Natural CO2 springs in Italy: a resource for examining long-term response of vegetation to rising atmospheric CO2 concentrations

It is estimated that more than 100 geothermal CO₂ springs exist in central-western Italy. Eight springs were selected in which the atmospheric CO₂ concentrations were consistently observed to be above the current atmospheric average of 354μmol mol^-1. CO₂ concentration measurements at some of the springs are reported. The springs are described, and their major topographic and vegetational features are reported. Preliminary observations made on natural vegetation growing around the gas vents are then illustrated. An azonal pattern of vegetation distribution occurs around every CO₂ spring regardless of soil type and phytoclimatic areas. This is composed of pioneer populations of a Northern Eurasiatic species (Agrostis canina L.) which is often associated with Scirpus lacustris L. The potential of these sites for studying the long-term response of vegetation to rising atmospheric CO₂ concentrations is discussed.     

Elevated atmospheric CO2: a nurse plant substitute for oak seedlings establishing in old fields

It has been hypothesized that elevated levels of atmospheric CO₂ (eCO₂) may facilitate the encroachment of woody plants into grasslands by reducing water stress. In east‐central Minnesota, sandy soils frequently create drought conditions for plants, and water limitation inhibits the establishment of oaks into old fields situated on these soils. Some have argued that eCO₂ should slow secondary succession by favoring fast‐growing early successional species. However, if oak encroachment into old fields is being inhibited by water stress, then eCO₂ could accelerate old‐field succession in this region. The purpose of this study was to test the hypothesis that eCO₂ will increase the establishment success of oak seedlings in an old field environment. The study was conducted with CO₂ levels controlled by free air CO₂ enrichment (FACE). In May 1999, four oak (Quercus ellipsoidalis) acorns were planted in each of 24 plots in each of six experimental FACE rings (n= 576), three of which received elevated levels (550 ppm) of CO₂. Half the plots in each ring were weeded during the first three summers of the experiment. In summer 2000, water input was manipulated during a 3.5‐week period, during which half the plots received regular watering while the other half received no water. Summer 2001 was dry, receiving 35% less rainfall than the mean level. Under hot and dry conditions, eCO₂ increased soil water levels in unweeded plots and enhanced oak establishment (survival and growth) in weeded plots. In 2006, after the eighth growing season following planting, survival was five times greater under elevated than ambient CO₂. The results showed that under hot and dry conditions, eCO₂ can act like a nurse plant for tree seedlings growing in bare and unshaded areas, increasing seedling survival and growth, and thereby expanding the establishment window for trees encroaching into a grassland environment.     

Effects of elevated atmospheric CO2 on net ecosystem CO2 exchange of a scrub–oak ecosystem

We report the results of a 2‐year study of effects of the elevated (current ambient plus 350 μmol CO₂ mol^?1) atmospheric CO₂ concentration (C_a) on net ecosystem CO₂ exchange (NEE) of a scrub–oak ecosystem. The measurements were made in open‐top chambers (OTCs) modified to function as open gas‐exchange systems. The OTCs enclosed samples of the ecosystem (ca. 10 m² surface area) that had regenerated after a fire, 5 years before, in either current ambient or elevated C_a. Throughout the study, elevated C_a increased maximum NEE (NEE_max) and the apparent quantum yield of the NEE (φ_NEE) during the photoperiod. The magnitude of the stimulation of NEE_max, expressed per unit ground area, was seasonal, rising from 50% in the winter to 180% in the summer. The key to this stimulation was effects of elevated C_a, and their interaction with the seasonal changes in the environment, on ecosystem leaf area index, photosynthesis and respiration. The separation of these factors was difficult. When expressed per unit leaf area the stimulation of the NEE_max ranged from 7% to 60%, with the increase being dependent on increasing soil water content (W_soil). At night, the CO₂ effluxes from the ecosystem (NEE_night) were on an average 39% higher in elevated C_a. However, the increase varied between 6% and 64%, and had no clear seasonality. The partitioning of NEE_night into its belowground (R_below) and aboveground (R_above) components was carried out in the winter only. A 35% and 27% stimulation of NEE_night in December 1999 and 2000, respectively, was largely due to a 26% and 28% stimulation of R_below in the respective periods, because R_below constituted ca. 87% of NEE_night. The 37% and 42% stimulation of R_above in December 1999 and 2000, respectively, was less than the 65% and 80% stimulation of the aboveground biomass by elevated C_a at these times. An increase in the relative amount of the aboveground biomass in woody tissue, combined with a decrease in the specific rate of stem respiration of the dominant species Quercus myrtifolia in elevated C_a, was responsible for this effect. Throughout this study, elevated C_a had a greater effect on carbon uptake than on carbon loss, in terms of both the absolute flux and relative stimulation. Consequently, for this scrub–oak ecosystem carbon sequestration was greater in the elevated C_a during this 2‐year study period.     

