δ13C-variations of leaves in forests as an indication of reassimilated CO2 from the soil

Summary An attempt has been made to evaluate the contribution of soil respired CO₂ to the total assimilation of a forest tree, by heeding the ¹³C-concentrations of CO₂ from the free atmosphere and from mineralization processes within the soil respectively. An expression has been derived, according to which the assimilated fraction of CO₂ from the soil at a particular height of a tree is given by the ¹³C-value of the corresponding leaves, ¹³C of atmospheric CO₂, ¹³C of soil respired CO₂ and the physiological state of the leaves expressed as the ratio of total respiration over gross photosynthesis and internal over external CO₂-concentration. In the particular case investigated, a ¹³C-difference of 5 has been determined from bottom to top of a beech tree which results in a CO₂ contribution from the soil of about 22% for the lower forest strata, while the total contribution of soil respired CO₂ accounts for about 5% of the overall assimilation.     

Stand microclimate and physiological activity of tree leaves in an oak-hornbeam forest

Summary In an uneven-aged, multi-species oak-hornbeam forest at Báb, SW Slovakia (former IBP Forest Research Site), a series of micrometeorological and ecophysiological measurements started in 1985. The aims of the work are to improve understanding of physiological processes (photosynthesis, respiration, and transpiration) of adult trees and stand microclimate, to collect data for simulation of the canopy (stand) photosynthesis and for ecological synthesis of the functioning of the forest ecosystem. In this paper, photosynthetically active radiation (PAR), air temperature (AT) and relative humidity (RH), wind speed (WS), and CO₂ concentration ([CO₂]) in and above the forest are characterized for the fully leaved season, using diurnal courses, vertical profiles and isodiagrams (isopleths). Approximately 50% of incident PAR was absorbed by the upper 4–5 m layer of leaves and only approximately 5% or less penetrated to the forest floor. Vertical gradients of AT and RH were generally low, but large differences in diurnal ranges of AT and RH were observed between vertical levels. The upper leaf canopy greatly reduced WS, and at a height of about 14 m above the ground it was close to zero. The highest diurnal [CO₂] maximum and variations occurred at 1 m above the ground, and the lowest above the forest. In good light conditions in the forest, the entire leaf canopy (overstorey and understorey canopy) is a large sink of CO₂. At night the forest stand is a source of CO₂, the largest internal source being the soil and forest floor.     

Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients

 Our objective was to evaluate the relative importance of gradients in light intensity and the isotopic composition of atmospheric CO₂ for variation in leaf carbon isotope ratios within a Pinus resinosa forest. In addition, we measured photosynthetic gas exchange and leaf carbon isotope ratios on four understory species (Dryopteris carthusiana, Epipactus helleborine, Hieracium floribundum, Rhamnus frangula), in order to estimate the consequence of the variation in the understory light microclimate for carbon gain in these plants. During midday, CO₂ concentration was relatively constant at vertical positions ranging from 15 m to 3 m above ground. Only at positions below 3 m was CO₂ concentration significantly elevated above that measured at 15 m. Based on the strong linear relationship between changes in CO₂ concentration and δ¹³C values for air samples collected during a diurnal cycle, we calculated the expected vertical profile for the carbon isotope ratio of atmospheric CO₂ within the forest. These calculations indicated that leaves at 3 m height and above were exposed to CO₂ of approximately the same isotopic composition during daylight periods. There was no significant difference between the daily mean δ¹³C values at 15 m (–7.77‰) and 3 m (–7.89‰), but atmospheric CO₂ was significantly depleted in ¹³C closer to the ground surface, with daily average δ¹³C values of –8.85‰ at 5 cm above ground. The light intensity gradient in the forest was substantial, with average photosynthetically active radiation (PAR) on the forest floor approximately 6% of that received at the top of the canopy. In contrast, there were only minor changes in air temperature, and so it is likely that the leaf-air vapour pressure difference was relatively constant from the top of the canopy to the forest floor. For red pine and elm tree samples, there was a significant correlation between leaf δ¹³C value and the height at which the leaf sample was collected. Leaf tissue sampled near the forest floor, on average, had lower δ¹³C values than samples collected near the top of the canopy. We suggest that the average light intensity gradient through the canopy was the major factor influencing vertical changes in tree leaf δ¹³C values. In addition, there was a wide range of variation (greater than 4‰) among the four understory plant species for average leaf δ¹³C values. Measurements of leaf gas exchange, under natural light conditions and with supplemental light, were used to estimate the influence of the light microclimate on the observed variation in leaf carbon isotope ratios in the understory plants. Our data suggest that one species, Epipactus helleborine, gained a substantial fraction of carbon during sunflecks. Received: 21 March 1996 / Accepted: 13 August 1996     

