Sap flow rates and sapwood density are critical factors in within- and between-tree variation in CO2 efflux from stems of mature Dacrydium cupressinum trees

Measurements of CO2 efflux from stems and branches, sap velocity, and respiratory activity of excised wood cores were conducted in Dacrydium cupressinum trees that differed in diameter, age, and canopy emergence. The objective of this study was to determine if consistent linkages exist among respiratory production of CO2 within stems, xylem transport of CO2, and the rate of CO2 diffusing from stem surfaces. Stem CO2 efflux was depressed during periods of sap flow compared with the efflux rate expected for a given stem temperature and was positively correlated with sapwood density. By contrast, no significant relationships were observed between CO2 efflux and the respiratory activity of wood tissues. Between 86 and 91% of woody tissue respiration diffused to the atmosphere over a 24-h period. However, at certain times of the day, xylem transport and internal storage of CO2 may account for up to 13-38% and 12-18%, respectively, of woody tissue respiration. These results demonstrate that differences in sap flow rates and xylem anatomy are critically important for explaining within- and between-tree variation in CO2 efflux from stems.     

Relationship between stem CO2 efflux, stem sap velocity and xylem CO2 concentration in young loblolly pine trees

We measured diel patterns of stem surface CO2 efflux (Es, micromol m(-2) s(-1)), sap velocity (vs, mm s(-1)) and xylem CO2 concentration ([CO2]) (Xs, %) in 8-year-old loblolly pine trees during the spring to determine how vs and Xs influence Es. All trees showed a strong diel hysteresis between Es and stem temperature, where at a given temperature, Es was lower during the day than at night. Diel variations in temperature-independent Es were correlated with vs (R2= 0.54), such that at maximum vs, Es was reduced between 18 and 40%. However, this correlation may not represent a cause-and-effect relationship. In a subset of trees, vs was artificially reduced by progressively removing the tree canopy. Reducing vs to near zero had no effect on Es and did not change the diel hysteretic response to temperature. Diel Xs tended to decrease with vs and increase with Es, however, in defoliated trees, large increases in Xs, when vs approximately 0, had no effect on Es. We conclude that at this time of the year, Es is driven primarily by respiration of cambium and phloem tissues and that sap flow and xylem transport of CO2 had no direct influence on Es.     

The concentration and efflux of tree stem CO2 and the role of xylem sap flow

The accurate assessment of actual tree stem respiration and its relation with temperature plays a considerable role in investigating the forest carbon cycle.An increasing number of research reports have indicated that tree stem respiration determined with the commonlyapplied chamber gas exchange measuring system does not follow expectations regarding temperature relationships.theory that the respired CO2 in a tree stem would all diffuse outward into the atmosphere,However,it neglects partial CO2 that is dissolved in the xylem sap and is carried away by the transpirational stream.Scientists have started to realize that the respired CO2 measured with the chamber gas exchange method is only a portion of the total stem respiration (CO2 efflux),while the other portion,which is sometimes very substantial in quantity (thought to occupy maybe 15%-75% of the total stem respiration),is transported to the upper part of the stem and to the canopy by sap flow.This suggests that the CO2 produced by respiration is re-allocated within the stem.Accordingly,the change in CO2 efflux could be reflected in the rates of sap flow in addition to its dependence on temperature.Proper methods and instruments are required to quantify the internal and external CO2 fluxes in the trunk and their interaction with related environmental factors.     

Transpiration alters the contribution of autotrophic and heterotrophic components of soil CO2 efflux

? An unbiased partitioning of autotrophic and heterotrophic components of soil CO(2) efflux is important to estimate forest carbon budgets and soil carbon sequestration. The contribution of autotrophic sources to soil CO(2) efflux (F(A)) may be underestimated during the daytime as a result of internal transport of CO(2) produced by root respiration through the transpiration stream. ? Here, we tested the hypothesis that carbon isotope composition of soil CO(2) efflux (δ(FS)) in a Eucalyptus plantation grown on a C(4) soil is enriched during the daytime, which will indicate a decrease in F(A) during the periods of high transpiration. ? Mean δ(FS) of soil CO(2) efflux decreased to -25.7‰ during the night and increased to -24.7‰ between 11:00 and 15:00 h when the xylem sap flux density was at its maximum. ? Our results indicate a decrease in the contribution of root respiration to soil CO(2) efflux during the day that may be interpreted as a departure of root-produced CO(2) in the transpiration stream, leading to a 17% underestimation of autotrophic contribution to soil CO(2) efflux on a daily timescale.     

