Carbon isotope discrimination in photosynthetic bark

We developed and tested a theoretical model describing carbon isotope discrimination during photosynthesis in tree bark. Bark photosynthesis reduces losses of respired CO₂ from the underlying stem. As a consequence, the isotopic composition of source CO₂ and the CO₂ concentration around the chloroplasts are quite different from those of photosynthesizing leaves. We found three lines of evidence that bark photosynthesis discriminates against ¹³C. First, in bark of Populus tremuloides, the '¹³C of CO₂ efflux increased from -24.2‰ in darkness to -15.8‰ in the light. In Pinus monticola, the '¹³C of CO₂ efflux increased from -27.7‰ in darkness to -10.2‰ in the light. Observed increases in '¹³C were generally in good agreement with predictions from the theoretical model. Second, we found that '¹³C of dark-respired CO₂ decreased following 2-3 h of illumination (P<0.01 for Populus tremuloides, P<0.001 for Pinus monticola). These decreases suggest that refixed photosynthate rapidly mixes into the respiratory substrate pool. Third, a field experiment demonstrated that bark photosynthesis influenced whole-tissue '¹³C. Long-term light exclusion caused a localized increase in the '¹³C of whole bark and current-year wood in branches of P. monticola (P<0.001 and P<0.0001, respectively). Thus bark photosynthesis was shown to discriminate against ¹³C and create a pool of photosynthate isotopically lighter than the dark respiratory pool in all three experiments. Failure to account for discrimination during bark photosynthesis could interfere with interpretation of the '¹³C in woody tissues or in woody-tissue respiration.     

13C content of ecosystem respiration is linked to precipitation and vapor pressure deficit

Variation in the carbon isotopic composition of ecosystem respiration ('¹³C_R) was studied for 3 years along a precipitation gradient in western Oregon, USA, using the Keeling plot approach. Study sites included six coniferous forests, dominated by Picea sitchensis, Tsuga heterophylla, Pseudotsuga menziesii, Pinus ponderosa, and Juniperus occidentalis, and ranged in location from the Pacific coast to the eastern side of the Cascade Mountains (a 250-km transect). Mean annual precipitation across these sites ranged from 227 to 2,760 mm. Overall '¹³C_R varied from -23.1 to -33.1‰, and within a single forest, it varied in magnitude by 3.5-8.5‰. Mean annual '¹³C_R differed significantly in the forests and was strongly correlated with mean annual precipitation. The carbon isotope ratio of carbon stocks (leaves, fine roots, litter, and soil organic matter) varied similarly with mean precipitation (more positive at the drier sites). There was a strong link between '¹³C_R and the vapor saturation deficit of air (vpd) 5-10 days earlier, both across and within sites. This relationship is consistent with stomatal regulation of gas exchange and associated changes in photosynthetic carbon isotope discrimination. Recent freeze events caused significant deviation from the '¹³C_R versus vpd relationship, resulting in higher than expected '¹³C_R values.     

Microbial community composition and function beneath temperate trees exposed to elevated atmospheric carbon dioxide and ozone

We hypothesized that changes in plant growth resulting from atmospheric CO₂ and O₃ enrichment would alter the flow of C through soil food webs and that this effect would vary with tree species. To test this idea, we traced the course of C through the soil microbial community using soils from the free-air CO₂ and O₃ enrichment site in Rhinelander, Wisconsin. We added either ¹³C-labeled cellobiose or ¹³C-labeled N-acetylglucosamine to soils collected beneath ecologically distinct temperate trees exposed for 3 years to factorial CO₂ (ambient and 200 µl l^-1 above ambient) and O₃ (ambient and 20 µl l^-1 above ambient) treatments. For both labeled substrates, recovery of ¹³C in microbial respiration increased beneath plants grown under elevated CO₂ by 29% compared to ambient; elevated O₃ eliminated this effect. Production of ¹³C-CO₂ from soils beneath aspen (Populus tremuloides Michx.) and aspen-birch (Betula papyrifera Marsh.) was greater than that beneath aspen-maple (Acer saccharum Marsh.). Phospholipid fatty acid analyses (¹³C-PLFAs) indicated that the microbial community beneath plants exposed to elevated CO₂ metabolized more ¹³C-cellobiose, compared to the microbial community beneath plants exposed to the ambient condition. Recovery of ¹³C in PLFAs was an order of magnitude greater for N-acetylglucosamine-amended soil compared to cellobiose-amended soil, indicating that substrate type influenced microbial metabolism and soil C cycling. We found that elevated CO₂ increased fungal activity and microbial metabolism of cellobiose, and that microbial processes under early-successional aspen and birch species were more strongly affected by CO₂ and O₃ enrichment than those under late-successional maple.     

