The functional significance of the stomatal size to density relationship: Interaction with atmospheric [CO2] and role in plant physiological behaviour.

The limits for stomatal conductance are set by stomatal size (SS) and density (SD). An inverse relationship between SS and SD has been observed in fossil and living plants. This has led to hypotheses proposing that the ratio of SS to SD influences the diffusion pathway for CO2 and degree of physiological stomatal control. However, conclusive evidence supportive of a functional role of the SS-SD relationship is not evident, and patterns in SS-SD may simply reflect geometric constraints in stomatal spacing over a leaf surface. We examine published and new data to investigate the potential functional significance of the relationship between SS and SD to atmospheric [CO2] in multiple generation adaptive responses and short-term acclamatory adjustment of stomatal morphology. Consistent patterns in SS and SD were not evident in fossil and living plants adapted to high [CO2] over many generations. However, evolutionary adaptation to [CO2] strongly affected SS and SD responses to elevated [CO2], with plants adapted to the 'low' [CO2] of the past 10 million years (Myr) showing adjustment of SS-SD, while members of the same species adapted to 'high' [CO2] showed no response. This may suggest that SS and SD responses to future [CO2] will likely constrain the stimulatory effect of 'CO2-fertilisation' on photosynthesis. Angiosperms generally possessed higher densities of smaller stomata that corresponded to a greater degree of physiological stomatal control consistent with selective pressures induced by declining [CO2] over the past 90 Myr. Atmospheric [CO2] has likely shaped stomatal size and density relationships alongside the interaction with stomatal physiological behaviour. The rate and predicted extent of future increases in [CO2] will have profound impacts on the selective pressures shaping SS and SD. Understanding the trade-offs involved in SS-SD and the interaction with [CO2], will be central to the development of more productive climate resilient crops.

Lipid Peroxidation and Chlorophyll Fluorescence of Photosystem II Performance during Drought and Heat Stress is Associated with the Antioxidant Capacities of C3 Sunflower and C4 Maize Varieties.

Agricultural production is predicted to be adversely affected by an increase in drought and heatwaves. Drought and heat damage cellular membranes, such as the thylakoid membranes where photosystem II occurs (PSII). We investigated the chlorophyll fluorescence (ChlF) of PSII, photosynthetic pigments, membrane damage, and the activity of protective antioxidants in drought-tolerant and -sensitive varieties of C3 sunflower and C4 maize grown at 20/25 and 30/35 °C. Drought-tolerant varieties retained PSII electron transport at lower levels of water availability at both temperatures. Drought and heat stress, in combination and isolation, had a more pronounced effect on the ChlF of the C3 species. For phenotyping, the maximum fluorescence was the most effective ChlF measure in characterizing varietal variation in the response of both species to drought and heat. The drought-tolerant sunflower and maize showed lower lipid peroxidation under drought and heat stress. The greater retention of PSII function in the drought-tolerant sunflower and maize at higher temperatures was associated with an increase in the activities of antioxidants (glutathione reductase, superoxide dismutase, catalase, peroxidase, and ascorbate peroxidase), whereas antioxidant activity declined in the drought-sensitive varieties. Antioxidant activity should play a key role in the development of drought- and heat-tolerant crops for future food security.

Microbiomes in Soils Exposed to Naturally High Concentrations of CO2 (Bossoleto Mofette Tuscany, Italy).

Direct and indirect effects of extremely high geogenic CO2 levels, commonly occurring in volcanic and hydrothermal environments, on biogeochemical processes in soil are poorly understood. This study investigated a sinkhole in Italy where long-term emissions of thermometamorphic-derived CO2 are associated with accumulation of carbon in the topsoil and removal of inorganic carbon in low pH environments at the bottom of the sinkhole. The comparison between interstitial soil gasses and those collected in an adjacent bubbling pool and the analysis of the carbon isotopic composition of CO2 and CH4 clearly indicated the occurrence of CH4 oxidation and negligible methanogenesis in soils at the bottom of the sinkhole. Extremely high CO2 concentrations resulted in higher microbial abundance (up to 4 × 109 cell g-1 DW) and a lower microbial diversity by favoring bacteria already reported to be involved in acetogenesis in mofette soils (i.e., Firmicutes, Chloroflexi, and Acidobacteria). Laboratory incubations to test the acetogenic and methanogenic potential clearly showed that all the mofette soil supplied with hydrogen gas displayed a remarkable CO2 fixation potential, primarily due to the activity of acetogenic microorganisms. By contrast, negligible production of acetate occurred in control tests incubated with the same soils, under identical conditions, without the addition of hydrogen. In this study, we report how changes in diversity and functions of the soil microbial community - induced by high CO2 concentration - create peculiar biogeochemical profile. CO2 emission affects carbon cycling through: (i) inhibition of the decomposition of the organic carbon and (ii) promotion of CO2-fixation via the acetyl-CoA pathway. Sites naturally exposed to extremely high CO2 levels could potentially represent an untapped source of microorganisms with unique capabilities to catalytically convert CO2 into valuable organic chemicals and fuels.

