Impacts of Hurricane Frances on Florida scrub-oak ecosystem processes: defoliation, net CO2 exchange and interactions with elevated CO2

Hurricane disturbances have profound impacts on ecosystem structure and function, yet their effects on ecosystem CO₂ exchange have not been reported. In September 2004, our research site on a fire‐regenerated scrub‐oak ecosystem in central Florida was struck by Hurricane Frances with sustained winds of 113 km h⁻¹ and wind gusts as high as 152 km h⁻¹. We quantified the hurricane damage on this ecosystem resulting from defoliation: we measured net ecosystem CO₂ exchange, the damage and recovery of leaf area, and determined whether growth in elevated carbon dioxide concentration in the atmosphere (C_a) altered this disturbance. The hurricane decreased leaf area index (LAI) by 21%, which was equal to 60% of seasonal variation in canopy growth during the previous 3 years, but stem damage was negligible. The reduction in LAI led to a 22% decline in gross primary production (GPP) and a 25% decline in ecosystem respiration (R_e). The compensatory declines in GPP and R_e resulted in no significant change in net ecosystem production (NEP). Refoliation began within a month after the hurricane, although this period was out of phase with the regular foliation period, and recovered 20% of the defoliation loss within 2.5 months. Full recovery of LAI, ecosystem CO₂ assimilation, and ecosystem respiration did not occur until the next growing season. Plants exposed to elevated C_a did not sustain greater damage, nor did they recover faster than plants grown under ambient C_a. Thus, our results indicate that hurricanes capable of causing significant defoliation with negligible damage to stems have negligible effects on NEP under current or future CO₂‐enriched environment.     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

Interactions between CO2 enhancement and N addition on net primary productivity and water‐use efficiency in a mesocosm with multiple subtropical tree species

Carbon dioxide (CO₂) enhancement (eCO₂) and N addition (aN) have been shown to increase net primary production (NPP) and to affect water‐use efficiency (WUE) for many temperate ecosystems, but few studies have been made on subtropical tree species. This study compared the responses of NPP and WUE from a mesocosm composing five subtropical tree species to eCO₂ (700 ppm), aN (10 g N m^?2 yr^?1) and eCO₂ × aN using open‐top chambers. Our results showed that mean annual ecosystem NPP did not changed significantly under eCO₂, increased by 56% under aN and 64% under eCO₂ × aN. Ecosystem WUE increased by 14%, 55%, and 61% under eCO₂, aN and eCO₂ × aN, respectively. We found that the observed responses of ecosystem WUE were largely driven by the responses of ecosystem NPP. Statistical analysis showed that there was no significant interactions between eCO₂ and aN on ecosystem NPP (P = 0.731) or WUE (P = 0.442). Our results showed that increasing N deposition was likely to have much stronger effects on ecosystem NPP and WUE than increasing CO₂ concentration for the subtropical forests. However, different tree species responded quite differently. aN significantly increased annual NPP of the fast‐growing species (Schima superba). Nitrogen‐fixing species (Ormosia pinnata) grew significantly faster only under eCO₂ × aN. eCO₂ had no effects on annual NPP of those two species but significantly increased annual NPP of other two species (Castanopsis hystrix and Acmena acuminatissima). Differential responses of the NPP among different tree species to eCO₂ and aN will likely have significant implications on the species composition of subtropical forests under future global change.     

Photosynthetic responses of 13 grassland species across 11 years of free‐air CO2 enrichment is modest,consistent and independent of N supply