CO2 exchange and thallus nitrogen across 75 contrasting lichen associations from different climate zones

Aiming to investigate whether a carbon-to-nitrogen equilibrium model describes resource allocation in lichens, net photosynthesis (NP), respiration (R), concentrations of nitrogen (N), chlorophyll (Chl), chitin and ergosterol were investigated in 75 different lichen associations collected in Antarctica, Arctic Canada, boreal Sweden, and temperate/subtropical forests of Tenerife, South Africa and Japan. The lichens had various morphologies and represented seven photobiont and 41 mycobiont genera. Chl a, chitin and ergosterol were used as indirect markers of photobiont activity, fungal biomass and fungal respiration, respectively. The lichens were divided into three groups according to photobiont: (1) species with green algae, (2) species with cyanobacteria, and (3) tripartite species with green algal photobionts and cyanobacteria in cephalodia. Across species, thallus N concentration ranged from 1 to 50 mg g^-1 dry wt., NP varied 50-fold, and R 10-fold. In average, green algal lichens had the lowest, cyanobacterial Nostoc lichens the highest and tripartite lichens intermediate N concentrations. All three markers increased with thallus N concentration, and lichens with the highest Chl a and N concentrations had the highest rates of both P and R. Chl a alone accounted for ca. 30% of variation in NP and R across species. On average, the photosynthetic efficiency quotient [K_F=(NP_max+R)/R)] ranged from 2.4 to 8.6, being higher in fruticose green algal lichens than in foliose Nostoc lichens. The former group invested more N in Chl a and this trait increased NP_max while decreasing R. In general terms, the investigated lichens invested N resources such that their maximal C input capacity matched their respiratory C demand around a similar (positive) equilibrium across species. However, it is not clear how this apparent optimisation of resource use is regulated in these symbiotic organisms.     

Tissue structure and respiration of stems of <Emphasis Type="Italic">Betula pendula</Emphasis> under contrasting ozone exposure and nutrition

Tissue structure and respiration (Rs) of stems were analyzed in Betula pendula grown throughout the growing season in either filtered air (control) or 90/40 nl O₃ l^-1 (day/night). Both regimes were split into high and low nutrient supply. High nutrition increased tissue and cell sizes within the stem xylem, phloem and periderm, whereas ozone (O₃) tended to reduce tissue widths, inhibiting in particular the cambial activity of xylem growth in low-fertilized, O₃-exposed plants (O₃/LF). Callose deposition was enhanced in the phloem sieve plates and tannins tended to condense in vacuoles of parenchyma cells under O₃ stress. Decline occurred close to lenticels, related to O₃ impact during shoot differentiation and was probably exacerbated by the limited assimilate translocation. Radial stem growth ceased 4 weeks earlier than in control plants; however, the area-based Rs was enhanced during intense growth in high-fertilized, O₃-exposed plants. Photosynthetic CO₂ refixation of stems reached about 50% of their dark respiration rate and the relative growth rate (RGR) did not differ between treatments. At high nutrition, RGR enhanced Rs to levels twice as high as the maintenance level. Unit construction costs appeared to be similar in each treatment, although CO₂ release on a volume-increment basis was lowered by 45% in O₃/LF plants. This latter effect is ascribed to lowered maintenance demands of a xylem remaining reduced in width by 50%. The high respiratory costs in the carbon balance of O₃/LF plants result from an enhanced leaf rather than stem respiration, given the high demand for stress compensation in the foliage.     