Consumption of coral propagules after mass spawning enhances larval quality of damselfish through maternal effects

Heat and stable isotope tracers were used to study axial and radial water transport in relation to sapwood anatomical characteristics and internal water storage in four canopy tree species of a seasonally dry tropical forest in Panama. Anatomical characteristics of the wood and radial profiles of sap flow were measured at the base, upper trunk, and crown of a single individual of Anacardium excelsum, Ficus insipida, Schefflera morototoni, and Cordia alliodora during two consecutive dry seasons. Vessel lumen diameter and vessel density did not exhibit a consistent trend axially from the base of the stem to the base of the crown. However, lumen diameter decreased sharply from the base of the crown to the terminal branches. The ratio of vessel lumen area to sapwood cross-sectional area was consistently higher at the base of the crown than at the base of the trunk in A. excelsum, F. insipida and C. alliodora, but no axial trend was apparent in S. morototoni. Radial profiles of the preceding wood anatomical characteristics varied according to species and the height at which the wood samples were obtained. Radial profiles of sap flux density measured with thermal dissipation sensors of variable length near the base of the crown were highly correlated with radial profiles of specific hydraulic conductivity (k(s)) calculated from xylem anatomical characteristics. The relationship between sap flux density and k(s) was species-independent. Deuterium oxide (D(2)O) injected into the base of the trunk of the four study trees was detected in the water transpired from the upper crown after only 1 day in the 26-m-tall C. alliodora tree, 2 days in the 28-m-tall F. insipida tree, 3 days in the 38-m-tall A. excelsum tree, and 5 days in the 22-m-tall S. morototoni tree. Radial transport of injected D(2)O was detected in A. excelsum, F. insipida and S. morototoni, but not C. alliodora. The rate of axial D(2)O transport, a surrogate for maximum sap velocity, was positively correlated with the predicted sapwood k(s) and with tree height normalized by the relative diurnal water storage capacity. Residence times for the disappearance of the D(2)O tracer in transpired water ranged from 2 days in C. alliodora to 22 days in A. excelsum and were positively correlated with a normalized index of diurnal water storage capacity. Capacitive exchange of water between stem storage compartments and the transpiration stream thus had a profound influence on apparent rates of axial water transport, the magnitude of radial water movement, and the retention time in the tree of water taken up by the roots. The inverse relationship between internal water exchange capacity and k(s) was consistent with a trade-off contributing to stability of leaf water status through highly efficient water transport at one extreme and release of stored water at the other extreme.     

Origin, fate and significance of CO2 in tree stems.

Although some CO(2) released by respiring cells in tree stems diffuses directly to the atmosphere, on a daily basis 15-55% can remain within the tree. High concentrations of CO(2) build up in stems because of barriers to diffusion in the inner bark and xylem. In contrast with atmospheric [CO(2)] of c. 0.04%, the [CO(2)] in tree stems is often between 3 and 10%, and sometimes exceeds 20%. The [CO(2)] in stems varies diurnally and seasonally. Some respired CO(2) remaining in the stem dissolves in xylem sap and is transported toward the leaves. A portion can be fixed by photosynthetic cells in woody tissues, and a portion diffuses out of the stem into the atmosphere remote from the site of origin. It is now evident that measurements of CO(2) efflux to the atmosphere, which have been commonly used to estimate the rate of woody tissue respiration, do not adequately account for the internal fluxes of CO(2). New approaches to quantify both internal and external fluxes of CO(2) have been developed to estimate the rate of woody tissue respiration. A more complete assessment of internal fluxes of CO(2) in stems will improve our understanding of the carbon balance of trees.     