If long‐term responses of photosynthesis and leaf diffusive conductance to rising atmospheric carbon dioxide (CO₂) levels are similar or predictably different among species, functional types, and ecosystem types, general global models of elevated CO₂ effects can effectively be developed. To address this issue we measured gas exchange rates of 13 perennial grassland species from four functional groups across 11 years of long‐term free‐air CO₂ enrichment (eCO₂, +180 ppm above ambient CO₂) in the BioCON experiment in Minnesota, USA. Eleven years of eCO₂ produced consistent but modest increases in leaf net photosynthetic rates of 10% on average compared with plants grown at ambient CO₂ concentrations across the 13 species. This eCO₂‐induced enhancement did not depend on soil N treatment, is much less than the average across other longer‐term studies, and represents strong acclimation (i.e. downregulation) as it is also much less than the instantaneous response to eCO₂. The legume and C3 nonlegume forb species were the most responsive among the functional groups (+13% in each), the C4 grasses the least responsive (+4%), and C3 grasses intermediate in their photosynthetic response to eCO₂ across years (+9%). Leaf stomatal conductance and nitrogen content declined comparably across species in eCO₂ compared with ambient CO₂ and to degrees corresponding to results from other studies. The significant acclimation of photosynthesis is explained in part by those eCO₂‐induced decreases in leaf N content and stomatal conductance that reduce leaf photosynthetic capacity in plants grown under elevated compared with ambient CO₂ concentrations. Results of this study, probably the longest‐term with the most species, suggest that carbon cycle models that assume and thereby simulate long‐lived strong eCO₂ stimulation of photosynthesis (e.g.> 25%) for all of Earth's terrestrial ecosystems should be viewed with a great deal of caution.     

Elevated CO2 did not affect the hydrological balance of a mature native Eucalyptus woodland

Elevated atmospheric CO₂ concentration (eC_a) might reduce forest water‐use, due to decreased transpiration, following partial stomatal closure, thus enhancing water‐use efficiency and productivity at low water availability. If evapotranspiration (E_t) is reduced, it may subsequently increase soil water storage (ΔS) or surface runoff (R) and drainage (D_g), although these could be offset or even reversed by changes in vegetation structure, mainly increased leaf area index (L). To understand the effect of eC_a in a water‐limited ecosystem, we tested whether 2 years of eC_a (~40% increase) affected the hydrological partitioning in a mature water‐limited Eucalyptus woodland exposed to Free‐Air CO₂ Enrichment (FACE). This timeframe allowed us to evaluate whether physiological effects of eC_a reduced stand water‐use irrespective of L, which was unaffected by eC_a in this timeframe. We hypothesized that eC_a would reduce tree‐canopy transpiration (E_tree), but excess water from reduced E_tree would be lost via increased soil evaporation and understory transpiration (E_floor) with no increase in ΔS, R or D_g. We computed E_t, ΔS, R and D_g from measurements of sapflow velocity, L, soil water content (θ), understory micrometeorology, throughfall and stemflow. We found that eC_a did not affect E_tree, E_floor, ΔS or θ at any depth (to 4.5 m) over the experimental period. We closed the water balance for dry seasons with no differences in the partitioning to R and D_g between C_a levels. Soil temperature and θ were the main drivers of E_floor while vapour pressure deficit‐controlled E_tree, though eC_a did not significantly affect any of these relationships. Our results suggest that in the short‐term, eC_a does not significantly affect ecosystem water‐use at this site. We conclude that water‐savings under eC_a mediated by either direct effects on plant transpiration or by indirect effects via changes in L or soil moisture availability are unlikely in water‐limited mature eucalypt woodlands.     

Direct and indirect effects of elevated atmospheric CO2 on net ecosystem production in a Chesapeake Bay tidal wetland