Publisher Correction: Impaired photosynthesis and increased leaf construction costs may induce floral stress during episodes of global warming over macroevolutionary timescales.

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Impaired photosynthesis and increased leaf construction costs may induce floral stress during episodes of global warming over macroevolutionary timescales.

Global warming events have coincided with turnover of plant species at intervals in Earth history. As mean global temperatures rise, the number, frequency and duration of heat-waves will increase. Ginkgo biloba was grown under controlled climatic conditions at two different day/night temperature regimes (25/20 °C and 35/30 °C) to investigate the impact of heat stress. Photosynthetic CO2-uptake and electron transport were reduced at the higher temperature, while rates of respiration were greater; suggesting that the carbon balance of the leaves was adversely affected. Stomatal conductance and the potential for evaporative cooling of the leaves was reduced at the higher temperature. Furthermore, the capacity of the leaves to dissipate excess energy was also reduced at 35/30 °C, indicating that photo-protective mechanisms were no longer functioning effectively. Leaf economics were adversely affected by heat stress, exhibiting an increase in leaf mass per area and leaf construction costs. This may be consistent with the selective pressures experienced by fossil Ginkgoales during intervals of global warming such as the Triassic - Jurassic boundary or Early Eocene Climatic Optimum. The physiological and morphological responses of the G. biloba leaves were closely interrelated; these relationships may be used to infer the leaf economics and photosynthetic/stress physiology of fossil plants.

Testing decontaminated sediments as a substrate for ornamentals in field nursery plantations.

When canals and harbours are dredged, huge amount of polluted sediments has to be stocked and transported to the landfill with incredibly high costs of management. Among the remediation techniques for the reclamation of polluted sediments and soils, phytoremediation represents a sustainable and effective technique though still not fully promoted or commercialized. In this study we have tested the suitability for plant nursing of a substrate resulting from sediments dredged from a canal and treated with phytoremediation. The experiment was set up in 2014. It aimed to test the physical, chemical and hydrological characteristics of two mixes of remediated sediments and agronomic soil (at 33% and 50% by volume) compared to control soil (100% agronomic soil), and to assess the growth of three ornamental species (Viburnum tinus L., Photinia x fraseri var. red robin, Eleagnus macrophylla Thunb.) together with the suitability for root balling. The mixed substrates produced good results in terms of water drainage, and were similar to the control in terms of soluble nutrients, guaranteeing and enhancing the aboveground and belowground growth of all the three species, especially V. tinus. In contrast, mixed substrates impaired root ball compaction with root ball breakage observed especially in 50% sediment/soil mix. Therefore, the use of remediated sediments in plant nursery can be limited to specific productions or practices.

Adaptation to high temperature mitigates the impact of water deficit during combined heat and drought stress in C3 sunflower and C4 maize varieties with contrasting drought tolerance.

Heat and drought stress frequently occur together, however, their impact on plant growth and photosynthesis (PN ) is unclear. The frequency, duration and severity of heat and drought stress events are predicted to increase in the future, having severe implications for agricultural productivity and food security. To assess the impact on plant gas exchange, physiology and morphology we grew drought tolerant and sensitive varieties of C3 sunflower (Helianthus annuus) and C4 maize (Zea mays) under conditions of elevated temperature for 4 weeks prior to the imposition of water deficit. The negative impact of temperature on PN was most apparent in sunflower. The drought tolerant sunflower retained ribulose-1,5-bisphosphate carboxylase/oxygenase (RubisCO) activity under heat stress to a greater extent than its drought sensitive counterpart. Maize exhibited no varietal difference in response to increased temperature. In contrast to previous studies, where a sudden rise in temperature induced an increase in stomatal conductance (Gs ), we observed no change or a reduction in Gs with elevated temperature, which alongside lower leaf area mitigated the impact of drought at the higher temperature. The drought tolerant sunflower and maize varieties exhibited greater investment in root-systems, allowing greater uptake of the available soil water. Elevated temperatures associated with heat-waves will have profound negative impacts on crop growth in both sunflower and maize, but the deleterious effect on PN was less apparent in the drought tolerant sunflower and both maize varieties. As C4 plants generally exhibit water use efficiency (WUE) and resistance to heat stress, selection on the basis of tolerance to heat and drought stress would be more beneficial to the yields of C3 crops cultivated in drought prone semi-arid regions.