Carbon gains by desiccation-tolerant plants at elevated CO2

1. There have been no reports of the long-term responses of the desiccation-tolerant (DT) plants to elevated CO₂. Xerophyta scabrida is a DT woody shrub, which loses chlorophylls and thylakoids during desiccation: a so-called poikilochlorophyllous desiccation-tolerant species (PDT). When the leaves of X. scabria are allowed to desiccate, the species shows many of the normal features of (P)DT plants.
2. However, the duration of photosynthesis in X. scabria is prolonged by 300% when the measurements are made at 700 as opposed to 350p.p.m. CO₂. The implication is that the carboxylating enzymes must still have been active at this time to enable appreciable photosynthetic activity. This response could have far-reaching implications for the success of such species in a future climate.
3. Lichens and mosses, representing the homoiochlorophyllous DTs (HDT), retain their chlorophyll content and photosynthetic apparatus during desiccation. We show the desiccation responses of two common HDT species ( Cladonia convoluta and Tortula ruralis ) to elevated CO₂ for comparison. Both HDT species showed increased net CO₂ uptake in the material grown at high CO₂ by more than 30% in moss and by more than 50% in lichen. It is concluded that desiccation-tolerant plants will be among the main beneficiaries of a high CO₂ future.     

Interactive effects of elevated [CO2] and drought on cherry (Prunus avium) seedlings I. Growth, whole-plant water use efficiency and water loss

Seeds of cherry ( Prunus avium ) were germinated and grown for two growing seasons in ambient (∼350 μmol mol⁻¹) or elevated (ambient+∼350 μmol mol⁻¹) CO₂ mole fractions in six open-top chambers. The seedlings were fertilized once a week, following Ingestad principles in order to supply mineral nutrients at free-access rates. In the first growing season gradual drought was imposed on rapidly growing cherry seedlings by withholding water for a 6-wk drying cycle. In the second growing season, the rapid onset of drought was imposed at the height of the growing season on the seedlings which had already experienced drought in the first growing season. Elevated [CO₂] significantly increased total dry-mass production in both water regimes, but did not ameliorate the growth response to drought of the cherry seedlings subjected to two sequential drying cycles. Water loss did not differ in either well watered or droughted seedlings between elevated and ambient [CO₂]; consequently whole-plant water- use efficiency (the ratio of total dry mass produced to total water consumption) was significantly increased. Similar patterns of carbon allocation between shoot and root were found in elevated and ambient [CO₂] when the seedlings were the same size. Thus, elevated [CO₂] did not improve drought tolerance, but it accelerated ontogenetic development irrespective of water status.     

Reduced water repellency of a grassland soil under elevated atmospheric CO2

Water repellency is a widespread characteristic of soils that can modify soil moisture content and distribution and is implicated in important processes such as aggregation and carbon sequestration. Repellency arises as a consequence of organic matter inputs; as elevated atmospheric CO₂ is known to modify such inputs, we tested the repellency of a grassland soil after 5 years of exposure to elevated CO₂ in a free air carbon dioxide enrichment experiment. Using a water droplet penetration time test, we found a significant reduction in repellency at elevated CO₂ in samples at field moisture content. As many of the processes potentially influenced by repellency have been shown to be modified at elevated CO₂ (e.g. soil aggregation, C sequestration, recruitment from seed), we suggest that further exploration of this phenomenon could enhance our understanding of CO₂ effects on ecosystem function. The mechanism responsible for the change in repellency has not been identified.     

N2 fixation by Acacia species increases under elevated atmospheric CO2

In the present study the effect of elevated CO₂ on growth and nitrogen fixation of seven Australian Acacia species was investigated. Two species from semi‐arid environments in central Australia (Acacia aneura and A. tetragonophylla) and five species from temperate south‐eastern Australia (Acacia irrorata, A. mearnsii, A. dealbata, A. implexa and A. melanoxylon) were grown for up to 148 d in controlled greenhouse conditions at either ambient (350 µmol mol^?1) or elevated (700 µmol mol^?1) CO₂ concentrations. After establishment of nodules, the plants were completely dependent on symbiotic nitrogen fixation. Six out of seven species had greater relative growth rates and lower whole plant nitrogen concentrations under elevated versus normal CO₂. Enhanced growth resulted in an increase in the amount of nitrogen fixed symbiotically for five of the species. In general, this was the consequence of lower whole‐plant nitrogen concentrations, which equate to a larger plant and greater nodule mass for a given amount of nitrogen. Since the average amount of nitrogen fixed per unit nodule mass was unaltered by atmospheric CO₂, more nitrogen could be fixed for a given amount of plant nitrogen. For three of the species, elevated CO₂ increased the rate of nitrogen fixation per unit nodule mass and time, but this was completely offset by a reduction in nodule mass per unit plant mass.     