The relationship between carbon dioxide fixation and chlorophyll a fluorescence during induction of photosynthesis in maize leaves at different temperatures and carbon dioxide concentrations

The rate of CO₂ fixation (F_c) and 680 nm chlorophyll fluorescence emission (F680) were measured simultaneously during induction of photosynthesis in Zea mays L. leaves under varying experimental conditions in order to assess the validity of fluorescence as an indicator of in vivo photosynthetic carbon assimilation. Z. mays leaves showed typical Kautsky fluorescence induction curves consisting of a fast rise in emission (O to P) followed by a slow quenching via a major transient (S-M) to a steady-state (T). After an initial lag, net CO₂ assimilation commenced at a point corresponding to the onset of the S-M transient on the F680 induction curve. Subsequently, F_c and F680 always arrived at a steady-state simultaneously. Decreasing the dark-adaption period increased the rate of induction of both parameters. Alteration of leaf temperature produced anti-parallel changes in induction characteristics of F_c and F680. Reducing the CO₂ level to below that required for saturation of photosynthesis also produced anti-parallel changes during induction, however, at CO₂ concentrations tenfold greater than the atmospheric level the rate of F680 quenching from P to T was appreciably reduced without a similar change in the induction of F_c. Removal of CO₂ at steady-state produced only a small increase in F680 and a correspondingly small decrease in F680 occurred when CO₂ was re-introduced. The complex relationship between chlorophyll fluorescence and carbon assimilation in vivo is discussed and the applicability of fluorescence as an indicator of carbon assimilation is considered.Abbreviations F_c rate of CO₂ fixation - F680 fluorescence emission at 680 nm     

Carbon fluxes to the soil in a mature temperate forest assessed by <Superscript>13</Superscript>C isotope tracing

Photosynthetic carbon uptake and respiratory C release from soil are major components of the global carbon balance. The use of ¹³C depleted CO₂ (¹³C = –30) in a free air CO₂ enrichment experiment in a mature deciduous forest permitted us to trace the carbon transfer from tree crowns to the rhizosphere of 100–120 years old trees. During the first season of CO₂ enrichment the CO₂ released from soil originated substantially from concurrent assimilation. The small contribution of recent carbon in fine roots suggests a much slower fine root turnover than is often assumed.¹³C abundance in soil air correlated best with temperature data taken from 4 to 10 days before air sampling time and is thus rapidly available for root and rhizosphere respiration. The spatial variability of ¹³C in soil air showed relationships to above ground tree types such as conifers versus broad-leaved trees. Considering the complexity and strong overlap of roots from different individuals in a forest, this finding opens an exciting new possibility of associating respiration with different species. What might be seen as signal noise does in fact contain valuable information on the spatial heterogeneity of tree-soil interaction.     