The rapid increase in atmospheric CO₂ concentrations (C_a) has resulted in extensive research efforts to understand its impact on terrestrial ecosystems, especially carbon balance. Despite these efforts, there are relatively few data comparing net ecosystem exchange of CO₂ between the atmosphere and the biosphere (NEE), under both ambient and elevated C_a. Here we report data on annual sums of CO₂ (NEE_net) for 19 years on a Chesapeake Bay tidal wetland for Scirpus olneyi (C₃ photosynthetic pathway)‐ and Spartina patens (C₄ photosynthetic pathway)‐dominated high marsh communities exposed to ambient and elevated C_a (ambient + 340 ppm). Our objectives were to (i) quantify effects of elevated C_a on seasonally integrated CO₂ assimilation (NEE_net = NEE_day + NEE_night, kg C m^?2 y^?1) for the two communities; and (ii) quantify effects of altered canopy N content on ecosystem photosynthesis and respiration. Across all years, NEE_net averaged 1.9 kg m^?2 y^?1 in ambient C_a and 2.5 kg m^?2 y^?1 in elevated C_a, for the C₃‐dominated community. Similarly, elevated C_a significantly (P < 0.01) increased carbon uptake in the C₄‐dominated community, as NEE_net averaged 1.5 kg m^?2 y^?1 in ambient C_a and 1.7 kg m^?2 y^?1 in elevated C_a. This resulted in an average CO₂ stimulation of 32% and 13% of seasonally integrated NEE_net for the C₃‐ and C₄‐dominated communities, respectively. Increased NEE_day was correlated with increased efficiencies of light and nitrogen use for net carbon assimilation under elevated C_a, while decreased NEE_night was associated with lower canopy nitrogen content. These results suggest that rising C_a may increase carbon assimilation in both C₃‐ and C₄‐dominated wetland communities. The challenge remains to identify the fate of the assimilated carbon.     

Atmospheric CO2 concentration does not directly affect leaf respiration in bean or poplar

It is a matter of debate if there is a direct (short‐term) effect of elevated atmospheric CO₂ concentration (C_a) on plant respiration in the dark. When C_a doubles, some authors found no (or only minor) changes in dark respiration, whereas most studies suggest a respiratory inhibition of 15–20%. The present study shows that the measurement artefacts – particularly leaks between leaf chamber gaskets and leaf surface, CO₂ memory and leakage effects of gas exchange systems as well as the water vapour (‘water dilution’) effect on DCO₂ measurement caused by transpiration – may result in larger errors than generally discussed. A gas exchange system that was used in three different ways – as a closed system in which C_a increased continuously from 200 to 4200 mmol (CO₂) mol^‐1 (air) due to respiration of the enclosed leaf; as an intermittently closed system that was repeatedly closed and opened during C_a periods of either 350 or 2000 mmol mol^‐1, and as an open system in which C_a varied between 350 and 2000 mmol mol^‐1– is described. In control experiments (with an empty leaf chamber), the respective system characteristics were evaluated carefully. When all relevant system parameters were taken into account, no effects of short‐term changes in CO₂ on dark CO₂ efflux of bean and poplar leaves were found, even when C_a increased to 4200 mmol mol^‐1. It is concluded that the leaf respiration of bean and poplar is not directly inhibited by elevated atmospheric CO₂.     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

Nutrient concentrations in a Littorella uniflora community at higher CO2 concentrations and reduced light intensities

1. Oligotrophic softwater lakes represent a special type of aquatic ecosystem with unique plant communities where generalisations from other aquatic plant communities to rising CO₂ in the water column may not apply. 2. In the present study, we set up large in situ mesocosms and supporting laboratory experiments with isoetid vegetation (Littorella uniflora) where water column CO₂ and light could be manipulated in order to test whether (i) light and CO₂ availability affect nutrient concentrations in isoetid vegetation, and (ii) if changes in light and CO₂ climate affect fluxes of inorganic nitrogen (N) and phosphorus (P) from sediment to water column, which potentially could result in increased growth of epiphytic algae. 3. The results showed that the standing stocks of phosphorus and nitrogen in the L. uniflora vegetation were significantly influenced by CO₂ concentration and light intensity. Both standing stocks of P and N were significantly higher in the mesocosm treatments with high CO₂ concentration than in those at low CO₂ concentration. Similarly, standing stocks of P and N enhanced with increasing light intensity. 4. Measurements of nutrient fluxes both in the field and the laboratory did not show any significant release of nutrients to the water column from plants or sediments at any of the light or CO₂ treatments. However, mats of epiphytic algae developed from the beginning of June to late September and caused a light reduction for the isoetid vegetation. 5. Increasing CO₂ concentrations in the water column may over time potentially result in a change in soft water plant communities.     

