Xylem transport of root-derived CO2 caused a substantial underestimation of belowground respiration during a growing season.

Previous research has indicated that a potentially large portion of root-respired CO2 can move internally through tree xylem, but these reports are relatively scarce and have generally been limited to short observations. Our main objective was to provide a continuous estimate of the quantity and variability of root-respired CO2 that moves either internally through the xylem (FT ) or externally through the soil to the atmosphere (FS ) over most of a growing season. Nine trees were measured in a Populus deltoides stand for 129 days from early June to mid-October. We calculated FT as the product of sap flow and dissolved [CO2 ] in the xylem (i.e., [CO2 *]) and calculated FS using the [CO2 ] gradient method. During the study, stem and soil CO2 concentrations, temperature, and sap flow were measured continuously. We determined that FT accounted for 33% of daily total belowground CO2 flux (i.e., FS  + FT ; FB ) during our observation period that spanned most of a growing season. Cumulative daily FT was lower than FS 74% of the time, equivalent to FS 26% of the time, and never exceeded FS . One-third of the total CO2 released by belowground respiration over most of the growing season in this forest stand followed the FT pathway rather than diffusing into the soil. The magnitude of FT indicates that measurements of FS alone substantially underestimate total belowground respiration in some forest ecosystems by systematically underestimating belowground autotrophic respiration. The variability in FT observed during the growing season demonstrated the importance of making long-term, high-frequency measurements of different flux pathways to better understand physiological and ecological processes and their implications to global change.

Stored root carbohydrates can maintain root respiration for extended periods.

Tight coupling between below-ground autotrophic respiration and the availability of recently assimilated carbon (C) has become a paradigm in the ecophysiological literature. Here, we show that stored carbohydrates can decouple respiration from assimilation for prolonged periods by mobilizing reserves from transport roots to absorptive roots. We permanently disrupted the below-ground transfer of recently assimilated C using stem girdling and root trenching and measured soil CO2 efflux for over 1 yr in longleaf pine (Pinus palustris), a species that has large reserves of stored carbohydrates in roots. Soil CO2 efflux was not influenced by girdling or trenching through the 14-month observation period. Stored carbohydrate concentrations in absorptive roots were not affected by the disrupted supply of current photosynthate for over 1 yr; however, carbohydrate concentrations in transport roots decreased. Our results indicate that root respiration can be decoupled from recent canopy assimilation and that stored carbohydrates can be mobilized from transport roots to absorptive roots to maintain respiration for over 1 yr. This refines the current paradigm that canopy assimilation and below-ground respiration are tightly coupled and provides evidence of the mechanism and dynamics responsible for decoupling the above- and below-ground processes.

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.

Pinus taeda forest growth predictions in the 21st century vary with site mean annual temperature and site quality.

Climate projections from 20 downscaled global climate models (GCMs) were used with the 3-PG model to predict the future productivity and water use of planted loblolly pine (Pinus taeda) growing across the southeastern United States. Predictions were made using Representative Concentration Pathways (RCP) 4.5 and 8.5. These represent scenarios in which total radiative forcing stabilizes before 2100 (RCP 4.5) or continues increasing throughout the century (RCP 8.5). Thirty-six sites evenly distributed across the native range of the species were used in the analysis. These sites represent a range in current mean annual temperature (14.9-21.6°C) and precipitation (1,120-1,680 mm/year). The site index of each site, which is a measure of growth potential, was varied to represent different levels of management. The 3-PG model predicted that aboveground biomass growth and net primary productivity will increase by 10%-40% in many parts of the region in the future. At cooler sites, the relative growth increase was greater than at warmer sites. By running the model with the baseline [CO2 ] or the anticipated elevated [CO2 ], the effect of CO2 on growth was separated from that of other climate factors. The growth increase at warmer sites was due almost entirely to elevated [CO2 ]. The growth increase at cooler sites was due to a combination of elevated [CO2 ] and increased air temperature. Low site index stands had a greater relative increase in growth under the climate change scenarios than those with a high site index. Water use increased in proportion to increases in leaf area and productivity but precipitation was still adequate, based on the downscaled GCM climate projections. We conclude that an increase in productivity can be expected for a large majority of the planted loblolly pine stands in the southeastern United States during this century.

