Effects of stem size on stem respiration and its flux components in yellow-poplar (Liriodendron tulipifera L.) trees.

Carbon dioxide (CO2) released from respiring cells in the stems of trees (RS) can diffuse radially to the atmosphere (EA) or dissolve in xylem sap and move internally in the tree (FT). Previous studies have observed that EA decreases as stem or branch diameter increases, but the cause of this relationship has not been determined, nor has the relationship been confirmed between stem diameter and RS, which includes both EA and FT. In this study, for the first time the mass balance technique was used to estimate RS of stems of Liriodendron tulipifera L. trees of different diameters, ranging from 16 to 60 cm, growing on the same site. The magnitude of the component fluxes scaled with tree size. Among the five trees, the contribution of EA to RS decreased linearly with increasing stem diameter and sapwood area while the contribution of FT to RS increased linearly with stem diameter and sapwood area. For the smallest tree EA was 86% of RS but it was only 46% of RS in the largest tree. As tree size increased a greater proportion of respired CO2 dissolved in sap and remained within the tree. Due to increase in FT with tree size, we observed that trees of different sizes had the same RS even though they had different EA. This appears to explain why the EA of stems and branches decreases as their size increases.

Stomatal conductance increases with rising temperature.

Stomatal conductance directly modifies plant water relations and photosynthesis. Many environmental factors affecting the stomatal conductance have been intensively studied but temperature has been largely neglected, even though it is one of the fastest changing environmental variables and it is rising due to climate change. In this study, we describe how stomata open when the temperature increases. Stomatal conductance increased by ca 40% in a broadleaf and a coniferous species, poplar (Populus deltoides x nigra) and loblolly pine (Pinus taeda) when temperature was increased by 10 °C, from 30 °C to 40 °C at a constant vapor pressure deficit of 1 kPa. The mechanism of regulating stomatal conductance by temperature was, at least partly, independent of other known mechanisms linked to water status and carbon metabolism. Stomatal conductance increased with rising temperature despite the decrease in leaf water potential, increase in transpiration, increase in intercellular CO2 concentration and was decoupled from photosynthesis. Increase in xylem and mesophyll hydraulic conductance coming from lower water viscosity may to some degree explain temperature dependent opening of stomata. The direct stomatal response to temperature allows plants to benefit from increased evaporative cooling during the heat waves and from lower stomatal limitations to photosynthesis but they may be jeopardized by faster depletion of soil water.

Increase in leaf temperature opens stomata and decouples net photosynthesis from stomatal conductance in Pinus taeda and Populus deltoides x nigra.

The effect of temperature on stomatal conductance (gs) and corresponding gas exchange parameters was studied in two tree species with contrasting leaf anatomy and ecophysiology-a broadleaf angiosperm, Populus deltoides x nigra (poplar), and a needle-leaf gymnosperm, Pinus taeda (loblolly pine). Experiments were conducted in growth chambers across a leaf temperature range of 19-48°C. Manipulations of temperature were done in well-watered and drought soil conditions and under ambient (400 ppm) and elevated (800 ppm) air CO2 concentrations. Increases in leaf temperature caused stomatal opening at both ambient and elevated [CO2]. The gs increased by 42% in poplar and by 40% in loblolly pine when leaf temperature increased from 30°C to 40°C at a vapour pressure difference of 1 kPa. Stomatal limitation to photosynthesis decreased in elevated temperature in loblolly pine but not in poplar. The ratio of net photosynthesis to gs depended on leaf temperature, especially at high temperatures. Evaporative cooling of transpiring leaves resulted in reductions in leaf temperature up to 9°C in well-watered poplar but only 1°C in drought-stressed poplar and in loblolly pine. As global mean temperatures rise and temperature extremes become more frequent and severe, understanding the effect of temperature on gs, and modelling that relationship, will become increasingly important.

Cell Wall Ultrastructure of Stem Wood, Roots, and Needles of a Conifer Varies in Response to Moisture Availability.