SHORT‐ AND LONG‐TERM EFFECTS OF ELEVATED CO2 ON PHOTOSYNTHESIS AND RESPIRATION IN THE MARINE MACROALGA HIZIKIA FUSIFORMIS (SARGASSACEAE,PHAEOPHYTA) GROWN AT LOW AND HIGH N SUPPLIES1

The short‐term and long‐term effects of elevated CO₂ on photosynthesis and respiration were examined in cultures of the marine brown macroalga Hizikia fusiformis (Harv.) Okamura grown under ambient (375 μL · L^?1) and elevated (700 μL · L^?1) CO₂ concentrations and at low and high N availability. Short‐term exposure to CO₂ enrichment stimulated photosynthesis, and this stimulation was maintained with prolonged growth at elevated CO₂, regardless of the N levels in culture, indicating no down‐regulation of photosynthesis with prolonged growth at elevated CO₂. However, the photosynthetic rate of low‐N‐grown H. fusiformis was more responsive to CO₂ enrichment than that of high‐N‐grown algae. Elevation of CO₂ concentration increased the value of K_1/2(Ci) (the half‐saturation constant) for photosynthesis, whereas high N supply lowered it. Neither short‐term nor long‐term CO₂ enrichment had inhibitory effects on respiration rate, irrespective of the N supply, under which the algae were grown. Under high‐N growth, the Q₁₀ value of respiration was higher in the elevated‐CO₂‐grown algae than the ambient‐CO₂‐grown algae. Either short‐ or long‐term exposure to CO₂ enrichment decreased respiration as a proportion of gross photosynthesis (Pg) in low‐N‐grown H. fusiformis. It was proposed that in a future world of higher atmospheric CO₂ concentration and simultaneous coastal eutrophication, the respiratory carbon flux would be more sensitive to changing temperature.     

Photosynthesis, carboxylation and leaf nitrogen responses of 16 species to elevated pCO2 across four free-air CO2 enrichment experiments in forest, grassland and desert

The magnitude of changes in carboxylation capacity in dominant plant species under long‐term elevated CO₂ exposure (elevated pC_a) directly impacts ecosystem CO₂ assimilation from the atmosphere. We analyzed field CO₂ response curves of 16 C₃ species of different plant growth forms in favorable growth conditions in four free‐air CO₂ enrichment (FACE) experiments in a pine and deciduous forest, a grassland and a desert. Among species and across herb, tree and shrub growth forms there were significant enhancements in CO₂ assimilation (A) by +40±5% in elevated pC_a (49.5–57.1 Pa), although there were also significant reductions in photosynthetic capacity in elevated pC_a in some species. Photosynthesis at a common pC_a (A_a) was significantly reduced in five species growing under elevated pC_a, while leaf carboxylation capacity (V_cmax) was significantly reduced by elevated pC_a in seven species (change of ?19±3% among these species) across different growth forms and FACE sites. Adjustments in V_cmax with elevated pC_a were associated with changes in leaf N among species, and occurred in species with the highest leaf N. Elevated pC_a treatment did not affect the mass‐based relationships between A or V_cmax and N, which differed among herbs, trees and shrubs. Thus, effects of elevated pC_a on leaf C assimilation and carboxylation capacity occurred largely through changes in leaf N, rather than through elevated pC_a effects on the relationships themselves. Maintenance of leaf carboxylation capacity among species in elevated pC_a at these sites depends on maintenance of canopy N stocks, with leaf N depletion associated with photosynthetic capacity adjustments. Since CO₂ responses can only be measured experimentally on a small number of species, understanding elevated CO₂ effects on canopy N_m and N_a will greatly contribute to an ability to model responses of leaf photosynthesis to atmospheric CO₂ in different species and plant growth forms.     

