Antonio Raschi

Joint control of terrestrial gross primary productivity by plant phenology and physiology

Significance Terrestrial gross primary productivity (GPP), the total photosynthetic CO2 fixation at ecosystem level, fuels all life on land. However, its spatiotemporal variability is poorly understood, because GPP is determined by many processes related to plant phenology and physiological activities. In this study, we find that plant phenological and physiological properties can be integrated in a robust index—the product of the length of CO2 uptake period and the seasonal maximal photosynthesis—to explain the GPP variability over space and time in response to climate extremes and during recovery after disturbance. Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate–carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy–covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO2 uptake period (CUP) and the seasonal maximal capacity of CO2 uptake (GPPmax). The product of CUP and GPPmax explained >90% of the temporal GPP variability in most areas of North America during 2000–2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r2 = 0.90) and GPP recovery after a fire disturbance in South Dakota (r2 = 0.88). Additional analysis of the eddy–covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPPmax than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPPmax and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space.

Response of foliar metabolism in mature trees of Quercus pubescens and Quercus ilex to long-term elevated CO2

Long-term effects on and adaptations of the carbon physiology of long-lived trees exposed to increasing atmospheric levels of CO2 are unknown. We compared two indigenous Quercus species, Q. ilex and Q. pubescens, growing in a natural CO2 spring located in central Italy and at a nearby control site. In May, 1995 photosynthetic rate at least doubled when measured with supplemental CO2 in both species and sites. Dark respiration was much higher at the CO2 spring site in both species. Foliar sugar and starch concentrations in Q. ilex exhibited significant site and diurnal differences (May and September). In July, 1995 there was little difference in the water potential values of the measured trees at the different sites over the diurnal period. Photosynthetic rate was higher for both species in the CO2 spring, particularly in the early morning and late afternoon. Mid-day stomatal closure reduced photosynthesis to similar levels. In the morning leaf conductance and transpiration were generally lower in the CO2 spring trees, contributing to higher instantaneous water use efficiency for both species. Isoprene emission rates were higher in Q. pubescens trees growing in the CO2 spring. The maximum difference between control and CO2 spring trees occurred in late afternoon. In contrast, Q. ilex exhibited isoprene emission near background level. Foliage and branch carbon and nitrogen status showed increased concentrations of starch and tannins in Q. ilex and of soluble sugars in Q. pubescens in the elevated CO2 environment, while nitrogen concentration decreased in both species. Wood gravity increased 6 and 3% in Q. ilex and Q. pubescens, respectively, growing in the CO2 spring. Q. ilex exhibited afternoon recovery of water potential compared to Q. pubescens which had better night-time recovery. Q. ilex and Q. pubescens exposed to elevated CO2 for prolonged periods exhibit different mechanisms for dealing with additional reduced carbon and do maintain an altered carbon physiology, even in midst of the regions characteristic summer drought.

Thermal optimality of net ecosystem exchange of carbon dioxide and underlying mechanisms.

• It is well established that individual organisms can acclimate and adapt to temperature to optimize their functioning. However, thermal optimization of ecosystems, as an assemblage of organisms, has not been examined at broad spatial and temporal scales. • Here, we compiled data from 169 globally distributed sites of eddy covariance and quantified the temperature response functions of net ecosystem exchange (NEE), an ecosystem-level property, to determine whether NEE shows thermal optimality and to explore the underlying mechanisms. • We found that the temperature response of NEE followed a peak curve, with the optimum temperature (corresponding to the maximum magnitude of NEE) being positively correlated with annual mean temperature over years and across sites. Shifts of the optimum temperature of NEE were mostly a result of temperature acclimation of gross primary productivity (upward shift of optimum temperature) rather than changes in the temperature sensitivity of ecosystem respiration. • Ecosystem-level thermal optimality is a newly revealed ecosystem property, presumably reflecting associated evolutionary adaptation of organisms within ecosystems, and has the potential to significantly regulate ecosystem-climate change feedbacks. The thermal optimality of NEE has implications for understanding fundamental properties of ecosystems in changing environments and benchmarking global models.