MRI of long-distance water transport: a comparison of the phloem and xylem flow characteristics and dynamics in poplar, castor bean, tomato and tobacco

We used dedicated magnetic resonance imaging (MRI) equipment and methods to study phloem and xylem transport in large potted plants. Quantitative flow profiles were obtained on a per-pixel basis, giving parameter maps of velocity, flow-conducting area and volume flow (flux). The diurnal xylem and phloem flow dynamics in poplar, castor bean, tomato and tobacco were compared. In poplar, clear diurnal differences in phloem flow profile were found, but phloem flux remained constant. In tomato, only small diurnal differences in flow profile were observed. In castor bean and tobacco, phloem flow remained unchanged. In all plants, xylem flow profiles showed large diurnal variation. Decreases in xylem flux were accompanied by a decrease in velocity and flow-conducting area. The diurnal changes in flow-conducting area of phloem and xylem could not be explained by pressure-dependent elastic changes in conduit diameter. The phloem to xylem flux ratio reflects what fraction of xylem water is used for phloem transport (Münch's counterflow). This ratio was large at night for poplar (0.19), castor bean (0.37) and tobacco (0.55), but low in tomato (0.04). The differences in phloem flow velocity between the four species, as well as within a diurnal cycle, were remarkably small (0.25-0.40 mm s(-1)). We hypothesize that upper and lower bounds for phloem flow velocity may exist: when phloem flow velocity is too high, parietal organelles may be stripped away from sieve tube walls; when sap flow is too slow or is highly variable, phloem-borne signalling could become unpredictable.     

Diurnal changes in branch diameter as indicator of water status of Hinoki cypress Chamaecyparis obtusa

We investigated use of strain gauges for monitoring the water status of trees by measuring changes in the diameter of the largest spreading branch of a 27-year-old Chamaecyparis obtusa tree. The change in xylem diameter in the branch is more closely related than the change in phloem diameter to the change in leaf water potential. Since the diurnal changes in diameter match the diurnal changes in water balance (sap flow velocity - transpiration), measuring the change in xylem diameter using a strain gauge is useful in evaluating the water status of C. obtusa.     

A new measurement technique reveals rapid post-illumination changes in the carbon isotope composition of leaf-respired CO2

We describe an open leaf gas exchange system coupled to a tunable diode laser (TDL) spectroscopy system enabling measurement of the leaf respiratory CO(2) flux and its associated carbon isotope composition (delta(13)C(Rl)) every 3 min. The precision of delta(13)C(Rl) measurement is comparable to that of traditional mass spectrometry techniques. delta(13)C(Rl) from castor bean (Ricinus communis L.) leaves tended to be positively related to the ratio of CO(2) produced to O(2) consumed [respiratory quotient (RQ)] after 24-48 h of prolonged darkness, in support of existing models. Further, the apparent fractionation between respiratory substrates and respired CO(2) within 1-8 h after the start of the dark period was similar to previous observations. In subsequent experiments, R. communis plants were grown under variable water availability to provide a range in delta(13)C of recently fixed carbohydrate. In leaves exposed to high light levels prior to the start of the dark period, CO(2) respired by leaves was up to 11 per thousand more enriched than phloem sap sugars within the first 10-15 min after plants had been moved from the light into the dark. The (13)C enrichment in respired CO(2) then decreased rapidly to within 3-7 per thousand of phloem sap after 30-60 min in the dark. This strong enrichment was not observed if light levels were low prior to the start of the dark period. Measurements of RQ confirmed that carbohydrates were the likely respiratory substrate for plants (RQ > 0.8) within the first 60 min after illumination. The strong (13)C enrichment that followed a high light-to-dark transition coincided with high respiration rates, suggesting that so-called light-enhanced dark respiration (LEDR) is fed by (13)C-enriched metabolites.     