The carbon isotope ratio of plant organic material reflects temporal and spatial variations in CO2 within tropical forest formations in Trinidad

Summary A method of monitoring and collecting CO₂ samples in the field has been developed which has been used to study both temporal and spatial variations in canopy CO₂ isotopic signatures in two contrasting tropical forest formations in Trinidad. These have been related to vertical gradients in the carbon isotope ratio (¹³C) of organic material in conjunction with measurements of other environmental parameters. The ¹³C of leaf material from two canopies showed a gradient with respect to height, more negative values being found low in the understorey. The deciduous secondary forest, (Simla) showed a difference of 4.6 and the semi-evergreen seasonal canopy (Aripo), 2.8. The range of ¹³C values at Simla was 4 less negative than those at Aripo. In order to relate these measurements to the interaction between diffusion or carboxylation limitation, and source CO₂ effects, variations in environmental parameters through the canopy have been compared with changes in CO₂ partial pressure (P _a) and isotopic composition ¹³C throughout the day during the dry season. Values of P _a20 m above the ground at Aripo varied from 380 vpm at dawn to 340 vpm at midday, at which time the partial pressure 15 cm above the ground was 375 vpm. The CO₂ partial pressure did not stabilise during the course of the day, and there was good correlation (r ²=0.82) between _a and P _a, with more negative values of _a occuring in the understorey. Diuraal changes of 2 were evident at all canopy positions. In the more open canopy at Simla, these gradients were similar, but less marked. Leaf-air vapour pressure deficit (VPD) showed no relationship with height, possibly as a result of minimal water flux from both the soil and the canopy due to low soil water content; VPD was 1.5 kPa higher at midday than dawn. A 3° C temperature gradient between the understorey and upper canopy was observed at Aripo but not in the more open Simla canopy. CO₂ partial pressure stabilised for only 4 h in the middle of the day, while other parameters showed no stable period. The proportion of floor respired CO₂ reassimilated at Aripo has been calculated as 26%, 19%, and 8% for the periods 0600–1000, 1000–1400, and 1400–1800 hours. In order to quantify source CO₂ effects, measurements of the environmental parameters and assimilation rate must be made at all canopy positions and throughout the day.     

Carbon-dioxide exchange in lichens: determination of transport and carboxylation characteristics

Measurements were made of net rates of CO₂ assimilation in lichens at various ambient concentrations of CO₂ in air and in helox (79% He, 21% O₂). Because of the faster rate of CO₂ diffusion in the pores of lichen thalli when filled with helox than when filled with air, a given net rate of assimilation was achieved at a lower ambient concentration of CO₂ in helox. The differences were used to estimate resistances to diffusion through the gas-filled pore systems in lichens. The technique was first tested with five lichen species, and then applied in a detailed study with Ramalina maciformis, in which gas-phase resistances were determined in samples at four different states of hydration and with two irradiances. By assuming, on the basis of previous evidence, that the phycobiont in R. maciformis is fully turgid and photosynthetically competent at the smallest hydration imposed (equilibration with vapour at 97% relative humidity), and that, with this state of hydration, diffusion of CO₂ to the phycobiont takes place through continuously gas-filled pores, it was possible also to determine both the dependence of net rate of assimilation in the phycobiont on local concentration of CO₂ in the algal layer, and, with the wetter samples, the extents to which diffusion of CO₂ to the phycobiont was impeded by water films. In equilibrium with air of 97% relative humidity, the thallus water content being 0.5 g per g dry weight, the resistance to CO₂ diffusion through the thallus was about twice as large as the resistance to CO₂ uptake within the phycobiont. Total resistance to diffusion increased rapidly with increase in hydration. At a water content of 2 g per g it was about 50 times as great as the resistance to uptake within the phycobiont and more than two-thirds of it was attributable to impedance of transfer by water. The influences of water content on rate of assimilation at various irradiances are discussed. The analysis shows that the local CO₂ compensation concentration of the phycobiont in R. maciformis is close to zero, indicating that photorespiratory release of CO₂ does not take place in the alga, Trebouxia sp., under the conditions of these experiments.Symbols and Units rate of CO₂ diffusion in air relative to that in carrier gas (unity if the carrier gas is air and 0.43 if is helox) - A₁ net rate of CO₂ uptake by the lichen - A_p gross rate of carboxylation minus photorespiratory decarboxylation in the phycobiont, i.e. net rate of light-activated CO₂ exchange - A_* maximum, CO₂-saturated magnitude of A_p - c concentration of CO₂ - c_a ambient concentration of CO₂ - c_i c_a minus difference in CO₂ concentration across air-filled pore space in the thallus - c₈ CO₂ concentration equivalent to partial pressure of CO₂ at the surface of the phycobiont - ₁ magnitude of c_a at which A₁ = 0 - _* magnitude of c_* at which A_p = 0 - R rate of dark respiration in the lichen (mycobiont and phycobiont) - R rate of dark respiration in region between the surface of the lichen and an arbitrary distance from the surface within the thallus - r resistance to CO₂ transfer from lichen surface to the surface of the phycobiont - r resistance to CO₂ transfer between effective source of dark respiration in the lichen and the surface of the phycobiont - r_g, r _g components of r and r, respectively, attributable to transfer in air-phase - r_w, r _w components of r and r, respectively, attributable to transfer in water-phase - r component of r between surface of lichen and an arbitrary distance from the surface within the thallus - r_* resistance to CO₂ transfer and carboxylation in the phycobiont - RH relative humidity     