The composition, integrity, and architecture of the macromolecular matrix of cell walls, collectively referred to as cell wall ultrastructure, exhibits variation across species and organs and among cell types within organs. Indirect approaches have suggested that modifications to cell wall ultrastructure occur in response to abiotic stress; however, modifications have not been directly observed. Glycome profiling was used to study cell wall ultrastructure by examining variation in composition and extractability of non-cellulosic glycans in cell walls of stem wood, roots, and needles of loblolly pine saplings exposed to high and low soil moisture. Soil moisture influenced physiological processes and the overall composition and extractability of cell wall components differed as a function of soil moisture treatments. The strongest response of cell wall ultrastructure to soil moisture was increased extractability of pectic backbone epitopes in the low soil moisture treatment. The higher abundance of these pectic backbone epitopes in the oxalate extract indicate that the loosening of cell wall pectic components could be associated with the release of pectic signals as a stress response. The increased extractability of pectic backbone epitopes in response to low soil moisture availability was more pronounced in stem wood than in roots or needles. Additional responses to low soil moisture availability were observed in lignin-associated carbohydrates released in chlorite extracts of stem wood, including an increased abundance of pectic arabinogalactan epitopes. Overall, these results indicate that cell walls of loblolly pine organs undergo changes in their ultrastructural composition and extractability as a response to soil moisture availability and that cell walls of the stem wood are more responsive to low soil moisture availability compared to cell walls of roots and needles. To our knowledge, this is the first direct evidence, delineated by glycomic analyses, that abiotic stress affects cell wall ultrastructure. This study is also unique in that glycome profiling of pine needles has never before been reported.

Temporal and spatial patterns of internal and external stem CO2 fluxes in a sub-Mediterranean oak.

To accurately estimate stem respiration (RS), measurements of both carbon dioxide (CO2) efflux to the atmosphere (EA) and internal CO2 flux through xylem (FT) are needed because xylem sap transports respired CO2 upward. However, reports of seasonal dynamics of FT and EA are scarce and no studies exist in Mediterranean species under drought stress conditions. Internal and external CO2 fluxes at three stem heights, together with radial stem growth, temperature, sap flow and shoot water potential, were measured in Quercus pyrenaica Willd. in four measurement campaigns during one growing season. Substantial daytime depressions in temperature-normalized EA were observed throughout the experiment, including prior to budburst, indicating that diel hysteresis between stem temperature and EA cannot be uniquely ascribed to diversion of CO2 in the transpiration stream. Low internal [CO2] (<0.5%) resulted in low contributions of FT to RS throughout the growing season, and RS was mainly explained by EA (>90%). Internal [CO2] was found to vary vertically along the stems. Seasonality in resistance to radial CO2 diffusion was related to shoot water potential. The low internal [CO2] and FT observed in our study may result from the downregulation of xylem respiration in response to a legacy of coppicing as well as high radial diffusion of CO2 through cambium, phloem and bark tissues, which was related to low water content of stems. Long-term studies analyzing temporal and spatial variation in internal and external CO2 fluxes and their interactions are needed to mechanistically understand and model respiration of woody tissues.

Seasonal and diel variation in xylem CO2 concentration and sap pH in sub-Mediterranean oak stems.

Since a substantial portion of respired CO2 remains within the stem, diel and seasonal trends in stem CO2 concentration ([CO2]) are of major interest in plant respiration and carbon budget research. However, continuous long-term stem [CO2] studies are scarce, and generally absent in Mediterranean climates. In this study, stem [CO2] was monitored every 15min together with stem and air temperature, sap flow, and soil water storage during a growing season in 16 stems of Quercus pyrenaica to elucidate the main drivers of stem [CO2] at different temporal scales. Fluctuations in sap pH were also assessed during two growing seasons to evaluate potential errors in estimates of the concentration of CO2 dissolved in xylem sap ([CO2*]) calculated using Henry's law. Stem temperature was the best predictor of stem [CO2] and explained more than 90% and 50% of the variability in stem [CO2] at diel and seasonal scales, respectively. Under dry conditions, soil water storage was the main driver of stem [CO2]. Likewise, the first rains after summer drought caused intense stem [CO2] pulses, suggesting enhanced stem and root respiration and increased resistance to radial CO2 diffusion. Sap flow played a secondary role in controlling stem [CO2] variations. We observed night-time sap pH acidification and progressive seasonal alkalinization. Thus, if the annual mean value of sap pH (measured at midday) was assumed to be constant, night-time sap [CO2*] was substantially overestimated (40%), and spring and autumn sap [CO2*] were misestimated by 25%. This work highlights that diel and seasonal variations in temperature, tree water availability, and sap pH substantially affect xylem [CO2] and sap [CO2*].