Leaf respiration of snow gum in the light and dark. Interactions between temperature and irradiance

We investigated the effect of temperature and irradiance on leaf respiration (R, non-photorespiratory mitochondrial CO(2) release) of snow gum (Eucalyptus pauciflora Sieb. ex Spreng). Seedlings were hydroponically grown under constant 20 degrees C, controlled-environment conditions. Measurements of R (using the Laisk method) and photosynthesis (at 37 Pa CO(2)) were made at several irradiances (0-2,000 micromol photons m(-2) s(-1)) and temperatures (6 degrees C-30 degrees C). At 15 degrees C to 30 degrees C, substantial inhibition of R occurred at 12 micromol photons m(-2) s(-1), with maximum inhibition occurring at 100 to 200 micromol photons m(-2) s(-1). Higher irradiance had little additional effect on R at these moderate temperatures. The irradiance necessary to maximally inhibit R at 6 degrees C to 10 degrees C was lower than that at 15 degrees C to 30 degrees C. Moreover, although R was inhibited by low irradiance at 6 degrees C to 10 degrees C, it recovered with progressive increases in irradiance. The temperature sensitivity of R was greater in darkness than under bright light. At 30 degrees C and high irradiance, light-inhibited rates of R represented 2% of gross CO(2) uptake (v(c)), whereas photorespiratory CO(2) release was approximately 20% of v(c). If light had not inhibited leaf respiration at 30 degrees C and high irradiance, R would have represented 11% of v(c). Variations in light inhibition of R can therefore have a substantial impact on the proportion of photosynthesis that is respired. We conclude that the rate of R in the light is highly variable, being dependent on irradiance and temperature.     

Dynamics of concentrations and nutrient fluxes in the xylem of Ricinus communis– diurnal course, impact of nutrient availability and nutrient uptake

Diurnal courses of nutrient transport in the xylem and their response to external availability of nutrients were studied. In soil culture, maximal concentrations in all analysed substances were observed during night‐time. Over experimental periods of up to 20 d, concentrations of some ions increased, most by accumulation in the soil. Stringent nutrient conditions were established in a novel pressure chamber. An aeroponic nutrient delivery system inside allows the sampling of xylem sap from intact plants under full control of the nutrient conditions at the root. Analysis of xylem transport under these highly defined conditions established that (1) diurnal variations in concentrations and fluxes in the xylem are dominated by plant‐internal processes; (2) concentrations of nutrients in the xylem sap are highly but specifically correlated with each other; (3) nitrate uptake and nitrate flux to the shoot are largely uncoupled; and (4) in continuous light, diurnal variations of xylem sap concentrations vanish. Step changes in nitrate concentrations of the nutrient solution established that (5) the concomitant increase in nitrate concentration and flux in the xylem is delayed by 2–3 h and is only transient. Diurnal variations of xylem sap composition and use of the new technique to elucidate xylem‐transport mechanisms are discussed.     

Evaluation of the heat pulse velocity technique for measurement of sap flow in rainforest and eucalypt forest species of south-eastern Australia

Sap flow in the stems of two cut saplings each of Eucalyptus maculata (a canopy eucalypt forest tree), Doryphora sassafras and Ceratopetalum apetalum (both canopy rainforest trees of south-eastern coastal Australia) was measured by the heat pulse velocity technique and compared with water uptake from a potometer. Scanning electron micrographs of wounding caused by implantation of temperature sensor and heater probes into the sapwood showed that wounding was similar in rainforest and eucalypt species and was elliptical in shape. A circular wound has been implicitly assumed in previous studies. Accurate measurements of sapling water use were obtained using the smaller transverse wound dimension rather than the larger longitudinal dimension because maximum disruption of sap flow through the xylem vessels occurred in the transverse plane. Accurate measurements of sap flux were obtained above a minimum threshold sap velocity. These velocities were 15·7,10·9 and 9·4 cm h^?1 for E. maculata, C. apetalum and D. sassafras, respectively. Below the threshold sap velocity, however, sap flow could not be accurately calculated from measurements of heat pulse velocity. The minimum threshold sap velocity appeared to be determined by probe construction and xylem anatomy. Despite the elliptical wounding and inaccurate measurement of sap flow below the threshold sap velocity, total sap flow over the experimental period for two saplings of each species was within 7% of water use measured by the potometer.     