Wood CO2 efflux in a primary tropical rain forest

The balance between photosynthesis and plant respiration in tropical forests may substantially affect the global carbon cycle. Woody tissue CO₂ efflux is a major component of total plant respiration, but estimates of ecosystem‐scale rates are uncertain because of poor sampling in the upper canopy and across landscapes. To overcome these problems, we used a portable scaffolding tower to measure woody tissue CO₂ efflux from ground level to the canopy top across a range of sites of varying slope and soil phosphorus content in a primary tropical rain forest in Costa Rica. The objectives of this study were to: (1) determine whether to use surface area, volume, or biomass for modeling and extrapolating wood CO₂ efflux, (2) determine if wood CO₂ efflux varied seasonally, (3) identify if wood CO₂ efflux varied by functional group, height in canopy, soil fertility, or slope, and (4) extrapolate wood CO₂ efflux to the forest. CO₂ efflux from small diameter woody tissue (<10 cm) was related to surface area, while CO₂ efflux from stems >10 cm was related to both surface area and volume. Wood CO₂ efflux showed no evidence of seasonality over 2 years. CO₂ efflux per unit wood surface area at 25° (F_A) was highest for the N‐fixing dominant tree species Pentaclethra macroloba, followed by other tree species, lianas, then palms. Small diameter F_A increased steeply with increasing height, and large diameter F_A increased with diameter. Soil phosphorus and slope had slight, but complex effects on F_A. Wood CO₂ efflux per unit ground area was 1.34±0.36 μmol m^?2 s^?1, or 508±135 g C m^?2 yr^?1. Small diameter wood, only 15% of total woody biomass, accounted for 70% of total woody tissue CO₂ efflux from the forest; while lianas, only 3% of total woody biomass, contributed one‐fourth of the total wood CO₂ efflux.     

Does tree species composition affect soil CO<Subscript>2</Subscript> emission and soil organic carbon storage in plantations?

Key message

Mixed tree plantations are potential silvicultural systems to increase soil carbon storage through altering litter and root inputs and soil physiochemical properties.

Abstract

Afforestation and reforestation are major strategies for global climate change mitigation. Different tree species composition can induce diverse changes in soil CO₂ emission and soil carbon sequestration in tree plantation. This study employed three plantations of monoculture and mixed Pinus yunnanensis and Eucalyptus globulus to estimate the effect of tree species composition on soil CO₂ emission and soil organic carbon storage in subtropical China. We found that tree species composition had a significant effect on the soil CO₂ emission and soil organic carbon storage. Soil CO₂ emission was lower in the mixed plantation than in the P. yunnanensis plantation, whereas it was higher than in the E. globulus plantation. Differences in soil CO₂ emission among the three plantations were determined by leaf litterfall mass, fine root biomass, soil available nitrogen, pH, soil bulk density, and soil C:N ratio. Soil organic carbon storage was 34.5 and 23.2 % higher in the mixed plantation than in the P. yunnanensis and E. globulus plantations, respectively. Higher soil organic carbon stock in the mixed plantation was attributed to lower C/N ratio of leaf litter and soil, greater fine root biomass and soil organic carbon content, and lower soil CO₂ emission. We conclude that mixed tree plantation can enhance soil carbon sequestration, but can decrease or increase soil CO₂ emission through altering litter and root inputs and soil physiochemical properties.