Water relations of coastal and estuarine Rhizophora mangle: xylem pressure potential and dynamics of embolism formation and repair

Physiological traits related to water transport were studied in Rhizophora mangle (red mangrove) growing in coastal and estuarine sites in Hawaii. The magnitude of xylem pressure potential (P_x), the vulnerability of xylem to cavitation, the frequency of embolized vessels in situ, and the capacity of R. mangle to repair embolized vessels were evaluated with conventional and recently developed techniques. The osmotic potential of the interstitial soil water (?_sw) surrounding the roots of R. mangle was c. -2.6LJ.52᎒^-3 and -0.4Lj.13᎒^-3 MPa in the coastal and estuarine sites, respectively. Midday covered (non-transpiring) leaf water potentials (O_L) determined with a pressure chamber were 0.6-0.8 MPa more positive than those of exposed, freely-transpiring leaves, and osmotic potential of the xylem sap (?_x) ranged from -0.1 to -0.3 MPa. Consequently, estimated midday values of P_x (calculated by subtracting ?_x from covered O_L) were about 1 MPa more positive than O_L determined on freely transpiring leaves. The differences in O_L between covered and transpiring leaves were linearly related to the transpiration rates. The slope of this relationship was steeper for the coastal site, suggesting that the hydraulic resistance was larger in leaves of coastal R. mangle plants. This was confirmed by both hydraulic conductivity measurements on stem segments and high-pressure flowmeter studies made on excised leafy twigs. Based on two independent criteria, loss of hydraulic conductivity and proportions of gas- and liquid-filled vessels in cryo-scanning electron microscope (cryo-SEM) images, the xylem of R. mangle plants growing at the estuarine site was found to be more vulnerable to cavitation than that of plants growing at the coastal site. However, the cryo-SEM analyses suggested that cavitation occurred more readily in intact plants than in excised branches that were air-dried in the laboratory. Cryo-SEM analyses also revealed that, in both sites, the proportion of gas-filled vessels was 20-30% greater at midday than at dawn or during the late afternoon. Refilling of cavitated vessels thus occurred during the late afternoon when considerable tension was present in neighboring vessels. These results and results from pressure-volume relationships suggest that R. mangle adjusts hydraulic properties of the water-transport system, as well as the leaf osmotic potential, in concert with the environmental growing conditions.     

Photosynthetic oxygen production and community respiration in the Indian sector of the Antarctic Ocean during austral summer

Photosynthetic oxygen production by phytoplankton and community respiration in the Indian sector of the Antarctic Ocean were estimated from changes in oxygen concentrations in light and dark bottles. Gross production varied between 0.1 and 5.1 µmol O₂ l^-1 day^-1. In the same water, community respiration (the sum of oxygen consumption by heterotrophs and phytoplankton) was 0.4-3.6 µmol O₂ l^-1 day^-1, which accounted for 47-343% of the gross production. Algal and heterotrophic respirations were distinguished using some assumptions. These estimates showed that heterotrophic respiration accounted for most of the community respiration (70-91% depending upon the assumptions), indicating that heterotrophic respiration plays an important role in the mineralization of phytoplankton production in the surveyed sea area. Gross production rate correlated with chlorophyll a concentration, showing that the photosynthetic production rate of oxygen depends on the abundance of phytoplankton. Moreover, there was a significant relationship between gross production and community respiration rates. These regression equations suggested that negative net production occurred under the usually low concentration of chlorophyll observed in the Indian sector of the Antarctic Ocean. Hence, the net exchange of carbon dioxide due to biological processes through the sea surface seemed to be not as large as expected in the Antarctic Ocean, although the number of data were limited at this stage.     