Calibration and application of FOREST-BGC in a Mediterranean area by the use of conventional and remote sensing data

The current work deals with the use in a Mediterranean environment of a simulation model of forest ecosystem processes which was originally created for temperate areas (FOREST-BGC). The model was calibrated and applied on two deciduous forest stands in Tuscany (Central Italy) by using conventional and remote sensing data as inputs. First, information on the two stands needed to initialise the model was derived from different sources, while meteorological data were extrapolated from a nearby station by an existing procedure (MT-Clim). Temporal profiles of leaf area index (LAI) were then derived both from direct ground measurement and from the processing of NOAA-AVHRR NDVI data. The model was calibrated using stand transpiration values obtained for 1997 by a sap flow method. Next, its performances were tested against the same transpiration values measured in 1998. The results obtained indicate that FOREST-BGC is capable of simulating water fluxes of Mediterranean forests when suitable LAI profiles are considered. Moreover, the derivation of these profiles from NDVI data can improve the model performance probably due to an enhanced consideration of the effects of the typical Mediterranean summer water stress. These results support the final objective of the work, which is the development of a procedure capable of integrating conventional and remote sensing data to operationally simulate water and carbon fluxes on a regional scale.

Carbon dioxide emissions at an Italian mineral spring: measurements of average CO2 concentration and air temperature

Abstract Emissions of carbon dioxide from vents at the Bossoleto mineral spring in Central Italy have been calculated to exceed 12 t day −1 . This emission leads to enhanced atmospheric concentrations of CO 2 over an area of more than 3000 m 2 . The vent gas is over 99% pure CO 2 , with a characteristic isotopic signature that is totally depleted in 14 C. At night, concentrations at the bottom of the bowl-like depression can increase to levels approaching 75%. In the morning, this high concentration of CO 2 is associated with a rapid temperature increase of over 10°C before the CO 2 disperses. This site is being used in a number of studie of the response of plant communities to long-term enhanced CO 2 concentrations. The problem of defining CO 2 concentrations in these studies was approached by comparing estimates determined by gas analysis measurements and isotopic analysis of leaf material. The isotopic method used 14 C as a tracer, integrating effective concentration over the life of a leaf by calculating from the ratio of 14 C measurements of plant material growing near the spring and at a control site. The estimates obtained using isotopic analysis of leaf material were similar to gas analysis measurements obtained during the day. This suggests that plants at this site are responding to the concentrations during the day, rather than the much higher night-time concentrations, making the system useful for biological research.

Physiological and morphological responses of grassland species to elevated atmospheric CO2 concentrations in FACE-systems and natural CO2 springs

Stomatal density, leaf conductance and water relations can be affected by an increase in the concentration of atmospheric CO2, and thus affect plant productivity. However, there is uncertainty about the effects of elevated CO2 on stomatal behaviour, water relations and plant productivity, owing to the lack of long-term experiments in representative natural ecosystems. In this work, variations in stomatal density and index, leaf water relations and plant biomass of semi-natural grassland communities were analysed under field conditions by comparing plants in three different experimental set-ups (natural CO2 springs, plastic tunnels and mini-FACE systems). Natural degassing vents continuously expose the surrounding vegetation to truly long-term elevated CO2 and can complement short-term manipulative experiments. Elevated CO2 concentration effects on stomata persist in the long term, though different species growing in the same environment show species-specific responses. The general decrease in stomatal conductance after exposure to elevated CO2 was not associated with clear changes in stomatal number on leaf surfaces. The hypothesis of long-term adaptive modifications to stomatal number and distribution of plants exposed to elevated CO2 was not supported by these experiments on grassland communities. Elastic cell wall properties were affected to some extent by elevated CO2. Above-ground biomass did not vary between CO2 treatments, leaf area index did not compensate for reduced stomatal conductance, and the root system had potentially greater soil exploration capacity. Considerable between-species variation in response to elevated CO2 may provide a mechanism for changing competitive interactions among plant species.

Interactions between drought and elevated CO2 on alfalfa plants

Summary Alfalfa ( Medicago sativa L.) plants were grown in open top chambers at ambient (340 ppm) and high (600 ppm) CO 2 concentrations. Twenty-five days after the first cutting one set of both plants was subjected to water deficit conditions by withholding water for 5 days. A chamber effect on proteolytic activity, monogalactosyl diacylglycerol to digalactosyl diacylglycerol molar ratio, total non-structural carbohydrates and soluble protein contents occurred. In contrast, no change in leaf water potential was observed between plants grown outdoors and inside the chambers. Plants grown at high CO 2 concentration showed a lower decrease in leaf water potential in comparison with plants grown at atmospheric CO 2 when subjected to water stress. Under high CO 2 concentration leaf nitrogen content decreased whereas starch accumulation and a higher proteolytic activity were recorded. Following water depletion, CO 2 -enriched plants showed a decrease in total non-structural carbohydrates and soluble proteins. In thylakoid membranes high CO 2 caused an increase in chlorophyll and lipid contents and a degradation of monogalactosyl diacylglycerol. A higher degree of unsaturation in the main thylakoid lipids was also observed. CO 2 -enriched plants were less affected by water stress as shown by reduced chlorophyll degradation and a higher membrane stability.