     

Stomatal conductance in mature deciduous forest trees exposed to elevated CO<Subscript>2</Subscript>

Stomatal conductance (g _s) of mature trees exposed to elevated CO₂ concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g _s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO₂ compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO₂ treatment. During the 2003 drought in the third treatment year, the CO₂ effect became reversed in Carpinus, resulting in higher g _s in trees exposed to elevated CO₂ compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO₂ enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO₂-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO₂.     

Woody-tissue respiration for Simarouba amara and Minquartia guianensis,two tropical wet forest trees with different growth habits

We measured CO₂ efflux from stems of two tropical wet forest trees, both found in the canopy, but with very different growth habits. The species were Simarouba amara, a fast-growing species associated with gaps in old-growth forest and abundant in secondary forest, and Minquartia guianensis, a slow-growing species tolerant of low-light conditions in old-growth forest. Per unit of bole surface, CO₂ efflux averaged 1.24 mol m^–2 s^–1 for Simarouba and 0.83 mol m^–2s^–1 for Minquartia. CO₂ efflux was highly correlated with annual wood production (r ²=0.65), but only weakly correlated with stem diameter (r ²=0.22). We also partitioned the CO₂ efflux into the functional components of construction and maintenance respiration. Construction respiration was estimated from annual stem dry matter production and maintenance respiration by subtracting construction respiration from the instantaneous CO₂ flux. Estimated maintenance respiration was linearly related to sapwood volume (39.6 mol m^–3s^–1 at 24.6° C, r ²=0.58), with no difference in the rate for the two species. Maintenance respiration per unit of sapwood volume for these tropical wet forest trees was roughly twice that of temperate conifers. A model combining construction and maintenance respiration estimated CO₂ very well for these species (r ²=0.85). For our sample, maintenance respiration was 54% of the total CO₂ efflux for Simarouba and 82% for Minquartia. For our sample, sapwood volume averaged 23% of stem volume when weighted by tree size, or 40% with no size weighting. Using these fractions, and a published estimate of aboveground dry-matter production, we estimate the annual cost of woody tissue respiration for primary forest at La Selva to be 220 or 350 g C m^–2 year^–1, depending on the assumed sapwood volume. These costs are estimated to be less than 13% of the gross production for the forest.     

Influence of vegetation and soil CO2 exchange on the concentration and stable oxygen isotope ratio of atmospheric CO2 within a Pinus resinosa canopy

Measurements were made of the concentration and stable oxygen isotopic ratio of carbon dioxide in air samples collected on a diurnal basis at two heights within a Pinus resinosa canopy. Large changes in CO₂ concentration and isotopic composition were observed during diurnal time courses on all three symple dates. In addition, there was strong vertical stratification in the forest canopy, with higher CO₂ concentrations and more negative ¹⁸O values observed closer to the soil surface. The observed daily increases in ¹⁸O values of forest CO₂ were dependent on relative humidity consistent with the modelled predictions of isotopic fractionation during photosynthetic gas exchange. During photosynthetic gas exchange, a portion of the CO₂ that enters the leaf and equilibrates with leaf water is not fixed and diffuses back out of the leaf with an altered oxygen isotopic ratio. The oxygen isotope ratio of CO₂ diffusing out of a leaf depends primarily on the ¹⁸O content of leaf water which changes in response to relative humidity. In contrast, soil respiration caused a decline in the ¹⁸O values of forest CO₂ at night, because CO₂ released from the soil has equilibrated with soil water which has a lower ¹⁸O content than leaf water. The observed relationship between diurnal changes in CO₂ concentration and oxygen isotopic composition in the forest environment were consistent with a gas mixing model that considered the relative magnitudes of CO₂ fluxes associated with photosynthesis, respiration and turbulent exchange between the forest and the bulk atmosphere.     

