₂, but was also a result of more rapid stomatal responses. Time constants describing stomatal responses were significantly reduced (29–33%) at elevated CO₂. As a result, water loss was decreased by as much as 57% (6.5% due to more rapid stomatal responses). Concurrent increases in leaf xylem pressure potential during periods of sunlight variability provided additional evidence that more rapid stomatal responses at elevated CO₂ enhanced plant water status. CO₂-induced alterations in the kinetics of stomatal responses to variable sunlight will likely enhance direct effects of elevated CO₂ on plant water relations in all ecosystems.     

Simple carbon assimilation response functions from atmospheric CO2, and daily temperature and shortwave radiation

A global ‘CO₂ fertilizer effect’ multiplier is often used in crop or ecosystem models because of its simplicity. However, this approach does not take into account the interaction between CO₂, temperature and light on assimilation. This omission can lead to significant under- or overestimation of the magnitude of beneficial effects from elevated CO₂, depending on environmental conditions. We use a mechanistic model of the biochemistry of photosynthesis to represent the response of net assimilation to different levels of CO₂, temperature and radiation, on the daily time scale. Instantaneous assimilation rates for an idealized canopy model are integrated through diurnal cycles of environmental variables derived from historical climate data at three locations in North America. The calculated CO₂ fertilizer effect is greatest at high light and warm temperatures. The results are summarized by assimilation response surfaces specified by the CO₂ concentration, the canopy leaf area index, and by daily values of temperature and radiation available from climatic records. These summary functions are suitable for incorporation into crop or ecosystem models for predicting carbon assimilation or biomass production on a daily time step. An example application of the function reveals that for a relatively cool, high latitude location, the beneficial effects from a CO₂ doubling would be negligible during the early spring, even assuming a + 4°C global warming scenario. In contrast, the beneficial effects from increasing CO₂ at a relatively warm, lower latitude location are greatest in the spring, but decline in late summer because of excessively warm temperatures with a + 4°C global warming.     

Elevated atmospheric CO2 alters stomatal responses to variable sunlight in a C4 grass

Native tallgrass prairie in NE Kansas was exposed to elevated (twice ambient) or ambient atmospheric CO₂ levels in open-top chambers. Within chambers or in adjacent unchambered plots, the dominant C₄ grass, Andropogon gerardii, was subjected to fluctuations in sunlight similar to that produced by clouds or within canopy shading (full sun > 1500 μmol m⁻² s⁻¹ versus 350 μmol m⁻² s⁻¹ shade) and responses in gas exchange were measured. These field experiments demonstrated that stomatal conductance in A. gerardii achieved new steady state levels more rapidly after abrupt changes in sunlight at elevated CO₂ when compared to plants at ambient CO₂. This was due primarily to the 50% reduction in stomatal conductance at elevated CO₂, but was also a result of more rapid stomatal responses. Time constants describing stomatal responses were significantly reduced (29–33%) at elevated CO₂. As a result, water loss was decreased by as much as 57% (6.5% due to more rapid stomatal responses). Concurrent increases in leaf xylem pressure potential during periods of sunlight variability provided additional evidence that more rapid stomatal responses at elevated CO₂ enhanced plant water status. CO₂-induced alterations in the kinetics of stomatal responses to variable sunlight will likely enhance direct effects of elevated CO₂ on plant water relations in all ecosystems.