Effects of branch solid Fe sulphate implants on xylem sap composition in field-grown peach and pear: changes in Fe, organic anions and pH

The effects of placing solid implants containing Fe sulfate in branches of Fe-deficient pear and peach trees on the composition of the xylem sap have been studied. Iron sulfate implants are commercially used in northeastern Spain to control iron chlorosis in fruit trees. Implants increased Fe concentrations and decreased organic acid concentrations in the xylem sap, whereas xylem sap pH was only moderately changed. The citrate to Fe ratios decreased markedly after implants, therefore improving the possibility that Fe could be reduced by the leaf plasma membrane enzyme reductase, known to be inhibited by high citrate/Fe ratios. In peach, the effects of the implants could be observed many months post treatment. In pear, some effects were still observed one year after the implants had taken place. Results obtained indicate that solid Fe sulfate implants were capable of significantly changing the chemical composition of the xylem sap in fruit trees.     

Changes in the composition of xylem sap during development of the spadix of skunk cabbage (Symplocarpus foetidus)

The spadix of skunk cabbage, Symplocarpus foetidus, is thermogenic and maintains an internal temperature of around 20 degrees C even when the ambient air temperature drops below freezing. This homeothermic heat production is observed only during the stigma stage, and thereafter ceases at the male stage when pollen is shed. To clarify the regulatory mechanism by which the stigma stage-specific heat production occurs in the spadix, sugars, organic acids, and amino acids in xylem sap were analyzed and compared with those of post-thermogenic plants. Interestingly, no significant difference was observed in the total volume of xylem sap per fresh weight of the spadix between thermogenic (31.2+/-24.7 microl h(-1) g(-1)) and post-thermogenic (50.5+/-30.4 microl h(-1) g(-1)) plants. However, concentrations of sugars (sucrose, glucose, and fructose), organic acids (malate and succinate), and amino acids (Asp, Asn, Glu, Gln, Gly, and Ala) in xylem sap decreased remarkably in post-thermogenic plants. Our results indicate that the composition of the xylem sap differs during the development of the spadix of S. foetidus.     

Hydraulic efficiency and safety of branch xylem increases with height in Sequoia sempervirens (D. Don) crowns

The hydraulic limitation hypothesis of Ryan & Yoder (1997, Bioscience 47, 235-242) suggests that water supply to leaves becomes increasingly difficult with increasing tree height. Within the bounds of this hypothesis, we conjectured that the vertical hydrostatic gradient which gravity generates on the water column in tall trees would cause a progressive increase in xylem 'safety' (increased resistance to embolism and implosion) and a concomitant decrease in xylem 'efficiency' (decreased hydraulic conductivity). We based this idea on the historically recognized concept of a safety-efficiency trade-off in xylem function, and tested it by measuring xylem conductivity and vulnerability to embolism of Sequoia sempervirens branches collected at a range of heights. Measurements of resistance of branch xylem to embolism did indeed show an increase in 'safety' with height. However, the expected decrease in xylem 'efficiency' was not observed. Instead, sapwood-specific hydraulic conductivities (Ks) of branches increased slightly, while leaf-specific hydraulic conductivities increased dramatically, with height. The latter could be largely explained by strong vertical gradients in specific leaf area. The increase in Ks with height corresponded to a decrease in xylem wall fraction (a measure of wall thickness), an increase in percentage of earlywood and slight increases in conduit diameter. These changes are probably adaptive responses to the increased transport requirements of leaves growing in the upper canopy where evaporative demand is greater. The lack of a safety-efficiency tradeoff may be explained by opposing height trends in the pit aperture and conduit diameter of tracheids and the major and semi-independent roles these play in determining xylem safety and efficiency, respectively.     