Effect of fruiting on carbon budgets of apple tree canopies

Summary Carbon budgets were calculated from net photosynthesis and dark respiration measurements for canopies of field-grown, 3-year-old apple trees (Malus domestica Borkh.) with maximum leaf areas of 5.4 m² in a temperature-controlled Perspex tree chamber, measured in situ over 2 years (July 1988 to October 1990) by computerized infrared gas analysis using a dedicated interface and software. Net photosynthesis (Pn) and carbon assimilation per leaf area peaked at respectively 8.3 and 7.7 mol CO₂ m^–2 s^–1 in April. Net photosynthesis (Pn) and dark respiration (Rd) per tree peaked at 3.6 g CO₂ tree^–1 h^–1 (Pn) and 1.2 g CO₂ tree^–1 h^–1 (Rd), equivalent to 4.2 mol CO₂ (Pn) and 1.4 mol CO₂ (Rd) m^–2 s^–1 with maximum carbon gain per tree in August and maximum dark respiration per tree in October 1988 and 1989. In May 1990, a tree was deblossomed. Pn (per tree) of the fruiting apple tree canopy exceeded that of the non-fruiting tree by 2–2.5 fold from June to August 1990, attributed to reduced photorespiration (RI), and resulting in a 2-fold carbon gain of the fruiting over the non-fruiting tree. Dark respiration of the fruiting tree canopy progressively exceeded, with increasing sink strength of the fruit, by 51% (June–August), 1.4-fold (September) and 2-fold (October) that of the non-fruiting tree due to leaf (i. e. not fruit) respiration to provide energy (a) to produce and maintain the fruit on the tree and (b) thereafter to facilitate the later carbohydrate translocation into the woody perennial parts of the tree. The fruiting tree reached its optium carbon budget 2–4 weeks earlier (August) then the non-fruiting tree (September 1990). In the winter, the trunk respired 2–100 g CO₂ month^–1 tree^–1. These data represent the first long-term examination of the effect of fruiting without fruit removal which shows increased dark respiration and with the increase progressing as the fruit developed.     

Elevated atmospheric partial pressure of carbon dioxide and dry matter production of konjak (Amorphophallus konjac K. Koch)

Konjak (Amorphophallus konjac K. Koch) was grown under normal (350 bar) or enriched (700 bar) CO₂ partial pressure in glasshouses kept at 33/26 °C. Doubling the CO₂ partial pressure resulted in twice the yield of corm because the net CO₂ assimilation rate doubled and, due to the simple source-sink relationship, the increased production was partitioned to the corm. The response to CO₂ of assimilation by konjak is discussed in relation to its original habitat in the tropics.     

Leaf and canopy responses of Lolium perenne to long-term elevated atmospheric carbon-dioxide concentration