Responses of tree species to heat waves and extreme heat events.

The number and intensity of heat waves has increased, and this trend is likely to continue throughout the 21st century. Often, heat waves are accompanied by drought conditions. It is projected that the global land area experiencing heat waves will double by 2020, and quadruple by 2040. Extreme heat events can impact a wide variety of tree functions. At the leaf level, photosynthesis is reduced, photooxidative stress increases, leaves abscise and the growth rate of remaining leaves decreases. In some species, stomatal conductance increases at high temperatures, which may be a mechanism for leaf cooling. At the whole plant level, heat stress can decrease growth and shift biomass allocation. When drought stress accompanies heat waves, the negative effects of heat stress are exacerbated and can lead to tree mortality. However, some species exhibit remarkable tolerance to thermal stress. Responses include changes that minimize stress on photosynthesis and reductions in dark respiration. Although there have been few studies to date, there is evidence of within-species genetic variation in thermal tolerance, which could be important to exploit in production forestry systems. Understanding the mechanisms of differing tree responses to extreme temperature events may be critically important for understanding how tree species will be affected by climate change.

Poplar saplings exposed to recurring temperature shifts of different amplitude exhibit differences in leaf gas exchange and growth despite equal mean temperature.

Most investigations of plant responses to changes in temperature have focused on a constant increase in mean day/night temperature without considering how differences in temperature cycles can affect physiological processes and growth. To test the effects of changes in growth temperature on foliar carbon balance and plant growth, we repeatedly exposed poplar saplings (Populus deltoides × nigra) to temperature cycles consisting of 5 days of a moderate (M, +5 °C) or extreme (E, +10 °C) increase in temperature followed by 5 days of a moderate (M, -5 °C) or extreme (E, -10 °C) decrease in temperature, with respect to a control treatment (C, 23.4 °C). The temperature treatments had the same mean temperature over each warm and cool cycle and over the entire study. Our goal was to examine the influence of recurring temperature shifts on growth. Net photosynthesis (A) was relatively insensitive to changes in growth temperature (from 20 to 35 °C), suggesting a broad range of optimum temperature for photosynthesis. Leaf respiration (R) exhibited substantial acclimation to temperature, having nearly the same rate at 13 °C as at 33 °C. There was no evidence that preconditioning through temperature cycles affected the response of A or R to treatment temperature fluctuations. Averaged across the complete warm/cool temperature cycle, the A : R ratio did not differ among the temperature treatments. While foliar carbon balance was not affected, the temperature treatments significantly affected growth. Whole-plant biomass was 1.5 times greater in the M treatment relative to the C treatment. Carbon allocation was also affected with shoot volume and biomass greater in the M and E treatments than in the C treatment. Our findings indicate that temperature fluctuations can have important effects on growth, though there were few effects on leaf gas exchange, and can help explain differences in growth that are not correlated with mean growth temperature.

Stem girdling affects the quantity of CO2 transported in xylem as well as CO2 efflux from soil.

There is recent clear evidence that an important fraction of root-respired CO2 is transported upward in the transpiration stream in tree stems rather than fluxing to the soil. In this study, we aimed to quantify the contribution of root-respired CO2 to both soil CO2 efflux and xylem CO2 transport by manipulating the autotrophic component of belowground respiration. We compared soil CO2 efflux and the flux of root-respired CO2 transported in the transpiration stream in girdled and nongirdled 9-yr-old oak trees (Quercus robur) to assess the impact of a change in the autotrophic component of belowground respiration on both CO2 fluxes. Stem girdling decreased xylem CO2 concentration, indicating that belowground respiration contributes to the aboveground transport of internal CO2 . Girdling also decreased soil CO2 efflux. These results confirmed that root respiration contributes to xylem CO2 transport and that failure to account for this flux results in inaccurate estimates of belowground respiration when efflux-based methods are used. This research adds to the growing body of evidence that efflux-based measurements of belowground respiration underestimate autotrophic contributions.