The effect of elevated carbon dioxide and ozone on leaf- and branch-level photosynthesis and potential plant-level carbon gain in aspen

Two aspen (Populus tremuloides Michx.) clones, differing in O₃ tolerance, were grown in a free-air CO₂ enrichment (FACE) facility near Rhinelander, Wisconsin, and exposed to ambient air, elevated CO₂, elevated O₃ and elevated CO₂+O₃. Leaf instantaneous light-saturated photosynthesis (P_S) and leaf areas (A) were measured for all leaves of the current terminal, upper (current year) and the current-year increment of lower (1-year-old) lateral branches. An average, representative branch was chosen from each branch class. In addition, the average photosynthetic rate was estimated for the short-shoot leaves. A summing approach was used to estimate potential whole-plant C gain. The results of this method indicated that treatment differences were more pronounced at the plant- than at the leaf- or branch-level, because minor effects within modules accrued in scaling to plant level. The whole-plant response in C gain was determined by the counteracting changes in P_S and A. For example, in the O₃-sensitive clone (259), inhibition of P_S in elevated O₃ (at both ambient and elevated CO₂) was partially ameliorated by an increase in total A. For the O₃-tolerant clone (216), on the other hand, stimulation of photosynthetic rates in elevated CO₂ was nullified by decreased total A.     

The Transport of Photosynthetic Products from the Chloroplasts of Tobacco Leaves

Tobacco leaves were fractionated by the non-aqueous method afterphotesynthesis for short periods in ¹⁴CO₂ and with or withoutsubsequent photosynthesis in ¹²CO₂ or respiration in the dark.Phosphate esters became labelled only slowly in the non-chloroplastparts of the leaf; glycine and serine which became rapidly labelledin the non-chloroplast leaf fraction appear to be concernedin the transport of newly assimilated carbon from the chloroplast.     

Forest carbon balance under elevated CO2

Free-air CO₂ enrichment (FACE) technology was used to expose a loblolly pine (Pinus taeda L.) forest to elevated atmospheric CO₂ (ambient + 200 µl l^-1). After 4 years, basal area of pine trees was 9.2% larger in elevated than in ambient CO₂ plots. During the first 3 years the growth rate of pine was stimulated by ~26%. In the fourth year this stimulation declined to 23%. The average net ecosystem production (NEP) in the ambient plots was 428 gC m^-2 year^-1, indicating that the forest was a net sink for atmospheric CO₂. Elevated atmospheric CO₂ stimulated NEP by 41%. This increase was primarily an increase in plant biomass increment (57%), and secondarily increased accumulation of carbon in the forest floor (35%) and fine root increment (8%). Net primary production (NPP) was stimulated by 27%, driven primarily by increases in the growth rate of the pines. Total heterotrophic respiration (R_h) increased by 165%, but total autotrophic respiration (R_a) was unaffected. Gross primary production was increased by 18%. The largest uncertainties in the carbon budget remain in separating belowground heterotrophic (soil microbes) and autotrophic (root) respiration. If applied to temperate forests globally, the increase in NEP that we measured would fix less than 10% of the anthropogenic CO₂ projected to be released into the atmosphere in the year 2050. This may represent an upper limit because rising global temperatures, land disturbance, and heterotrophic decomposition of woody tissues will ultimately cause an increased flux of carbon back to the atmosphere.     

Refixation of respiratory CO2 in the ears of C3 cereals

The spatial arrangement of tissues within the ears of cereals,and gas exchange measurements on intact ears of barley and durumwheat suggest that respiratory CO₂ associated with grain-fillingprocesses, may be refixed close to its site of evolution. Apparentrefixation of respiratory CO₂ in intact ears was compared withthat in flag leaves, by feeding both organs with ¹⁴C-labelledsucrose and trapping ¹⁴CO₂ released by respiration. Apparentrefixation in the ears was twice that measured in flag leafblades of durum wheat genotype Durelle. In ears, the capacityof refixation of respiratory CO₂ at 210 mmol mol^–1 O₂ranged from 55% in barley genotype Roxana to 75% in barley genotypeHatif, and 60% in duwm wheat genotype Bidi 17. A low O₂ concentrationincreased refixation of respiratory CO₂ by up to 90%, 92% and82%, respectively. The occurrence of CO₂ refixation in the field,in a set of 12 barley genotypes grown under irrigated and rainfedMediterranean field conditions, was consistent with observedcarbon isotope ratios (     