Interaction Between Drought and Elevated CO2 in the Response of Alfalfa Plants to Oxidative Stress

Summary Alfalfa ( Medicago sativa L.) plants were grown for 2 years in open top chambers both at ambient (340 ppm) and at high CO 2 concentrations (600 ppm). During the second-year regrowth cycle, plants were subjected to drought for 5 days. A chamber effect on the oxidative status of cells, total asorbate contents, glutathione reductase, and Ca 2+ ATPase and Ca 2+ ITPase activities occurred. Ascorbate was found to be more affected than glutathione by CO 2 enrichment and an interaction between CO 2 and drought prevented the decrease in ascorbate/dehydroascorbate ratio and increased the reduced glutathione/oxidised glutathione ratio as compared with ambient-CO 2 plants. In the microsomal fraction the presence of multiple Ca 2+ pumps, i.e. the plasma membrane-type and endoplasmic reticulum-type pumps, was highlighted by their differential substrate preference for ATP and ITP, though only one major peptide of 130 kD was recognised by an antibody against Ca 2+ ATPase. The pump was degraded in CO 2 -enriched plants and its preference for ATP was reduced. The activity of the Ca 2+ pump was also less susceptible to oxidation in drought conditions. CO 2 -enriched plants were less affected by drought as shown by a higher reducing status of their cells and by a reduced requirement for Ca 2+ ATPase activity to maintain Ca 2+ homeostasis.

Responses of two olive tree (Olea europaea L.) cultivars to elevated CO2 concentration in the field

Five-year-old plants of two olive cultivars (Frantoio and Moraiolo) grown in large pots were exposed for 7 to 8 months to ambient (AC) or elevated (EC) CO2 concentration in a free-air CO2 enrichment (FACE) facility. Exposure to EC enhanced net photosynthetic rate (PN) and decreased stomatal conductance, leading to greater instantaneous transpiration efficiency. Stomata density also decreased under EC, while the ratio of intercellular (Ci) to atmospheric CO2 concentration and chlorophyll content did not differ, except for the cv. Moraiolo after seven months of exposure to EC. Analysis of the relationship between photosynthesis and Ci indicated no significant change in carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxygenase after five months of exposure to EC. Based on estimates derived from the PN-Ci relationship, there were no apparent treatment differences in daytime respiration, CO2 compensation concentration, CO2-saturated photosynthetic rate, or photosynthetic rate at the mean Ci, but there was a reduction in stomata limitation to PN at EC. Thus 5-year-old olive trees did not exhibit down regulation of leaf-level photosynthesis in their response to EC, though some indication of adjustment was evident for the cv. Frantoio with respect to the cv. Moraiolo.

Coordination of stomatal physiological behavior and morphology with carbon dioxide determines stomatal control

PREMISE OF THE STUDY Stomatal control is determined by the ability to alter stomatal aperture and/or the number of stomata on the surface of new leaves in response to growth conditions. The development of stomatal control mechanisms to the concentration of CO₂within the atmosphere ([CO₂]) is fundamental to our understanding of plant evolutionary history and the prediction of gas exchange responses to future [CO₂]. METHODS In a controlled environment, fern and angiosperm species were grown in atmospheres of ambient (400 ppm) and elevated (2000 ppm) [CO₂]. Physiological stomatal behavior was compared with the stomatal morphological response to [CO₂]. KEY RESULTS An increase in [CO₂] or darkness induced physiological stomatal responses ranging from reductions (active) to no change (passive) in stomatal conductance. Those species with passive stomatal behavior exhibited pronounced reductions of stomatal density in new foliage when grown in elevated [CO₂], whereas species with active stomata showed little morphological response to [CO₂]. Analysis of the physiological and morphological stomatal responses of a wider range of species suggests that patterns of stomatal control to [CO₂] do not follow a phylogenetic pattern associated with plant evolution. CONCLUSIONS Selective pressures may have driven the development of divergent stomatal control strategies to increased [CO₂]. Those species that are able to actively regulate guard cell turgor are more likely to respond to [CO₂] through a change in stomatal aperture than stomatal number. We propose a model of stomatal control strategies in response to [CO₂] characterized by a trade-off between short-term physiological behavior and longer-term morphological response.