Tolerance or avoidance: drought frequency determines the response of an N2 -fixing tree.

Climate change is increasing drought frequency, which may affect symbiotic N2 fixation (SNF), a process that facilitates ecosystem recovery from disturbance. Here, we assessed the effect of drought frequency on the ecophysiology and SNF rate of a common N2 -fixing tree in eastern US forests. We grew Robinia pseudoacacia seedlings under the same mean soil moisture, but with different drought frequency caused by wet-dry cycles of varying periodicity. We found no effect of drought frequency on final biomass or mean SNF rate. However, seedlings responded differently to wet and dry phases depending on drought frequency. Under low-frequency droughts, plants fixed carbon (C) and nitrogen (N) at similar rates during wet and dry phases. Conversely, under high-frequency droughts, plants fixed C and N at low rates during dry phases and at high rates during wet phases. Our findings suggest that R. pseudoacacia growth is resistant to increased drought frequency because it employs two strategies - drought tolerance or drought avoidance, followed by compensation. SNF may play a role in both by supplying N to leaf tissues for acclimation and by facilitating compensatory growth following drought. Our findings point to SNF as a mechanism for plants and ecosystems to cope with drought.

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.

Global patterns and predictors of stem CO2 efflux in forest ecosystems.

Stem CO2 efflux (ES) plays an important role in the carbon balance of forest ecosystems. However, its primary controls at the global scale are poorly understood and observation-based global estimates are lacking. We synthesized data from 121 published studies across global forest ecosystems and examined the relationships between annual ES and biotic and abiotic factors at individual, biome, and global scales, and developed a global gridded estimate of annual ES . We tested the following hypotheses: (1) Leaf area index (LAI) will be highly correlated with annual ES at biome and global scales; (2) there will be parallel patterns in stem and root CO2 effluxes (RA) in all forests; (3) annual ES will decline with forest age; and (4) LAI coupled with mean annual temperature (MAT) and mean annual precipitation (MAP) will be sufficient to predict annual ES across forests in different regions. Positive linear relationships were found between ES and LAI, as well as gross primary production (GPP), net primary production (NPP), wood NPP, soil CO2 efflux (RS), and RA . Annual ES was correlated with RA in temperate forests after controlling for GPP and MAT, suggesting other additional factors contributed to the relationship. Annual ES tended to decrease with stand age. Leaf area index, MAT and MAP, predicted 74% of variation in ES at global scales. Our statistical model estimated a global annual ES of 6.7 ± 1.1 Pg C yr(-1) over the period of 2000-2012 with little interannual variability. Modeled mean annual ES was 71 ± 43, 270 ± 103, and 420 ± 134 g C m(2) yr(-1) for boreal, temperate, and tropical forests, respectively. We recommend that future studies report ES at a standardized constant temperature, incorporate more manipulative treatments, such as fertilization and drought, and whenever possible, simultaneously measure both aboveground and belowground CO2 fluxes.

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.

Root-derived CO(2) efflux via xylem stream rivals soil CO(2) efflux.

Respiration consumes a large portion of annual gross primary productivity in forest ecosystems and is dominated by belowground metabolism. Here, we present evidence of a previously unaccounted for internal CO(2) flux of large magnitude from tree roots through stems. If this pattern is shown to persist over time and in other forests, it suggests that belowground respiration has been grossly underestimated. Using an experimental Populus deltoides plantation as a model system, we tested the hypothesis that a substantial portion of the CO(2) released from belowground autotrophic respiration remains within tree root systems and is transported aboveground through the xylem stream rather than diffusing into the soil atmosphere. On a daily basis, the amount of CO(2) that moved upward from the root system into the stem via the xylem stream (0.26 mol CO(2) m(-2) d(-1)) rivalled that which diffused from the soil surface to the atmosphere (0.27 mol CO(2) m(-2) d(-1)). We estimated that twice the amount of CO(2) derived from belowground autotrophic respiration entered the xylem stream as diffused into the soil environment. Our observations indicate that belowground autotrophic respiration consumes substantially more carbohydrates than previously recognized and challenge the paradigm that all root-respired CO(2) diffuses into the soil atmosphere.