No photosynthetic down-regulation in sweetgum trees (Liquidambar styraciflua L.) after three years of CO2 enrichment at the Duke Forest FACE experiment

Photosynthetic capacity and leaf properties of sun and shade leaves of overstorey sweetgum trees (Liquidambar styraciflua L.) were compared over the first 3 years of growth in ambient or ambient + 200 μL L^?1 CO₂ at the Duke Forest Free Air CO₂ Enrichment (FACE) experiment. We were interested in whether photosynthetic down‐regulation to CO₂ occurred in sweetgum trees growing in a forest ecosystem, whether shade leaves down‐regulated to a greater extent than sun leaves, and if there was a seasonal component to photosynthetic down‐regulation. During June and September of each year, we measured net photosynthesis (A) versus the calculated intercellular CO₂ concentration (C_i) in situ and analysed these response curves using a biochemical model that described the limitations imposed by the amount and activity of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Vc_max) and by the rate of ribulose‐1,5‐bisphosphate (RuBP) regeneration mediated by electron transport (J_max). There was no evidence of photosynthetic down‐regulation to CO₂ in either sun or shade leaves of sweetgum trees over the 3 years of measurements. Elevated CO₂ did not significantly affect Vc_max or J_max. The ratio of Vc_max to J_max was relatively constant, averaging 2·12, and was not affected by CO₂ treatment, position in the canopy, or measurement period. Furthermore, CO₂ enrichment did not affect leaf nitrogen per unit leaf area (N_a), chlorophyll or total non‐structural carbohydrates of sun or shade leaves. We did, however, find a strong relationship between N_a and the modelled components of photosynthetic capacity, Vc_max and J_max. Our data over the first 3 years of this experiment corroborate observations that trees rooted in the ground may not exhibit symptoms of photosynthetic down‐regulation as quickly as tree seedlings growing in pots. There was a strong sustained enhancement of photosynthesis by CO₂ enrichment whereby light‐saturated net photosynthesis of sun leaves was stimulated by 63% and light‐saturated net photosynthesis of shade leaves was stimulated by 48% when averaged over the 3 years. This study suggests that this CO₂ enhancement of photosynthesis will be sustained in the Duke Forest FACE experiment as long as soil N availability keeps pace with photosynthetic and growth processes.     

18O composition of CO2 and H2O ecosystem pools and fluxes in a tallgrass prairie: Simulations and comparisons to measurements

In this paper we describe measurements and modeling of ¹⁸O in CO₂ and H₂O pools and fluxes at a tallgrass prairie site in Oklahoma. We present measurements of the δ¹⁸O value of leaf water, depth‐resolved soil water, atmospheric water vapor, and Keeling plot δ¹⁸O intercepts for net soil‐surface CO₂ and ecosystem CO₂ and H₂O fluxes during three periods of the 2000 growing season. Daytime discrimination against C¹⁸OO, as calculated from measured above‐canopy CO₂ and δ¹⁸O gradients, is also presented. To interpret the isotope measurements, we applied an integrated land‐surface and isotope model (ISOLSM) that simulates ecosystem H₂¹⁸O and C¹⁸OO stocks and fluxes. ISOLSM accurately predicted the measured isotopic composition of ecosystem water pools and the δ¹⁸O value of net ecosystem CO₂ and H₂O fluxes. Simulations indicate that incomplete equilibration between CO₂ and H₂O within C₄ plant leaves can have a substantial impact on ecosystem discrimination. Diurnal variations in the δ¹⁸O value of above‐canopy vapor had a small impact on the predicted δ¹⁸O value of ecosystem water pools, although sustained differences had a large impact. Diurnal variations in the δ¹⁸O value of above‐canopy CO₂ substantially affected the predicted ecosystem discrimination. Leaves dominate the ecosystem ¹⁸O‐isoflux in CO₂ during the growing season, while the soil contribution is relatively small and less variable. However, interpreting daytime measurements of ecosystem C¹⁸OO fluxes requires accurate predictions of both soil and leaf ¹⁸O‐isofluxes.     