The relationship between leaf photosynthetic capacity (p _{n, max}), net canopy CO₂- and H₂O-exchange rate (NCER and E _t, respectively) and canopy dry-matter production was examined in Lollium perenne L. cv. Vigor in ambient (363±30 l· l^-1) and elevated (631±43 l·l^-1) CO₂ concentrations. An open system for continuous and simultaneous regulation of atmospheric CO₂ concentration and NCER and E _t measurement was designed and used over an entire growth cycle to calculate a carbon and a water balance. While NCER_max of full-grown canopies was 49% higher at elevated CO₂ level, stimulation of p _n, max was only 46% (in spite of a 50% rise in one-sided stomatal resistance for water-vapour diffusion), clearly indicating the effect of a higher leaf-area index under high CO₂ (approx. 10% in one growing period examined). A larger amount of CO₂-deficient leaves resulted in higher canopy dark-respiration rates and higher canopy light compensation points. The structural component of the high-CO₂ effect was therefore a disadvantage at low irradiance, but a far greater benefit at high irradiance. Higher canopy darkrespiration rates under elevated CO₂ level and low irradiance during the growing period are the primary causes for the increase in dry-matter production (19%) being much lower than expected merely based on the NCER_max difference. While total water use was the same under high and low CO₂ levels, water-use efficiency increased 25% on the canopy level and 87% on a leaf basis. In the course of canopy development, allocation towards the root system became greater, while stimulation of shoot dry-matter accumulation was inversely affected. Over an entire growing season the root/shoot production ratio was 22% higher under high CO₂ concentration.Abbreviations and symbols C350 ambient CO₂, 363±30 l·l^-1 - C600 high CO₂, 631±43 l·l^-1 - c _a atmospheric CO₂ level - c _i CO₂ concentration in the intracellular spaces of the leaf - E_t canopy evapotranspiration - I _o canopy light compensation point - NCER canopy CO₂-exchange rate - p _n leaf photosynthetic rate - PPFD photosynthetic photon flux density - r _a leaf boundary-layer resistance - RD canopy dark-respiration rate - r _s stomatal resistance - WUE water use efficiency     

Effect of changes in water content on photosynthesis,transpiration and discrimination against 13CO2 and C18O16O in Pleurozium and Sphagnum

Photosynthetic gas exchange characteristics of two common boreal forest mosses, Sphagnum (section acutifolia) and Pleurozium schreberi, were measured continuously during the time required for the moss to dry out from full hydration. Similar patterns of change in CO₂ assimilation with variation in water content occurred for both species. The maximum rates of CO₂ assimilation for Sphagnum (approx. 7 mol m^–2 s^–1) occurred at a water content of approximately 7 (fresh weight/dry weight) while for Pleurozium the maximum rate (approx. 2 mol m^–2 s^–1) occurred at a water content of approximately 6 (fresh weight/dry weight). Above and below these water contents CO₂ assimilation declined. In both species total conductance to water vapour (expressed as a percentage of the maximum rates) remained nearly constant at a water content above 9 (fresh weight/dry weight), but below this level declined in a strong linear manner. Short-term, on-line ¹³CO₂ and C¹⁸O¹⁶O discrimination varied substantially with changes in moss water content and associated changes in the ratio of chloroplast CO₂ to ambient CO₂ partial pressure. At full hydration (maximum water content) both Sphagnum and Pleurozium had similar values of ¹³CO₂ discrimination (approx. 15). Discrimination against ¹³CO₂ increased continuously with reductions in water content to a maximum of 27 in Sphagnum and 22 in Pleurozium. In a similar manner C¹⁸C¹⁶O discrimination increased from approximately 30 at full hydration in both species to a maximum of 150 in Sphagnum and 90 in Pleurozium, at low water content. The observed changes in C¹⁸O¹⁶O were strongly correlated to predictions of a mechanistic model of discrimination processes. Field measurements of moss water content suggested that photosynthetic gas exchange by moss in the understory of a black spruce forest was regularly limited by low water content.     

Canopy surface topography in a French Guiana forest and the folded forest theory

The canopy surface is an undulating surface that follows the irregular contours of the upper tree crowns and defines the inner and the outer limits of the forest volume. In French Guiana, the height of the canopy surface was surveyed in both a primary and a 20-years old clear-felled secondary forest plot. The topographic surface was displayed in a three-dimensional mesh, where X and Y are horizontal co-ordinates, and Z is the canopy height measured from the ground with an optical telemeter. The statistical dispersion of Z-data, and the spatial tree height variations, are interpreted at different levels of ecosystem organisation, from forest type (primary or secondary forest) to the trees themselves, following the folded forest model theory (Oldeman 1992, 1994). The vertical growth of trees creates a convex pattern in the relief of canopy surface, whereas gaps make concavities which delimit impact of perturbation on the forest structure. These events are either the result of the dynamic of single trees (emergent and decayed trees), or arise from the dynamic of a group of trees working together (group of emergent trees or complex gaps). At the plot scale, the elementary events, convexities and concavities, are gathered on similar topo-sequences, and form canopy units either higher or lower than the average canopy height. This study suggests that the topography of the canopy surface is defined by a complex nested system from trees, to groups of trees, to canopy units, within a delimited floristic and physical environment.     