The Respiration of Mature Field Bean (Vicia faba L.) Leaves During Prolonged Darkness

Dark respiration in attached and detached mature leaves of thefield bean (Vicia faba L.) was studied whilst leaves experiencedup to 60 h of darkness. The results showed: (1) the initialrespiration rate to vary according to the irradiance duringthe previous photoperiod; (2) the dark respiration rate (perunit area) of attached leaves to be essentially constant duringa normal 12 h night although there was a rapid loss in leafd. wt during this time; (3) after 12 h, the respiration rateof attached leaves decayed to an asymptotic value at about 36h; (4) the respiration rate of leaves detached at the end ofthe photoperiod and maintained in the dark on deionised water,decayed only after 36 h of darkness; (5) there was no differencebetween the respiration rate of attached and detached leavesduring the normal 12 h night. It is concluded that the dark respiration of attached fieldbean leaves is intially related to the synthesis and translocationof sucrose in addition to maintenance. After about 36 h, whenthe rate of CO₂ efflux is more or less steady, the CO₂ effluxreflects the intensity of maintenance processes only. The maintenancerespiration rate (determined after 60 h in the dark) rangedfrom 062 to 151 mg CO₂ (g d. wt)^–1 h^–1 but wasrelatively unaffected by several applied treatments. Vicia faba L., field bean, respiration, maintenance, nitrate, non-structural carbohydrate, export     

The respiratory energy requirements involved in nocturnal carbohydrate export from starch-storing mature source leaves and their contribution to leaf dark respiration

The present study explores the potential contribution of theenergy requirements associated with nocturnal carbohydrate exportto (1) the fraction of dark respiration correlating with leafnitrogen concentration and (2) the dark respiration of maturesource leaves. To this end, we determined the nocturnal carbohydrate-exportrates from leaves with an optimal nitrogen supply, and the correlationbetween the nitrogen concentration and the dark respirationof leaves. The specific energy costs of carbohydrate exportfrom starch-storing source leaves were determined both experimentallyand theoretically. The present estimate of the specific energycost involved in carbohydrate export as obtained by linear regression(0.70 mol CO₂ [mol sucrose]^–1), agrees well with bothliterature data obtained by different methods (0.47 to 1.26mol CO₂ [mol sucrose]^–1) and the theoretically calculatedrange for starch-storing species (0.40 to 1.20 mol CO₂ [molsucrose]^–1). The conversion of starch in the chloroplastto sucrose in the cytosol is a major energy-requiring process.Maximally 42 to ‘107’% of the slope of the relationshipbetween respiration rate and organic nitrogen concentrationof primary bean leaves, may be ascribed to the energy costsassociated with nocturnal export of carbohydrates. Total energycosts associated with export were derived from the product ofthe specific costs of carbohydrate export and the export rates,either measured on full-grown (primary) leaves of potato andbean or derived from the literature. These export costs account,on average, for 29% of the dark respiration rate in starch-storingspecies. We conclude that nocturnal carbohydrate export is amajor energy-requiring process in starch-storing species Key words: Carbohydrate export, leaf dark respiration, nitrogen concentration, respiratory costs, specific energy cost     

Photosynthetic induction in saplings of three shade-tolerant tree species: comparing understorey and gap habitats in a French Guiana rain forest