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.     

Reduction of forest floor respiration by fertilization on both carbon dioxide-enriched and reference 17-year-old loblolly pine stands

Elevated atmospheric carbon dioxide (CO₂^e) increases soil respiration rates in forest, grassland, agricultural and wetland systems as a result of increased growth, root biomass and enhanced biological activity of soil microorganisms. Less is known about how forest floor fluxes respond to the combined effects of elevated CO₂ and nutrient amendments; until now no experiments have been in place with large forest trees to allow even preliminary investigations. We investigated changes in forest floor respiration (S_ff) in a Pinus taeda L. plantation fumigated with CO₂ by using free‐air CO₂ enrichment (FACE) technology and given nutrient amendments. The prototype FACE apparatus (FACEp; 707 m²) was constructed in 1993, 10 years after planting, on a moderate fertility site in Duke Forest, North Carolina, USA, enriching the stand to 55 Pa (CO₂^e). A nearby ambient CO₂ (CO₂^a) plot (117 m²) was designated at the inception of the study as a reference (Ref). Both FACEp and Ref plot were divided in half and urea fertilizer was applied to one half at an annual rate of 11.2 g N m^?2 in the spring of 1998, 1999 and 2000. Forest floor respiration was monitored continuously for 220 days – March through November 2000 – by using two Automated Carbon Efflux Systems. Thirty locations (491 cm² each) were sampled in both FACEp and Ref, about half in each fertility treatment. Forest floor respiration was strongly correlated with soil temperature at 5 cm. Rates of S_ff were greater in CO₂^e relative to CO₂^a (an enhancement of ～178 g C m^?2) during the measurement period. Application of fertilizer resulted in a statistically significant depression of respiration rates in both the CO₂^a and CO₂^e plots (a reduction of ～186 g C m^?2). The results suggest that closed canopy forests on moderate fertility sites cycle back to the atmosphere more assimilated carbon (C) than similar forests on sites of high fertility. We recognize the limitations of this non‐replicated study, but its clear results offer strong testable hypotheses for future research in this important area.     

Hydrologic balance in an intact temperate forest ecosystem under ambient and elevated atmospheric CO2 concentration

Abstract Increasing atmospheric CO₂ concentration decreases stomatal conductance in many species, but the savings of water from reduced transpiration may permit the forest to retain greater leaf area index (L). Therefore, the net effect on water use in forest ecosystems under a higher CO₂ atmosphere is difficult to predict. The free air CO₂ enrichment (FACE) facility (n = 3) in a 14‐m tall (in 1996) Pinus taeda L. stand was designed to reduce uncertainties in predicting such responses. Continuous measurements of precipitation, throughfall precipitation, sap flux, and soil moisture were made over 3.5 years under ambient (CO₂^a) and elevated (CO₂^e) ambient + 200 µmol mol^?1). Annual stand transpiration under ambient CO₂ conditions accounted for 84–96% of latent heat flux measured with the eddy‐covariance technique above the canopy. Under CO₂^e, P. taeda transpired less per unit of leaf area only when soil drought was severe. Liquidambar styraciflua, the other major species in the forest, used progressively less water, settling at 25% reduction in sap flux density after 3.5 years under CO₂^e. Because P. taeda dominated the stand, and severe drought periods were of relatively short duration, the direct impact of CO₂^e on water savings in the stand was undetectable. Moreover, the forest used progressively more water under CO₂^e, probably because soil moisture availability progressively increased, probably owing to a reduction in soil evaporation caused by more litter buildup in the CO₂^e plots. The results suggest that, in this forest, the effect of CO₂^e on transpiration was greater indirectly through enhanced litter production than directly through reduced stomatal conductance. In forests composed of species more similar to L. styraciflua, water savings from stomatal closure may dominate the response to CO₂^e.     