Sex-specific physiological and growth responses to elevated atmospheric CO2 in Silene latifolia Poiret

Dioecy is found in nearly half of the angiosperm families, but little is known about how rising atmospheric CO₂ concentration will affect male and female individuals of dioecious species. We examined gender‐specific physiological and growth responses of Silene latifolia Poiret, a widespread dioecious species, to a doubled atmospheric CO₂ concentration in environmentally controlled growth chambers. Elevated CO₂ significantly increased photosynthesis in both male and female plants and by a similar magnitude. Males and females did not differ in net photosynthetic rate, but females had significantly greater biomass production than males, regardless of CO₂ concentrations. Vegetative mass increased by 39% in males and in females, whereas reproductive mass increased by 82% in males and 97% in females at elevated CO₂. As a result, proportionately more carbon was allocated to reproduction in male and female plants at elevated CO₂. Higher CO₂ increased individual seed mass significantly, but had no effect on the number or mass of seeds per female plant. Our results demonstrated that rising atmospheric CO₂ will alter the allocation patterns in both male and female S. latifolia Poiret plants by shifting proportionally more photosynthate to reproduction.     

Non-stomatal limitation of CO2 assimilation in three tree species during natural drought conditions

The effect of drought on CO₂ assimilation and leaf conductance was studied in three northern hardwood species: Quercus rubra L., Acer rubrum L. and Populus grandidentata Michx. Leaf gas exchange characteristics at two CO₂ levels (320 and 620 μl I⁻¹) and temperatures from 20 to 35°C were measured at the end of a dry period and shortly after 10 cm of rainfall. The effects of drought varied with species, temperature and CO₂ level. Calculated values of internal CO₂ concentration showed little or no decline during drought. Differences in assimilation, before vs after the rains, were most apparent at the higher CO₂ level. These latter two observations indicate nonstomatal disruption of CO₂ assimilation during the dry period. In P. grandidentata there was a substantial interaction between drought and temperature, with a resultant shift in the temperature for maximum assimilation to lower temperatures during drought. During drought, internal CO₂ concentrations increased sharply in all three species under the combined conditions of high temperatures and the higher CO₂ level.     

Leaf age effects of elevated CO2-grown white oak leaves on spring-feeding lepidopterans

Folivorous insect responses to elevated CO₂-grown tree species may be complicated by phytochemical changes as leaves age. For example, young expanding leaves in tree species may be less affected by enriched CO₂-alterations in leaf phytochemistry than older mature leaves due to shorter exposure times to elevated CO₂ atmospheres. This, in turn, could result in different effects on early vs. late instar larvae of herbivorous insects. To address this, seedlings of white oak (Quercus alba L.), grown in open-top chambers under ambient and elevated CO₂, were fed to two important early spring feeding herbivores; gypsy moth (Lymantria dispar L.), and forest tent caterpillar (Malacosoma disstria Hübner). Young, expanding leaves were presented to early instar larvae, and older fully expanded or mature leaves to late instar larvae. Young leaves had significantly lower leaf nitrogen content and significantly higher total nonstructural carbohydrate:nitrogen ratio as plant CO₂ concentration rose, while nonstructural carbohydrates and total carbon-based phenolics were unaffected by plant CO₂ treatment. These phytochemical changes contributed to a significant reduction in the growth rate of early instar gypsy moth larvae, while growth rates of forest tent caterpillar were unaffected. The differences in insect responses were attributed to an increase in the nitrogen utilization efficiency (NUE) of early instar forest tent caterpillar larvae feeding on elevated CO₂-grown leaves, while early instar gypsy moth larval NUE remained unchanged among the treatments. Later instar larvae of both insect species experienced larger reductions in foliage quality on elevated CO₂-grown leaves than earlier instars, as the carbohydrate:nitrogen ratio of leaves substantially increased. Despite this, neither insect species exhibited changes in growth or consumption rates between CO₂ treatments in the later instar. An increase in NUE was apparently responsible for offsetting reduced foliar nitrogen for the late instar larvae of both species.