The photosynthetic induction response under constant and fluctuating light was examined in naturally occurring saplings (about 0.5-2 m in height) of three shade-tolerant tree species, Pourouma bicolor spp digitata, Dicorynia guianensis, and Vouacapoua americana, growing in bright gaps and in the shaded understorey in a Neotropical rain forest. Light availability to saplings was estimated by hemispherical photography. Photosynthetic induction was measured in the morning on leaves that had not yet experienced direct sunlight. In Dicorynia, the maximum net photosynthesis rate (A_max) was similar between forest environments (ca 4 µmol m^-2 s^-1), whereas for the two other species, it was twice as high in gaps (ca 7.5) as in the understorey (ca 4.5). However, the time required to reach 90% of A_max did not differ among species, and was short, 7-11 min. Biochemical induction was fast in leaves of Pourouma, as about 3 min were needed to reach 75% of maximum carboxylation capacity (V_cmax); the two other species needed 4-5 min. When induction continued after reaching 75% of V_cmax, stomatal conductance increased in Pourouma only (ca 80%), causing a further increase in its net photosynthesis rate. When fully induced leaves were shaded for 20 min, loss of induction was moderate in all species. However, gap saplings of Dicorynia had a rapid induction loss (ca 80%), which was mainly due to biochemical limitation as stomatal conductance decreased only slowly. When leaves were exposed to a series of lightflecks separated by short periods of low light, photosynthetic induction increased substantially and to a similar extent in all species. Although A_max was much lower in old than in young leaves as measured in Dicorynia and Vouacapoua, variables of the dynamic response of photosynthesis to a change in light tended to be similar between young and old leaves. Old leaves, therefore, might remain important for whole-plant carbon gain, especially in understorey environments. The three shade-tolerant species show that, particularly in low light, they are capable of efficient sunfleck utilization.     

Very high productivity of the C4 aquatic grass Echinochloa polystachya in the Amazon floodplain confirmed by net ecosystem CO2 flux measurements

Fluxes of CO₂ and H₂O vapour from dense stands of the C4 emergent macrophyte grass Echinochloa polystachya were measured by eddy covariance in both the low water (LW) and high water (HW, flooded) phases of the annual Amazon river cycle at Manaus, Brazil. Typical clear-sky midday CO₂ uptake rates by the vegetation stand (including detritus, sediment or water surface) were 30 and 35 µmol CO₂ (ground) m^-2 s^-1 in the LW and HW periods, respectively. A rectangular hyperbola model fitted the responses of "instantaneous" (20- or 30-min average) net CO₂ exchange rates to incident photosynthetic photon flux densities (PFD) well. Stand evaporation rates were linearly related to PFD. The major difference in CO₂ uptake rates between the two periods was the larger respiration flux during LW due to the CO₂ efflux from sediment, roots and litter. Integrated 20- or 30-min fluxes were used to derive relationships between daily CO₂ and H₂O vapour fluxes and incident radiation. The daily CO₂ fluxes were almost linearly related to incident radiation, but there was evidence of saturation at the highest daily radiation totals. Annual productivity estimated from the daily model in 1996-1997 agreed closely with that previously estimated for 1985-1986 from a leaf-scale photosynthetic model, but were some 15% less than those derived at that time from biomass harvests. Both CO₂ uptake and water use efficiency were comparable with those found in fertilised maize fields in warm temperate conditions.     

Effects of Prolonged Darkness on the Sensitivity of Leaf Respiration to Carbon Dioxide Concentration in C3and C4Species