Isotope exchange reactions with radiolabeled sulfur compounds in anoxic seawater

The isotope exchange between³⁵S-labeled sulfur compounds of sulfate (SO₄ ^2–), elemental sulfur (S⁰), polysulfide (S_n ^2–), hydrogen sulfide (HS^–: H₂S + HS^– + S^2–), iron sulfide (FeS), and pyrite (FeS₂) was studied at pH 7.6 and 20 °C in anoxic, sterile seawater. Isotope exchange was observed between S⁰, S₂ ^2– HS^–, and FeS, but not between³⁵S labeled SO₄ ^2– or FeS₂ and the other sulfur compounds. Polysulfide mediated the isotope exchange between S⁰ and bisulfide (HS^–). The isotope exchange between S⁰ and S_n ^2–) reached 50% of equilibrium within < 2 min while exchange between S₂ ^2– and HS^– approached equilibrium within 0.5-1 h. In all the experiments HS^–, revealed a fraction exchange from 0.79 to 1.00. Isotope exchange between S^2– and FeS took place only via S₂ ^2– and/or HS^–. The isotope exchange between iron sulfide and the other sulfur compounds was not complete within 24 h as shown by a fraction exchange of 0.07–0.83. This lack of equilibrium (fraction exchange < 1) was due to the isotope exchange between dissolved compounds and surfaces of sulfur particles. The isotopic exchange reactions limit the usefulness of radiotracers in process studies of the inorganic sulfur species. Exchange reactions will also affect the stable isotope distribution among the sulfur species. The kinetics of the isotopic exchange reactions, however, depend on both pH and temperature.     

Remote sensing of forest gas exchange: Considerations derived from a tomographic perspective

The global exchange of gas (CO₂, H₂O) and energy (sensible and latent heat) between forest ecosystems and the atmosphere is often assessed using remote sensing (RS) products. Although these products are essential in quantifying the spatial variability of forest–atmosphere exchanges, large uncertainties remain from a measurement bias towards top of canopy fluxes since optical RS data are not sensitive for the vertically integrated forest canopy. We hypothesize that a tomographic perspective opens new pathways to advance upscaling gas exchange processes from leaf to forest stands and larger scales. We suggest a 3D modelling environment comprising principles of ecohydrology and radiative transfer modelling with measurements of micrometeorological variables, leaf optical properties and forest structure, and assess 3D fields of net CO₂ assimilation (A_n) and transpiration (T) in a Swiss temperate forest canopy. 3D simulations were used to quantify uncertainties in gas exchange estimates inherent to RS approaches and model assumptions (i.e. a big‐leaf approximation in modelling approaches). Our results reveal substantial 3D heterogeneity of forest gas exchange with top of canopy A_n and T being reduced by up to 98% at the bottom of the canopy. We show that a simplified use of RS causes uncertainties in estimated vertical gas exchange of up to 300% and that the spatial variation of gas exchange in the footprint of flux towers can exceed diurnal dynamics. We also demonstrate that big‐leaf assumptions can cause uncertainties up to a factor of 10 for estimates of A_n and T. Concluding, we acknowledge the large potential of 3D assessments of gas exchange to unravelling the role of vertical variability and canopy structure in regulating forest–atmosphere gas and energy exchange. Such information allows to systematically link canopy with global scale controls on forest functioning and eventually enables advanced understanding of forest responses to environmental change.