Impaired Stomatal Control Is Associated with Reduced Photosynthetic Physiology in Crop Species Grown at Elevated [CO2].

Physiological control of stomatal conductance (Gs) permits plants to balance CO2-uptake for photosynthesis (PN) against water-loss, so optimizing water use efficiency (WUE). An increase in the atmospheric concentration of carbon dioxide ([CO2]) will result in a stimulation of PN and reduction of Gs in many plants, enhancing carbon gain while reducing water-loss. It has also been hypothesized that the increase in WUE associated with lower Gs at elevated [CO2] would reduce the negative impacts of drought on many crops. Despite the large number of CO2-enrichment studies to date, there is relatively little information regarding the effect of elevated [CO2] on stomatal control. Five crop species with active physiological stomatal behavior were grown at ambient (400 ppm) and elevated (2000 ppm) [CO2]. We investigated the relationship between stomatal function, stomatal size, and photosynthetic capacity in the five species, and then assessed the mechanistic effect of elevated [CO2] on photosynthetic physiology, stomatal sensitivity to [CO2] and the effectiveness of stomatal closure to darkness. We observed positive relationships between the speed of stomatal response and the maximum rates of PN and Gs sustained by the plants; indicative of close co-ordination of stomatal behavior and PN. In contrast to previous studies we did not observe a negative relationship between speed of stomatal response and stomatal size. The sensitivity of stomata to [CO2] declined with the ribulose-1,5-bisphosphate limited rate of PN at elevated [CO2]. The effectiveness of stomatal closure was also impaired at high [CO2]. Growth at elevated [CO2] did not affect the performance of photosystem II indicating that high [CO2] had not induced damage to the photosynthetic physiology, and suggesting that photosynthetic control of Gs is either directly impaired at high [CO2], sensing/signaling of environmental change is disrupted or elevated [CO2] causes some physical effect that constrains stomatal opening/closing. This study indicates that while elevated [CO2] may improve the WUE of crops under normal growth conditions, impaired stomatal control may increase the vulnerability of plants to water deficit and high temperatures.

The impact of built-up surfaces on land surface temperatures in Italian urban areas.

Urban areas are characterized by the very high degree of soil sealing and continuous built-up areas: Italy is one of the European countries with the highest artificial land cover rate, which causes a substantial spatial variation in the land surface temperature (LST), modifying the urban microclimate and contributing to the urban heat island effect. Nevertheless, quantitative data regarding the contribution of different densities of built-up surfaces in determining urban spatial LST changes is currently lacking in Italy. This study, which aimed to provide clear and quantitative city-specific information on annual and seasonal spatial LST modifications resulting from increased urban built-up coverage, was conducted generally throughout the whole year, and specifically in two different periods (cool/cold and warm/hot periods). Four cities (Milan, Rome, Bologna and Florence) were included in the study. The LST layer and the built-up-surface indicator were obtained via use of MODIS remote sensing data products (1km) and a very high-resolution map (5m) of built-up surfaces recently developed by the Italian National Institute for Environmental Protection and Research. The relationships between the dependent (mean daily, daytime and nighttime LST values) and independent (built-up surfaces) variables were investigated through linear regression analyses, and comprehensive built-up-surface-related LST maps were also developed. Statistically significant linear relationships (p<0.001) between built-up surfaces and spatial LST variations were observed in all the cities studied, with a higher impact during the warm/hot period than in the cool/cold ones. Daytime and nighttime LST slope patterns depend on the city size and relative urban morphology. If implemented in the existing city plan, the urban maps of built-up-surface-related LST developed in this study might be able to support more sustainable urban land management practices by identifying the critical areas (Hot-Spots) that would benefit most from mitigation actions by local authorities, land-use decision makers, and urban planners.

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.

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

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 (r(2) = 0.90) and GPP recovery after a fire disturbance in South Dakota (r(2) = 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.