Predicting responses of plant and global carbon balance to theincreasing concentration of carbon dioxide in the atmosphererequires an understanding of the response of plant respirationto carbon dioxide concentration ([CO₂]). Direct effects of thecarbon dioxide concentration at which rates of respiration ofplant tissue are measured are quite variable and their effectsremain controversial. One possible source of variation in responsivenessis the energy status of the tissue, which could influence thecontrol coefficients of enzymes, such as cytochrome-c oxidase,whose activity is sensitive to [CO₂]. In this study we comparedresponses of respiration rate to [CO₂] over the range of 60to 1000 µmol mol^-1in fully expanded leaves of four C₃andfour C₄herbaceous species. Responses were measured near themiddle of the normal 10 h dark period, and also after another24 h of darkness. On average, rates of respiration were reducedabout 70% by the prolonged dark period, and leaf dry mass perunit area decreased about 30%. In all species studied, the relativedecrease in respiration rate with increasing [CO₂] was largerafter prolonged darkness. In the C₃species, rates measured at1000 µmol mol^-1CO₂averaged 0.89 of those measured at 60µmol mol^-1in the middle of the normal dark period, and0.70-times when measured after prolonged darkness. In the C₄species,rates measured at 1000 µmol mol^-1CO₂averaged 0.79 of thoseat 60 µmol mol^-1CO₂in the middle of the normal dark period,and 0.51-times when measured after prolonged darkness. In threeof the C₃species and one of the C₄species, the decrease in theabsolute respiration rate between 60 and 1000 µmol mol^-1CO₂wasessentially the same in the middle of the normal night periodand after prolonged darkness. In the other species, the decreasein the absolute rate of respiration with increase in [CO₂] wassubstantially less after prolonged darkness than in the middleof the normal night period. These results indicated that increasingthe [CO₂] at the time of measurement decreased respiration inall species examined, and that this effect was relatively largerin tissues in which the respiration rate was substrate-limited.The larger relative effect of [CO₂] on respiration in tissuesafter prolonged darkness is evidence against a controlling roleof cytochrome-c oxidase in the direct effects of [CO₂] on respiration.Copyright 2001 Annals of Botany Company Carbon dioxide, respiration, Abutilon theophrasti(L.), Amaranthus retroflexus(L.),Amaranthus hypochondriacus (L.), Datura stramonium(L.), Helianthus annuus(L.), Solanum melongena(L.), Sorghum bicolor(L. Moench), Zea mays     

Stem radius changes and their relation to stored water in stems of young Norway spruce trees

Changes in the stem radius of young Norway spruce [Picea abies (L.) Karst.] were related to changes in stem water content in order to investigate the relationship between diurnal stem size fluctuations and internally stored water. Experiments were performed on living trees and on cut stem segments. The defoliated stem segments were dried under room conditions and weight (W), volume (V), and xylem water potential (O_s) were continuously monitored for 95 h. Additionally, photos of cross-sections of fresh and air-dried stem segments were taken. For stem segments we found that the change in V was linearly correlated to the change in W as long as O_s was >-2.3ǂ.3 MPa (phase transition point). Stem contraction occurred almost solely in the elastic tissues of the bark (cambium, phloem, and parenchyma), and the stem radius changes were closely coupled to bark water content. For living trees, it is therefore possible to estimate the daily contribution of "bark water" to transpiration from knowledge of the stem size and continuous measurements of the stem radius fluctuations. When O_s reaches the phase-transition point, water is also withdrawn from the inelastic tissue of the stem (xylem), which - in the experiment with stem segments - was indicated by an increasing ratio between (V and (W. We assume that for O_s below the transition point, air is sucked into the tracheids (cavitation) and water is also withdrawn from the xylem. Due to the fact that in living P. abies O_s rarely falls below -2.3ǂ.3 MPa and the xylem size is almost unaffected by radius fluctuations, dendrometers are useful instruments with which to derive the diurnal changes in the bark water contents of Norway spruce trees.     

Synthesis and Dissolution of Starch labelled with 14Carbon in Tobacco Leaf Tissue

Tobacco leaves depleted of starch, were detached and allowedto assimilate equal amounts of ¹⁴CO₂ and ¹²CO₂in succession,and vice versa. Distribution of radioactivity in starch, andsugars was determined after assimilation and after disks cutfrom the leaves had been kept in darkness for times up to 40hours. The amount and activity of the CO₂ was also determined.¹⁴C and ¹²C were incorporated in equal amounts into starch independentlyof the order in which they were supplied. In contrast sucrosehad high activity ¹⁴C was given last, and hexose a low one.The reverse was true when ¹²C was given last. Activity of respiratoryCO₂ was slightly higher when ¹⁴C was assimilated last as comparedwith ¹²C. In the dark only ¹⁴C or ¹²C was at first lost fromstarch, in accordance or removal of discrete layers. Analyticalresults show that starch is the main respiratory substrate andto account for the redistribution of radioactivity in passageto CO₂ it is concluded that sucrose occurs at two sites separatedinter-or intra-cellularly, one of which is in equilibrium withthe system intrconverting starch and CO₂ and at the other hexosesare produced by inversion. A starch-like polysaccharide is formedduring assimilation which persists in the dark and there isa significant contribution to respiration of carbon from non-carbohydratesources when leaf disks are kept on the dark.     

