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Rates of CO2 efflux of stems and branches are highly variable among and within trees and across stands. Scaling factors have only partially succeeded in accounting for the observed variations. In this study, the resistance to radial CO2 diffusion was quantified for tree stems of an eastern cottonwood (Populus deltoides Bartr. ex Marsh.) clone by direct manipulation of the CO2 concentration ([CO2]) of xylem sap under controlled conditions. Tree-specific linear relationships between rates of stem CO2 efflux (JO) and xylem [CO2] were found. The resistance to radial CO2 diffusion differed 6-fold among the trees and influenced the balance between the amount of CO2 retained in the xylem v. that which diffused to the atmosphere. Therefore, we hypothesised that variability in the resistance to radial CO2 diffusion might be an overlooked cause for the inconsistencies and large variations in woody tissue CO2 efflux. It was found that transition from light to dark conditions caused a rapid increase in JO and xylem [CO2], both in manipulated trees and in an intact tree with no sap manipulation. This resulted in an increased resistance to radial CO2 diffusion during the dark, at least for trees with smaller daytime resistances. Stem diameter changes measured in the intact tree supported the idea that higher actual respiration rates occurred at night owing to higher metabolism in relation to an improved water status and higher turgor pressure.

Oxidative respiration is strongly temperature driven. However, in woody stems, efflux of CO(2) to the atmosphere (E (A)), commonly used to estimate the rate of respiration (R (S)), and stem temperature (T (st)) have often been poorly correlated, which we hypothesized was due to transport of respired CO(2) in xylem sap, especially under high rates of sap flow (f (s)). To test this, we measured E (A), T (st), f (s) and xylem sap CO(2) concentrations ([CO(2)*]) in 3-year-old Populus deltoides trees under different weather conditions (sunny and rainy days) in autumn. We also calculated R (S) by mass balance as the sum of both outward and internal CO(2) fluxes and hypothesized that R (S) would correlate better with T (st) than E (A). We found that E (A) sometimes correlated well with T (st), but not on sunny mornings and afternoons or on rainy days. When the temperature effect on E (A) was accounted for, a clear positive relationship between E (A) and xylem [CO(2)*] was found. [CO(2)*] varied diurnally and increased substantially at night and during periods of rain. Changes in [CO(2)*] were related to changes in f (s) but not T (st). We conclude that changes in both respiration and internal CO(2) transport altered E (A). The dominant component flux of R (S) was E (A). However, on a 24-h basis, the internal transport flux represented 9-18% and 3-7% of R (S) on sunny and rainy days, respectively, indicating that the contribution of stem respiration to forest C balance may be larger than previously estimated based on E (A) measurements. Unexpectedly, the relationship between R (S) and T (st) was sometimes weak in two of the three trees. We conclude that in addition to temperature, other factors such as water deficits or substrate availability exert control on the rate of stem respiration so that simple temperature functions are not sufficient to predict stem respiration.

Stem photosynthesis can contribute significantly to woody plant carbon balance, particularly in times when leaves are absent or in 'open' crowns with sufficient light penetration. We explored the significance of woody tissue (stem) photosynthesis for the carbon income in three California native plant species via measurements of chlorophyll concentrations, radial stem growth, bud biomass and stable carbon isotope composition of sugars in different plant organs. Young plants of Prunus ilicifolia, Umbellularia californica and Arctostaphylos manzanita were measured and subjected to manipulations at two levels: trunk light exclusion (100 and 50%) and complete defoliation. We found that long-term light exclusion resulted in a reduction in chlorophyll concentration and radial growth, demonstrating that trunk assimilates contributed to trunk carbon income. In addition, bud biomass was lower in covered plants compared to uncovered plants. Excluding 100% of the ambient light from trunks on defoliated plants led to an enrichment in 13 C of trunk phloem sugars. We attributed this effect to a reduction in photosynthetic carbon isotope discrimination against 13 C that in turn resulted in an enrichment in 13 C of bud sugars. Taken together our results reveal that stem photosynthesis contributes to the total carbon income of all species including the buds in defoliated plants.