Assimilation of xylem-transported CO2 is dependent on transpiration rate but is small relative to atmospheric fixation.

The effect of transpiration rate on internal assimilation of CO2 released from respiring cells has not previously been quantified. In this study, detached branches of Populus deltoides were allowed to take up (13)CO2-labelled solution at either high (high label, HL) or low (low label, LL) (13)CO2 concentrations. The uptake of the (13)CO2 label served as a proxy for the internal transport of respired CO2, whilst the transpiration rate was manipulated at the leaf level by altering the vapour pressure deficit (VPD) of the air. Simultaneously, leaf gas exchange was measured, allowing comparison of internal CO2 assimilation with that assimilated from the atmosphere. Subsequent (13)C analysis of branch and leaf tissues revealed that woody tissues assimilated more label under high VPD, corresponding to higher transpiration, than under low VPD. More (13)C was assimilated in leaf tissue than in woody tissue under the HL treatment, whereas more (13)C was assimilated in woody tissue than in leaf tissue under the LL treatment. The ratio of (13)CO2 assimilated from the internal source to CO2 assimilated from the atmosphere was highest for the branches under the HL and high VPD treatment, but was relatively small regardless of VPD×label treatment combination (up to 1.9%). These results showed that assimilation of internal CO2 is highly dependent on the rate of transpiration and xylem sap [CO2]. Therefore, it can be expected that the relative contribution of internal CO2 recycling to tree carbon gain is strongly dependent on factors controlling transpiration, respiration, and photosynthesis.

The effect of heat waves, elevated [CO2 ] and low soil water availability on northern red oak (Quercus rubra L.) seedlings.

The frequency and intensity of heat waves are predicted to increase. This study investigates whether heat waves would have the same impact as a constant increase in temperature with the same heat sum, and whether there would be any interactive effects of elevated [CO2 ] and soil moisture content. We grew Quercus rubra seedlings in treatment chambers maintained at either ambient or elevated [CO2 ] (380 or 700 µmol CO2 mol(-1) ) with temperature treatments of ambient, ambient +3 °C, moderate heat wave (+6 °C every other week) or severe heat wave (+12 °C every fourth week) temperatures. Averaged over a 4-week period, and the entire growing season, the three elevated temperature treatments had the same average temperature and heat sum. Half the seedlings were watered to a soil water content near field capacity, half to about 50% of this value. Foliar gas exchange measurements were performed morning and afternoon (9:00 and 15:00 hours) before, during and after an applied heat wave in August 2010. Biomass accumulation was measured after five heat wave cycles. Under ambient [CO2 ] and well-watered conditions, biomass accumulation was highest in the +3 °C treatment, intermediate in the +6 °C heat wave and lowest in the +12 °C heat wave treatment. This response was mitigated by elevated [CO2 ]. Low soil moisture significantly decreased net photosynthesis (Anet ) and biomass in all [CO2 ] and temperature treatments. The +12 °C heat wave reduced afternoon Anet by 23% in ambient [CO2 ]. Although this reduction was relatively greater under elevated [CO2 ], Anet values during this heat wave were still 34% higher than under ambient [CO2 ]. We concluded that heat waves affected biomass growth differently than the same amount of heat applied uniformly over the growing season, and that the plant response to heat waves also depends on [CO2 ] and soil moisture conditions.

Transport of root-respired CO2 via the transpiration stream affects aboveground carbon assimilation and CO2 efflux in trees.

Upward transport of CO2 via the transpiration stream from belowground to aboveground tissues occurs in tree stems. Despite potentially important implications for our understanding of plant physiology, the fate of internally transported CO2 derived from autotrophic respiratory processes remains unclear. We infused a ¹³CO2-labeled aqueous solution into the base of 7-yr-old field-grown eastern cottonwood (Populus deltoides) trees to investigate the effect of xylem-transported CO2 derived from the root system on aboveground carbon assimilation and CO2 efflux. The ¹³C label was transported internally and detected throughout the tree. Up to 17% of the infused label was assimilated, while the remainder diffused to the atmosphere via stem and branch efflux. The largest amount of assimilated ¹³C was found in branch woody tissues, while only a small quantity was assimilated in the foliage. Petioles were more highly enriched in ¹³C than other leaf tissues. Our results confirm a recycling pathway for respired CO2 and indicate that internal transport of CO2 from the root system may confound the interpretation of efflux-based estimates of woody tissue respiration and patterns of carbohydrate allocation.