Carbon dioxide fertilisation and supressed respiration induce enhanced spring biomass production in a mixed species temperate meadow exposed to moderate carbon dioxide enrichment.

The rising concentration of carbon dioxide in the atmosphere ([CO2]) has a direct effect on terrestrial vegetation through shifts in the rates of photosynthetic carbon uptake and transpirational water-loss. Free Air CO2 Enrichment (FACE) experiments aim to predict the likely responses of plants to increased [CO2] under normal climatic conditions. The Giessen FACE system operates a lower [CO2] enrichment regime (480µmolmol-1) than standard FACE (550-600µmolmol-1), permitting the analysis of a mixed species temperate meadow under a [CO2] level equivalent to that predicted in 25-30 years. We analysed the physiological and morphological responses of six species to investigate the effect of moderate [CO2] on spring biomass production. Carbon dioxide enrichment stimulated leaf photosynthetic rates and supressed respiration, contributing to enhanced net assimilation and a 23% increase in biomass. The capacity for photosynthetic assimilation was unaffected by [CO2] enrichment, with no downregulation of rates of carboxylation of Rubisco or regeneration of ribulose-1,5-bisphosphate. Foliar N content was also not influenced by increased [CO2]. Enhanced [CO2] reduced stomatal size, but stomatal density and leaf area index remained constant, suggesting that the effect on gas exchange was minimal.

Stomatal index responses of Agrostis canina to CO2 and sulphur dioxide: implications for palaeo-[CO2] using the stomatal proxy.

• Stomatal index values of fossil plants are widely used in reconstructing palaeo-[CO(2)]. This depends upon the assumption that the stomatal index is determined by the atmospheric concentration of CO(2) ([CO(2)]). This study investigates whether fumigation with, and resistance to, sulphur dioxide (SO(2)) induces a reduction in the stomatal index that may affect stomatal reconstructions of palaeo-[CO(2)] coinciding with episodes of global-scale volcanism. • Agrostis canina from Mefite di Ansanto, Italy, grow in atmospheres of elevated-[CO(2)], SO(2) and hydrogen sulphide (H(2)S). Mefite A. canina were compared with a control population in a 'common-garden' experiment and a controlled-environment study under elevated-[CO(2)] and SO(2) fumigation. • In A. canina, resistance to toxic volcanic gases is not associated with reduced stomatal index, and fumigation with SO(2) does not cause a decrease in stomatal initiation. The two populations of A. canina analyzed in this study exhibit different stomatal index-[CO(2)] 'responses', with control plants showing a reduction in stomatal index and Mefite plants showing no response. • Stomatal reconstructions of palaeo-[CO(2)] during past episodes of global-scale volcanism probably reflect atmospheric [CO(2)] and not [SO(2)]. The lack of a reduction in the stomatal index in response to elevated [CO(2)] in the Mefite plants, suggests that resistance to toxic gases and/or long-term growth at high [CO(2)] reduces, or negates, sensitivity of the stomatal index-[CO(2)] relationship, or that stomatal index-[CO(2)] in the Mefite plants is attuned to [CO(2)] fluctuations at much higher concentrations.

Soil Respiration in European Grasslands in Relation to Climate and Assimilate Supply.

Soil respiration constitutes the second largest flux of carbon (C) between terrestrial ecosystems and the atmosphere. This study provides a synthesis of soil respiration (R(s)) in 20 European grasslands across a climatic transect, including ten meadows, eight pastures and two unmanaged grasslands. Maximum rates of R(s) (R(s(max) )), R(s) at a reference soil temperature (10°C; R(s(10) )) and annual R(s) (estimated for 13 sites) ranged from 1.9 to 15.9 µmol CO(2) m(-2) s(-1), 0.3 to 5.5 µmol CO(2) m(-2) s(-1) and 58 to 1988 g C m(-2) y(-1), respectively. Values obtained for Central European mountain meadows are amongst the highest so far reported for any type of ecosystem. Across all sites R(s(max) ) was closely related to R(s(10) ).Assimilate supply affected R(s) at timescales from daily (but not necessarily diurnal) to annual. Reductions of assimilate supply by removal of aboveground biomass through grazing and cutting resulted in a rapid and a significant decrease of R(s). Temperature-independent seasonal fluctuations of R(s) of an intensively managed pasture were closely related to changes in leaf area index (LAI). Across sites R(s(10) ) increased with mean annual soil temperature (MAT), LAI and gross primary productivity (GPP), indicating that assimilate supply overrides potential acclimation to prevailing temperatures. Also annual R(s) was closely related to LAI and GPP. Because the latter two parameters were coupled to MAT, temperature was a suitable surrogate for deriving estimates of annual R(s) across the grasslands studied. These findings contribute to our understanding of regional patterns of soil C fluxes and highlight the importance of assimilate supply for soil CO(2) emissions at various timescales.