Daytime depressions in stem FCO2 correlate with the daily dynamics of turgor, as a measure of the water status in the living stem tissues: it is suggested that water status of tree stems is a potentially important determinant of stem FCO2, as it influences the rate of growth and maintenance processes in the living tissues of the stem.

A young potted oak (Quercus robur L.) tree was subjected to drought by interrupting the water supply for 9 days. The tree was placed in a growth chamber in which daily patterns of temperature and radiation were constant. The effects of drought on the water and carbon status of the stem were examined by measuring stem sap flow rate, stem water potential, stem diameter variations, stem CO(2) efflux rate (F(CO2)) and xylem CO(2) concentration ([CO(2)*]). Before and after the drought treatment, diurnal fluctuations in F(CO2) and [CO(2)*] corresponded well with variations in stem temperature (T(st)). Daytime depressions in F(CO2) did not occur. During the drought treatment, F(CO2) still responded to stepwise changes in temperature, but diurnal fluctuations in F(CO2) were no longer correlated with diurnal fluctuations in T(st). From the moment daily growth rate of the stem became zero, diurnal fluctuations in F(CO2) became closely correlated with diameter variations, exhibiting clear daytime depressions. The depressions in F(CO2) were likely the result of a reduction in metabolic activity caused by the lowered daytime stem water status. Xylem [CO(2)*] showed clear daytime depressions in response to drought. When the tree was re-watered, F(CO2) and [CO(2)*] exhibited sharp increases, coinciding with an increase in stem diameter. After resumption of the water supply, daytime depressions in F(CO2) and [CO(2)*] disappeared and diurnal fluctuations in F(CO2) and [CO(2)*] corresponded again with variations in T(st).

The spatial and diurnal variability of sap flow in a mature beech tree (Fagus sylvatica L.) was investigated on days with different climatic conditions (sunny, cloudy, rainy), during the summer. Sap flux density (nu) was measured with six heat field deformation probes placed around the stem circumference. Each probe measured nu at six sapwood depths. Daily nu exhibited clear radial variation, and the shape of the radial profile of nu differed substantially among circumferential positions. At some positions, daily nu decreased monotonically towards the stem centre, whereas at others it showed an almost monotonic increase. Hence, the conducting sapwood area of the beech was highly asymmetrical. At all positions., conducting sapwood reached beyond the deepest sampled sapwood depth, precluding correct estimations of total sap flow. The radial profile of nu also differed among measuring days. A general trend was that the inner sapwood contributed relatively more to total sap flow under better weather conditions. Besides variations among days, the shape of the radial profile of nu also showed within-day variations. The contribution of the inner sapwood to total sap flow increased in the afternoon, with increasing vapour pressure deficit and photosynthetic active radiation. Because of large circumferential and temporal variability, no general function for the radial profile of nu could be developed

Respiration rates are reported to increase exponentially with temperature. Respiration rates of woody tissues are commonly measured as CO 2 efflux rates (F CO 2 ) from that tissue. However, this paper describes clear variations in stem F CO 2 that were not related to temperature for the case of a young beech (Fagus sylvatica L.) and oak (Quercus robur L.) tree during the dormant season. The CO 2 concentration ([CO 2 ]) in the xylem of the beech tree showed similar temperature-independent variations. The trees were grown in a growth chamber in which radiation patterns and temperature were kept constant. F CO 2 was measured with an IRGA connected to cuvettes surrounding a stem segment. Xylem [CO 2 ] was measured in situ using a CO 2 microelectrode. Depressions in F CO 2 and [CO 2 ] occurred during the light period, despite equal temperatures in the light and dark period. Explanations found in literature for discrepancies in the exponential relationship between temperature and F CO 2 are the influence of (1) sap flow or (2) decreased cell water content. However, (1) the variations were observed in the dormant season, when no sap flow was observed yet, and (2) reduced cell water content was not likely to be apparent as differences in stem transpiration rates between the dark and light period were not significant. Hence, previously formulated theories failed to explain our results. This work therefore provides a new ground for discussion on other possible causes of daytime depressions in F CO 2 . One might be the refixation of respired CO 2 by corticular photosynthesis in the stem parts adjacent to the stem segment enclosed by the cuvette.