Internal recycling of respired CO 2 may be important for plant functioning under changing climate regimes.

Recent studies have provided evidence of a large flux of root-respired CO 2 in the transpiration stream of trees. In our study, we investigated the potential impact of this internal CO 2 transport on aboveground carbon assimilation and CO 2 efflux. To trace the transport of root-respired CO 2, we infused a (13)C label at the stem base of field-grown Populus deltoides Bartr. ex. Marsh trees. The (13)C label was transported to the top of the stem and throughout the crown via the transpiration stream. Up to 17% of the (13)C label was assimilated by chlorophyll-containing tissues. Our results provide evidence of a mechanism for recycling respired CO 2 within trees. Such a mechanism may have important implications for how plants cope with predicted increases in intensity and frequency of droughts. Here, we speculate on the potential significance of this recycling mechanism within the context of plant responses to climate change and plants currently inhabiting arid environments.

The effect of induced heat waves on Pinus taeda and Quercus rubra seedlings in ambient and elevated CO2 atmospheres.

Here, we investigated the effect of different heat-wave intensities applied at two atmospheric CO2 concentrations ([CO2]) on seedlings of two tree species, loblolly pine (Pinus taeda) and northern red oak (Quercus rubra). Seedlings were assigned to treatment combinations of two levels of [CO2] (380 or 700 µmol mol(-1)) and four levels of air temperature (ambient, ambient +3°C, or 7-d heat waves consisting of a biweekly +6°C heat wave, or a monthly +12°C heat wave). Treatments were maintained throughout the growing season, thus receiving equal heat sums. We measured gas exchange and fluorescence parameters before, during and after a mid-summer heat wave. The +12°C heat wave, significantly reduced net photosynthesis (Anet) in both species and [CO2] treatments but this effect was diminished in elevated [CO2]. The decrease in Anet was accompanied by a decrease in Fv'/Fm' in P. taeda and ΦPSII in Q. rubra. Our findings suggest that, if soil moisture is adequate, trees will experience negative effects in photosynthetic performance only with the occurrence of extreme heat waves. As elevated [CO2] diminished these negative effects, the future climate may not be as detrimental to plant communities as previously assumed.

Effects of predicted future and current atmospheric temperature and [CO2] and high and low soil moisture on gas exchange and growth of Pinus taeda seedlings at cool and warm sites in the species range.

Predicted future changes in air temperature and atmospheric CO(2) concentration ([CO(2)]), coupled with altered precipitation, are expected to substantially affect tree growth. Effects on growth may vary considerably across a species range, as temperatures vary from sub-optimal to supra-optimal for growth. We performed an experiment simultaneously at two locations in the current range of loblolly pine, a cool site and a warm site, to examine the effect of future climate conditions on growth of loblolly pine seedlings in contrasting regions of the species range. At both sites 1-year-old loblolly pine seedlings were grown in current (local ambient temperature and [CO(2)]) and predicted future atmospheric conditions (ambient +2 °C temperature and 700 µmol mol(-1) [CO(2)]). Additionally, high and low soil moisture treatments were applied within each atmospheric treatment at each site by altering the amount of water provided to the seedlings. Averaged across water treatments, photosynthesis (A(net)) was 31% greater at the cool site and 34% greater at the warm site in elevated temperature and [CO(2)] compared with ambient temperature. Biomass accumulation was also stimulated by 38% at the cool site and by 24% at the warm site in that treatment. These results suggest that a temperature increase of 2 °C coupled with an increase in [CO(2)] (predicted future climate) will create conditions favorable for growth of this species. Reduced soil moisture decreased growth in both current and predicted atmospheric conditions. Biomass accumulation and A(net) were reduced by ∼39 and 17%, respectively, in the low water treatment. These results suggest that any benefit of future atmospheric conditions may be negated if soil moisture is reduced by altered precipitation patterns.