Measurement of stem respiration of sycamore (Platanus occidentalis L.) trees involves internal and external fluxes of CO2 and possible transport of CO2 from roots

CO(2) released by respiring cells in tree stems can either diffuse to the atmosphere or dissolve in xylem sap. In this study, the internal and external fluxes of CO(2) released from respiring stems of five sycamore (Platanus occidentalis L.) trees were calculated. Mean rates of stem respiration were highest in mid-afternoon and lowest at night, and were positively correlated with air temperature. Over a 24 h period, on average 34% of the CO(2) released by respiring cells in the measured stem segment remained within the tree. CO(2) efflux to the atmosphere consisted of similar proportions of CO(2) derived from local respiring cells (55%) and CO(2) that had been transported in the xylem (45%), indicating that CO(2) efflux does not accurately estimate respiration. A portion of the efflux of transported CO(2) appeared to have originated in the root system. A modification of the method for calculating stem respiration based on internal and external fluxes of CO(2) was developed to separate efflux due to local respiration from efflux of transported CO(2).     

Thermal dissipation probe measurements of sap flow in the xylem of trees documenting dynamic relations to variable transpiration given by instantaneous weather changes and the activities of a mistletoe xylem parasite

The thermal dissipation probe was described in the early 1930s for the demonstration of a volume and mass flow of sap in the conductive elements of the xylem in trees. It was subsequently developed further and is now widely used in physiological ecology including measurements in the field. Thermal dissipation demonstrates the occurrence of sap flow and allows determination of its velocity. Here we report simultaneous continuous measurements of sap flow using the thermal dissipation technique and of transpiration by infrared gas analysis for diurnal and annual cycles in a deciduous and an evergreen oak tree, Quercus robur L. and Quercus turneri Willd., respectively, in a deciduous and an evergreen conifer, Larix decidua Mill. and Pinus griffithii McClell., respectively, and the host/mistletoe consortium of the deciduous linden Tilia mandschurica Rupr. & Max. and the evergreen Viscum album L. We show (1) that in diurnal cycles sap flow closely follows dynamic changes of the rate of transpiration elicited by daily fluctuations of weather parameters (sunshine, cloudiness, air temperature and humidity), (2) that in annual cycles sap flow reflects autumnal yellowing and shedding of leaves of the deciduous trees. We report for the first time comparative measurements of sap flow towards mistletoe shoots and host branches in a parasite/host consortium. This demonstrates (3) that mistletoes maintain considerably larger sap flow rates in their xylem conduits than the adjacent host branches dragging the transpiration stream of their host towards their own shoots. We also show (4) that even after the deciduous host has shed its leaves and itself does not transpire any more the evergreen mistletoe towards its shoots can maintain the persistence of a continuous sap flow via the stem and branches of the host as long as frost does not prevent that. The work presented underlines the contention that transpiration is the driving force for sap flow with continuous files of water in the xylem. It shows for the first time that mistletoes direct the flow of water through host roots and stems towards its own shoots by not only performing stronger transpiration as it is known from the literature but also by maintaining larger sap flow rates in the xylem conduits of its stems.     

荷木树干CO_2释放通量与木质部液流和CO_2浓度的关系

采用红外气体分析仪,于2008年10月17-19日连续3个昼夜原位监测了荷木的树干CO_2释放通量、树干温度、木质部液流密度和CO_2浓度.结果表明:树干CO_2释放通量(EA)日变化呈S形曲线,不同径级间差异显著.EA与树干温度呈显著幂函数关系(0.24

Abstract：

By using a Li-820 infra-red CO_2 gas analyzer, an in situ measurement of Schima super-ba stem CO_2 efflux was conducted for three consecutive days from 17 to 19 October 2008. In the meantime, the stem temperature, xylem sap efflux density, and xylem CO_2 concentration were measured. The stem CO_2 efflux had a diurnal variation of "S" pattern, and differed significantly with stem diameter. There was a significant exponential relationship between stem CO_2 efflux and stem temperature (0. 24 < R~2 < 0. 78). The temperature coefficient (b) and regression coeffi-cient (R~2) were higher at nighttime than at daytime, and the Q_(10) value ranged from 2. 01 to 2. 79. The stem CO_2 efflux correlated significantly with the xylem CO_2 concentration, and the best regression curve was cubic (R~2= 0. 48). Excluding the effects of stem temperature, the stem CO_2 efflux showed a significant negative correlation with xylem sap flux density (r =-0.462). Therefore, only using simple temperature function to estimate stem CO_2 efflux would yield a significant error, and xylem sap flux should be taken into consideration in the stem CO_2 emux estimation.     