Modelling assimilation and intercellular CO2 from measured conductance: a synthesis of approaches

A spectrum of models that estimate assimilation rate A from intercellular carbon dioxide concentration (C_i) and measured stomatal conductance to CO₂ (g_c) were investigated using leaf‐level gas exchange measurements. The gas exchange measurements were performed in a uniform loblolly pine stand (Pinus taeda L.) using the Free Air CO₂ Enrichment (FACE) facility under ambient and elevated atmospheric CO₂ for 3 years. These measurements were also used to test a newly proposed framework that combines basic properties of the A–C_i curve with a Fickian diffusion transport model to predict the relationship between C_i/C_a and g_c, where C_a is atmospheric carbon dioxide concentration. The widely used Ball–Berry model and five other models as well as the biochemical model proposed by Farquhar et al. (1980) were also reformulated to express variations in C_i/C_a as a function of their corresponding driving mechanisms. To assess the predictive capabilities of these approaches, their respective parameters were estimated from independent measurements of long‐term stable carbon isotope determinations (δ¹³C), meteorological variables, and ensemble A–C_i curves. All eight approaches reproduced the measured A reasonably well, in an ensemble sense, from measured water vapour conductance and modeled C_i/C_a. However, the scatter in the instantaneous A estimates was sufficiently large for both ambient and elevated C_a to suggest that other transient processes were not explicitly resolved by all eight parameterizations. An important finding from our analysis is that added physiological complexity in modeling C_i/C_a (when g_c is known) need not always translate to increased accuracy in predicting A. Finally, the broader utility of these approaches to estimate assimilation and net ecosystem exchange is discussed in relation to elevated atmospheric CO₂.     

Effect of eutrophication on air–sea CO2 fluxes in the coastal Southern North Sea: a model study of the past 50 years

The RIVERSTRAHLER model, an idealized biogeochemical model of the river system, has been coupled to MIRO‐CO₂, a complex biogeochemical model describing diatom and Phaeocystis blooms and carbon and nutrient cycles in the marine domain, to assess the dual role of changing nutrient loads and increasing atmospheric CO₂ as drivers of air–sea CO₂ exchanges in the Southern North Sea with a focus on the Belgian coastal zone (BCZ). The whole area, submitted to the influence of two main rivers (Seine and Scheldt), is characterized by variable diatom and Phaeocystis colonies blooms which impact on the trophic status and air–sea CO₂ fluxes of the coastal ecosystem. For this application, the MIRO‐CO₂ model is implemented in a 0D multibox frame covering the eutrophied Eastern English Channel and Southern North Sea and receiving loads from the rivers Seine and Scheldt. Model simulations are performed for the period between 1951 and 1998 using real forcing fields for sea surface temperature, wind speed and atmospheric CO₂ and RIVERSTRAHLER simulations for river carbon and nutrient loads. Model results suggest that the BCZ shifted from a source of CO₂ before 1970 (low eutrophication) towards a sink during the 1970–1990 period when anthropogenic DIN and P loads increased, stimulating C fixation by autotrophs. In agreement, a shift from net annual heterotrophy towards autotrophy in BCZ is simulated from 1980. The period after 1990 is characterized by a progressive decrease of P loads concomitant with a decrease of primary production and of the CO₂ sink in the BCZ. At the end of the simulation period, the BCZ ecosystem is again net heterotroph and acts as a source of CO₂ to the atmosphere. R‐MIRO‐CO₂ scenarios testing the relative impact of temperature, wind speed, atmospheric CO₂ and river loads variability on the simulated air–sea CO₂ fluxes suggest that the trend in air–sea CO₂ fluxes simulated between 1951 and 1998 in the BCZ was mainly controlled by the magnitude and the ratio of inorganic nutrient river loads. Quantitative nutrient changes control the level of primary production while qualitative changes modulate the relative contribution of diatoms and Phaeocystis to this flux and hence the sequestration of atmospheric CO₂.     