Higher growth temperatures decreased net carbon assimilation and biomass accumulation of northern red oak seedlings near the southern limit of the species range.

If an increase in temperature will limit the growth of a species, it will be in the warmest portion of the species distribution. Therefore, in this study we examined the effects of elevated temperature on net carbon assimilation and biomass production of northern red oak (Quercus rubra L.) seedlings grown near the southern limit of the species distribution. Seedlings were grown in chambers in elevated CO(2) (700 µmol mol(-1)) at three temperature conditions, ambient (tracking diurnal and seasonal variation in outdoor temperature), ambient +3 °C and ambient +6 °C, which produced mean growing season temperatures of 23, 26 and 29 °C, respectively. A group of seedlings was also grown in ambient [CO(2)] and ambient temperature as a check of the growth response to elevated [CO(2)]. Net photosynthesis and leaf respiration, photosynthetic capacity (V(cmax), J(max) and triose phosphate utilization (TPU)) and chlorophyll fluorescence, as well as seedling height, diameter and biomass, were measured during one growing season. Higher growth temperatures reduced net photosynthesis, increased respiration and reduced height, diameter and biomass production. Maximum net photosynthesis at saturating [CO(2)] and maximum rate of electron transport (J(max)) were lowest throughout the growing season in seedlings grown in the highest temperature regime. These parameters were also lower in June, but not in July or September, in seedlings grown at +3 °C above ambient, compared with those grown in ambient temperature, indicating no impairment of photosynthetic capacity with a moderate increase in air temperature. An unusual and potentially important observation was that foliar respiration did not acclimate to growth temperature, resulting in substantially higher leaf respiration at the higher growth temperatures. Lower net carbon assimilation was correlated with lower growth at higher temperatures. Total biomass at the end of the growing season decreased in direct proportion to the increase in growth temperature, declining by 6% per 1 °C increase in mean growing season temperature. Our observations suggest that increases in air temperature above current ambient conditions will be detrimental to Q. rubra seedlings growing near the southern limit of the species range.

Stem respiration and carbon dioxide efflux of young Populus deltoides trees in relation to temperature and xylem carbon dioxide concentration.

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.

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.

Resistance to radial CO2 diffusion contributes to between-tree variation in CO2 efflux of Populus deltoides stems.

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.

Assessing variation in the radial profile of sap flux density in Pinus species and its effect on daily water use.

We monitored sap flux density (v) diurnally in nine mature southeastern pine (Pinus spp.) trees with a thermal dissipation probe that spanned the sapwood radius. We found the expected pattern of high v near the cambium and decreasing v with depth toward the center of the tree; however, the pattern was not constant within a day or between trees. Radial profiles of trees were steeper earlier in the day and became less steep later in the day. As a result, time-dependent changes in the shape of the radial profile of v were sometimes correlated with daily changes in evaporative demand. As the radial profile became less steep, the inner xylem contributed relatively more to total tree sap flow than it did earlier in the day. We present a 3-parameter Gaussian function that can be used to describe the radial distribution of v in trees. Parameters in the function represent depth in the xylem from the cambium, maximum v, depth in the xylem where maximum v occurs, and the rate of radial change in v with radial depth (beta). Values of beta varied significantly between trees and with time, and were sometimes correlated with air vapor pressure deficit (D). We hypothesize that this occurred during periods of high transpiration when the water potential gradient became great enough to move water in the inner sapwood despite its probable high hydraulic resistance. We examined discrepancies among estimates of daily water use based on single-point, two-point and multi-point (i.e., every 20 mm in the sapwood) measurements. When radial distribution of v was not considered, a single-point measurement resulted in errors as large as 154% in the estimate of daily water use relative to the estimate obtained from a multi-point measurement. Measuring v at two close sample points (10 and 30 mm) did not improve the estimate; however, estimates derived from v measured at two distant sample points (10 and 70 mm) significantly improved the estimate of daily water use, although errors were as great as 32% in individual trees. The variability in v with depth in the xylem, over time, and between trees indicates that measurements of the radial distribution of v are necessary to accurately estimate water flow in trees with large sapwood areas.