Temporal variation in δ<Superscript>13</Superscript>C of ecosystem respiration in the Pacific Northwest: links to moisture stress

We measured seasonal and interannual variations in delta(13)C values within the carbon reservoirs (leaves and soil) and CO(2) fluxes (soil and ecosystem respired CO(2)) of an old growth coniferous forest in the Pacific Northwest USA with relation to local meteorological conditions. There were significant intra-annual and interannual differences in the carbon isotope ratios of CO(2) respired at both the ecosystem (delta(13)C(R)) and the soil levels (delta(13)C(R-soil)), but only limited variations in the carbon isotope ratios of carbon stocks. The delta(13)C(R) values varied by as much as 4.4 per thousand over a growing season, while delta(13)C(R-soil )values changed as much as 6.2 per thousand. The delta(13)C of soil organic carbon (delta(13)C(SOC)) and needle organic carbon (delta(13)C(P)) exhibited little or no significant changes over the course of this study. Carbon isotope discrimination within leaves (Delta(p)) showed systematic decreases with increased canopy height, but remained fairly constant throughout the year (Delta(p)=17.9 per thousand -19.2 per thousand at the top of the canopy, Delta(p)=19.6 per thousand -20.9 per thousand at mid-canopy, Delta(p)=23.3 per thousand -25.1 per thousand at the canopy base). The temporal variation in the delta(13)C of soil and ecosystem respired CO(2) was correlated ( r=0.93, P<0.001) with soil moisture levels, with dry summer months having the most (13)C-enriched values. The dynamic seasonal changes in delta(13)C of respired CO(2) are hypothesized to be the result of fast cycling of recently fixed carbon back to the atmosphere. One scaling consequence of the seasonal and interannual variations in delta(13)C(R) is that inversion-based carbon-cycle models dependent on observed atmospheric CO(2) concentration and isotope values may be improved by incorporating dynamic delta(13)C(R) values to interpret regional carbon sink strength.     

Effect of CO₂ on the ventilatory sensitivity to rising body temperature during exercise

We examined the degree to which ventilatory sensitivity to rising body temperature (the slope of the regression line relating ventilation and body temperature) is altered by restoration of arterial PCO(2) to the eucapnic level during prolonged exercise in the heat. Thirteen subjects exercised for ~60 min on a cycle ergometer at 50% of peak O(2) uptake with and without inhalation of CO(2)-enriched air. Subjects began breathing CO(2)-enriched air at the point that end-tidal Pco(2) started to decline. Esophageal temperature (T(es)), minute ventilation (V(E)), tidal volume (V(T)), respiratory frequency (f(R)), respiratory gases, middle cerebral artery blood velocity, and arterial blood pressure were recorded continuously. When V(E), V(T), f(R), and ventilatory equivalents for O(2) uptake (V(E)/VO(2)) and CO(2) output (V(E)/VCO(2)) were plotted against changes in T(es) from the start of the CO(2)-enriched air inhalation (ΔT(es)), the slopes of the regression lines relating V(E), V(T), V(E)/VO(2), and V(E)/VCO(2) to ΔT(es) (ventilatory sensitivity to rising body temperature) were significantly greater when subjects breathed CO(2)-enriched air than when they breathed room air (V(E): 19.8 ± 10.3 vs. 8.9 ± 6.7 l·min(-1)·°C(-1), V(T): 18 ± 120 vs. -81 ± 92 ml/°C; V(E)/VO(2): 7.4 ± 5.5 vs. 2.6 ± 2.3 units/°C, and V(E)/VCO(2): 7.6 ± 6.6 vs. 3.4 ± 2.8 units/°C). The increase in Ve was accompanied by increases in V(T) and f(R). These results suggest that restoration of arterial PCO(2) to nearly eucapnic levels increases ventilatory sensitivity to rising body temperature by around threefold.