Leaf traits and tree rings suggest different water-use and carbon assimilation strategies by two co-occurring Quercus species in a Mediterranean mixed-forest stand in Tuscany, Italy.

We compared the water-use characteristics of co-occurring mature Quercus cerris L. and Quercus pubescens Willd. trees growing in resource-limited (mainly water) hilly habitats in Tuscany, Italy. The species differed in their distribution along soil water gradients and in their access to, and use of, water, even though the study year was wetter than average, though with a summer drought. Compared with Q. cerris, Q. pubescens had greater access to soil water (less negative predawn water potentials) and a more conservative water-use strategy based on its relatively low stomatal conductance, high instantaneous water-use efficiency, less negative midday water potential and high soil-to-leaf hydraulic conductance. Quercus cerris had less conservative water-use characteristics than Q. pubescens, exhibiting relatively high stomatal conductance, low instantaneous water-use efficiency, more negative midday water potentials and low soil-to-leaf hydraulic conductance; however, Q. cerris had higher photosynthetic rates than Q. pubescens. Photosynthesis and stomatal conductance were positively correlated in both species. Although a strong correlation between ring widths and precipitation patterns was not found, some dry periods influenced ring-width growth. Quercus pubescens has always grown faster than Q. cerris, probably because of more efficient water use, although stand dynamics (driven by exogenous disturbance factors, including coppicing, browsing and competition) cannot be excluded. Ring-width variability, as well as tree-ring growth in dry years, which should be unaffected by stand dynamics, were higher in Q. pubescens than in Q. cerris. Moreover, Q. pubescens recovered completely after the drought in the seventies, even showing higher tree-ring growth than in the recent past, whereas Q. cerris showed a minor growth decline followed by a recovery to values comparable with those observed before the 1970s drought. Beginning in the early eighties, tree-ring growth decreased in both species, though Q. pubescens showed consistently higher values than Q. cerris. These differences can be explained by differences in water-use efficiency. Despite differences between the species in water use and water status, the results are consistent with the interpretation that both are drought tolerant, but that Q. pubescens is at an advantage on xeric ridges because of its greater ability to access soil water and use it more conservatively compared with Q. cerris.

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.

Seasonal changes in tissue elasticity and water transport efficiency in three co-occurring Mediterranean shrubs under natural long-term CO2 enrichment.

Seasonal changes in hydraulic properties and tissue elasticity were evaluated in Erica arborea L., Myrtuscommunis L. and Juniperus communis L., three Mediterranean shrubs that differ in adaptations to drought. These parameters were analysed over 12 months under field conditions, by comparing plants grown in the proximity of a natural CO2 spring (about 700 µmol mol-1 atmospheric CO2 concentration, [CO2]) with plants in ambient conditions. Plants at the CO2-spring site have been growing for generations at elevated [CO2]. At both sites, stem hydraulic and structural properties followed the prevailing climatic constraints. However, these shrub species co-occurring in the same environment differed in their capacity to tolerate water deficits, in xylem efficiency, and in strategies for regulating water movement between plant compartments. Either an increase or a decrease in tissue elasticity was effective in promoting resistance to drought stress, depending on the species. Long-term elevated [CO2] influenced all the studied parameters. Species-dependent differences existed in hydraulic architecture between the CO2-spring plants and control plants of E. arborea and M. communis, while J. communis plants rarely showed differences between sites. Less distinct differences between sites were observed for wood structure. The three species showed somewhat lower tissue elasticity under elevated [CO2], in particular during stress periods. The effects of elevated [CO2] on stem hydraulic pathway and structure and shoot elastic properties persist in the long term, but differ in absolute values and sign among the studied species and with the seasonal course, and thus might alter competitive relations between these shrubs.

Water relations, stomatal response and transpiration of Quercus pubescens trees during summer in a Mediterranean carbon dioxide spring.