Integration of CO2 flux and remotely-sensed data for primary production and ecosystem respiration analyses in the Northern Great Plains: potential for quantitative spatial extrapolation

Aim Extrapolation of tower CO₂ fluxes will be greatly facilitated if robust relationships between flux components and remotely sensed factors are established. Long‐term measurements at five Northern Great Plains locations were used to obtain relationships between CO₂ fluxes and photosynthetically active radiation (Q), other on‐site factors, and Normalized Difference Vegetation Index (NDVI) from the SPOT VEGETATION data set. Location CO₂ flux data from the following stations and years were analysed: Lethbridge, Alberta 1998–2001; Fort Peck, MT 2000, 2002; Miles City, MT 2000–01; Mandan, ND 1999–2001; and Cheyenne, WY 1997–98. Results Analyses based on light‐response functions allowed partitioning net CO₂ flux (F) into gross primary productivity (P_g) and ecosystem respiration (R_e). Weekly averages of daytime respiration, γ_day, estimated from light responses were closely correlated with weekly averages of measured night‐time respiration, γ_night (R² 0.64 to 0.95). Daytime respiration tended to be higher than night‐time respiration, and regressions of γ_day on γ_night for all sites were different from 1 : 1 relationships. Over 13 site‐years, gross primary production varied from 459 to 2491 g CO₂ m^?2 year^?1, ecosystem respiration from 996 to 1881 g CO₂ m^?2 year^?1, and net ecosystem exchange from ?537 (source) to +610 g CO₂ m^?2 year^?1 (sink). Maximum daily ecological light‐use efficiencies, ?_d,max = P_g/Q, were in the range 0.014 to 0.032 mol CO₂ (mol incident quanta)^?1. Main conclusions Ten‐day average P_g was significantly more highly correlated with NDVI than 10‐day average daytime flux, P_d (R² = 0.46 to 0.77 for P_g‐NDVI and 0.05 to 0.58 for P_d‐NDVI relationships). Ten‐day average R_e was also positively correlated with NDVI, with R² values from 0.57 to 0.77. Patterns of the relationships of P_g and R_e with NDVI and other factors indicate possibilities for establishing multivariate functions allowing scaling‐up local fluxes to larger areas using GIS data, temporal NDVI, and other factors.     

Altered sediment biota and lagoon habitat carbonate dynamics due to sea cucumber bioturbation in a high‐pCO2 environment

The effects of global change on biological systems and functioning are already measurable, but how ecological interactions are being altered is poorly understood. Ecosystem resilience is strengthened by ecological functionality, which depends on trophic interactions between key species and resilience generated through biogenic buffering. Climate‐driven alterations to coral reef metabolism, structural complexity and biodiversity are well documented, but the feedbacks between ocean change and trophic interactions of non‐coral invertebrates are understudied. Sea cucumbers, some of the largest benthic inhabitants of tropical lagoon systems, can influence diel changes in reef carbonate dynamics. Whether they have the potential to exacerbate or buffer ocean acidification over diel cycles depends on their relative production of total alkalinity (A_T) through the dissolution of ingested calcium carbonate (CaCO₃) sediments and release of dissolved inorganic carbon (C_T) through respiration and trophic interactions. In this study, the potential for the sea cucumber, Stichopus herrmanni, a bêche‐de‐mer (fished) species listed as vulnerable to extinction, to buffer the impacts of ocean acidification on reef carbonate chemistry was investigated in lagoon sediment mesocosms across diel cycles. Stichopus herrmanni directly reduced the abundance of meiofauna and benthic primary producers through its deposit‐feeding activity under present‐day and near‐future pCO₂. These changes in benthic community structure, as well as A_T (sediment dissolution) and C_T (respiration) production by S. herrmanni, played a significant role in modifying seawater carbonate dynamics night and day. This previously unappreciated role of tropical sea cucumbers, in support of ecosystem resilience in the face of global change, is an important consideration with respect to the bêche‐de‐mer trade to ensure sea cucumber populations are sustained in a future ocean.