14C transfer between the spring ephemeral Erythronium americanum and sugar maple saplings via arbuscular mycorrhizal fungi in natural stands

We investigated in the field the carbon (C) transfer between sugar maple (Acer saccharum) saplings and the spring ephemeral Erythronium americanum via the mycelium of arbuscular mycorrhizal (AM) fungi. Sugar maple saplings and E. americanum plants were planted together in pots placed in the ground of a maple forest in 1999. Ectomycorrhizal yellow birches (Betula alleghaniensis) were added as control plants. In spring 2000, during leaf expansion of sugar maple saplings, the leaves of E. americanum were labelled with ¹⁴CO₂. Seven days after labelling, radioactivity was detected in leaves, stem and roots of sugar maples. Specific radioactivity in sugar maples was 13-fold higher than in yellow birches revealing the occurrence of a direct transfer of ¹⁴C between the AM plants. The quantity of ¹⁴C transferred to sugar maple saplings was negatively correlated with the percentage of ¹⁴C allocated to the storage organ of E. americanum. A second labelling was performed in autumn 2000 on sugar maple leaves during annual growth of E. americanum roots. Radioactivity was detected in 7 of 22 E. americanum root systems and absent in yellow birches. These results suggest that AM fungi connecting different understorey species can act as reciprocal C transfer bridges between plant species in relation with the phenology of the plants involved.     

Fine-root biomass and fluxes of soil carbon in young stands of paper birch and trembling aspen as affected by elevated atmospheric CO2 and tropospheric O3

Rising atmospheric CO₂ may stimulate future forest productivity, possibly increasing carbon storage in terrestrial ecosystems, but how tropospheric ozone will modify this response is unknown. Because of the importance of fine roots to the belowground C cycle, we monitored fine-root biomass and associated C fluxes in regenerating stands of trembling aspen, and mixed stands of trembling aspen and paper birch at FACTS-II, the Aspen FACE project in Rhinelander, Wisconsin. Free-air CO₂ enrichment (FACE) was used to elevate concentrations of CO₂ (average enrichment concentration 535 µl l^-1) and O₃ (53 nl l^-1) in developing forest stands in 1998 and 1999. Soil respiration, soil pCO₂, and dissolved organic carbon in soil solution (DOC) were monitored biweekly. Soil respiration was measured with a portable infrared gas analyzer. Soil pCO₂ and DOC samples were collected from soil gas wells and tension lysimeters, respectively, at depths of 15, 30, and 125 cm. Fine-root biomass averaged 263 g m^-2 in control plots and increased 96% under elevated CO₂. The increased root biomass was accompanied by a 39% increase in soil respiration and a 27% increase in soil pCO₂. Both soil respiration and pCO₂ exhibited a strong seasonal signal, which was positively correlated with soil temperature. DOC concentrations in soil solution averaged ~12 mg l^-1 in surface horizons, declined with depth, and were little affected by the treatments. A simplified belowground C budget for the site indicated that native soil organic matter still dominated the system, and that soil respiration was by far the largest flux. Ozone decreased the above responses to elevated CO₂, but effects were rarely statistically significant. We conclude that regenerating stands of northern hardwoods have the potential for substantially greater C input to soil due to greater fine-root production under elevated CO₂. Greater fine-root biomass will be accompanied by greater soil C efflux as soil respiration, but leaching losses of C will probably be unaffected.