Variations in water relations and stomatal response of Quercus pubescens Willd. were analyzed under Mediterranean field conditions during two consecutive summers (1993 and 1994) at two locations characterized by different atmospheric CO(2) concentrations because of the presence at one of them of a CO(2) spring. Trees at the CO(2) spring site have been growing for generations in elevated atmospheric CO(2) concentrations. The heat-pulse velocity technique was used to estimate water use of trees during a 5-month period from June to November 1994. At the end of the sap flow measurements, the trees were harvested and foliage and sapwood area measured. At both sites, maximum leaf conductance was related to predawn shoot water potential. Effects of summer drought on plant water relations, including whole-plant transpiration, were severe, but leaf conductance and water potential recovered to predrought values after major rainfall in September. Leaf conductance, predawn water potential, and sometimes sap flow, decreased in parallel with increases in hydraulic resistance, reaching a minimum in midsummer. Hydraulic resistance was higher in trees at the control site than in trees at the CO(2) spring site. The effect of elevated CO(2) concentration on leaf conductance was less at high leaf-to-air water vapor pressure difference than at low leaf-to-air water vapor pressure difference. Mean and diurnal sap fluxes were consistently higher in trees at the control site than in trees at the CO(2) spring site. During the summer period, plant water use varied between the two sites. Trees at the control site had consistently higher sap flow at corresponding values of sapwood cross-sectional area than trees at the CO(2) spring site. Because trees at the CO(2) spring site supported a smaller foliage area for a corresponding sapwood cross-sectional area than trees at the control site, the overall mean sap flux/foliage area ratio did not differ between sites. The results are discussed in terms of effects of elevated CO(2) concentration on plant water use at the organ and whole-tree scale.

Seasonal embolism and xylem vulnerability in deciduous and evergreen Mediterranean trees influenced by proximity to a carbon dioxide spring.

We investigated how proximity to natural CO(2) springs affected the seasonal patterns of xylem embolism in Quercus ilex L., Quercus pubescens Willd., Fraxinus ornus L., Populus tremula L. and Arbutus unedo L., which differ in leaf phenology and wood anatomy. Xylem embolism was evaluated in both artificially dehydrated branches and in hydrated apical branches collected at monthly intervals during a 20-month sampling period. Initial specific hydraulic conductivity was also evaluated. We found species-dependent differences in xylem hydraulic properties in response to elevated CO(2) concentration. Populus tremula was the most embolized and A. unedo was the least embolized of the species examined. Effects of elevated CO(2) were significant in Q. pubescens, P. tremula and A. unedo, whereas the overall response to elevated CO(2) was less evident in F. ornus and Q. ilex. Specific hydraulic conductivity differed among species but not between sites, although the interaction between species and site was significant. Differences in xylem vulnerability between trees growing near to the CO(2) spring and those growing in control areas were small. Although differences in hydraulic properties in response to elevated CO(2) concentration were small, they may be of great importance in determining future community composition in Mediterranean-type forest ecosystems. The possible causes and ecological significance of such differences are discussed in relation to elevated CO(2) concentration and other environmental conditions.

Influence of increased atmospheric CO(2) concentration on quality of plant material and litter decomposition.

Nitrogen (N) and lignin concentrations in plant tissues and litter of plants grown in greenhouses or open-top chambers in elevated atmospheric CO(2) concentration were compared with those of plants grown in ambient air in short-term studies. We also compared the N concentration of plant material of Quercus ilex L. and Q. pubescens Willd. growing in the vicinity of natural CO(2)-springs with that of the same species growing at a control site. In the short-term studies, elevated CO(2) caused significant decreases in tissue N concentration and the extent of the decrease varied with species. Nitrogen amendment of the soil lessened the CO(2)-enrichment effect. Lignin concentration was modified by elevated CO(2) and the effect was species specific, but no general positive or negative trend was evident. A comparison of trees growing under natural conditions near a natural CO(2)-spring and at a control site revealed no site differences in N concentration of the plant material. A comparison of published results on decomposition rates of litter produced in elevated atmospheric CO(2) and in ambient air indicated that CO(2) enrichment can cause both enhancements and decreases of carbon mineralization. We conclude that (1) long-term responses to elevated CO(2) could differ from the results obtained from short-term studies and that (2) biodiversity could be an important factor altering the sign of the feedback on atmospheric CO(2) concentration. We also discuss the implications of our finding of a long-term, inhibitory effect of the initial N concentration of litter on the decomposition rate of litter and its consequence on ecosystem feedback.