Forest stand and canopy development unaltered by 12 years of CO2 enrichment.

Canopy structure-the size and distribution of tree crowns and the spatial and temporal distribution of leaves within them-exerts dominant control over primary productivity, transpiration and energy exchange. Stand structure-the spatial arrangement of trees in the forest (height, basal area and spacing)-has a strong influence on forest growth, allocation and resource use. Forest response to elevated atmospheric CO2 is likely to be dependent on the canopy and stand structure. Here, we investigated elevated CO2 effects on the forest structure of a Liquidambar styraciflua L. stand in a free-air CO2 enrichment experiment, considering leaves, tree crowns, forest canopy and stand structure. During the 12-year experiment, the trees increased in height by 5 m and basal area increased by 37%. Basal area distribution among trees shifted from a relatively narrow distribution to a much broader one, but there was little evidence of a CO2 effect on height growth or basal area distribution. The differentiation into crown classes over time led to an increase in the number of unproductive intermediate and suppressed trees and to a greater concentration of stand basal area in the largest trees. A whole-tree harvest at the end of the experiment permitted detailed analysis of canopy structure. There was little effect of CO2 enrichment on the relative leaf area distribution within tree crowns and there was little change from 1998 to 2009. Leaf characteristics (leaf mass per unit area and nitrogen content) varied with crown depth; any effects of elevated CO2 were much smaller than the variation within the crown and were consistent throughout the crown. In this young, even-aged, monoculture plantation forest, there was little evidence that elevated CO2 accelerated tree and stand development, and there were remarkably small changes in canopy structure. Questions remain as to whether a more diverse, mixed species forest would respond similarly.

Facultative symbiosis with a saprotrophic soil fungus promotes potassium uptake in American sweetgum trees.

Several species of soil free-living saprotrophs can sometimes establish biotrophic symbiosis with plants, but the basic biology of this association remains largely unknown. Here, we investigate the symbiotic interaction between a common soil saprotroph, Clitopilus hobsonii (Agaricomycetes), and the American sweetgum (Liquidambar styraciflua). The colonized root cortical cells were found to contain numerous microsclerotia-like structures. Fungal colonization led to increased plant growth and facilitated potassium uptake, particularly under potassium limitation (0.05 mM K+ ). The expression of plant genes related to potassium uptake was not altered by the symbiosis, but colonized roots contained the transcripts of three fungal genes with homology to K+ transporters (ACU and HAK) and channel (SKC). Heterologously expressed ChACU and ChSKC restored the growth of a yeast K+ -uptake-defective mutant. Upregulation of ChACU transcript under low K+ conditions (0 and 0.05 mM K+ ) compared to control (5 mM K+ ) was demonstrated in planta and in vitro. Colonized plants displayed a larger accumulation of soluble sugars under 0.05 mM K+ than non-colonized plants. The present study suggests reciprocal benefits of this novel tree-fungus symbiosis under potassium limitation mainly through an exchange of additional carbon and potassium between both partners.

Higher soil capacity of intercepting heavy rainfall in mixed stands than in pure stands in riparian forests.

Changes in global precipitation patterns would make wet regions more humid and extreme precipitation events occur frequently, followed by widespread flooding. Riparian forests are more capable of withstanding floods than inland forests because they are frequently exposed to short-term flooding events. Although many previous studies have investigated the soil water dynamics of terrestrial forests, little is known about how the soil water of riparian forests responds to different amounts of rainfall and which factors mainly regulate the soil water-holding capacity. Here, we employed stable hydrogen isotope to explore the contribution of different magnitudes of rainfall (7.9, 18.6 and 34.1 mm) to the soil water in two types of riparian forests (pure vs. mixed stand of Populus deltoides) in the middle-lower reaches of the Yangtze River, China. We further used structure equation modelling to determine the relative importance of soil properties and vegetation biomass in affecting the contribution of different magnitudes of rainfall to soil water. Our results revealed that there was no significant difference between these two stand types in the contributions of light and moderate rainfall to soil water, while the contribution of heavy rainfall to soil water (CHRSW) in mixed stand was significantly higher than that in pure stand (74.3% vs. 62.9%), suggesting that mixed stand soil has higher water-holding capacity than pure stand soil. Furthermore, soil properties were the best predictor affecting CHRSW, which explained 68% and 59% of the variation in the CHRSW on the 1st and 8th days after rainfall, respectively. Moreover, the root biomass could indirectly affect the CHRSW. Overall, mixed stand soil had a greater capacity in intercepting and storing rainwater than pure stand soil, implying that the mixed stand plantation, rather than the pure stand, should be recommended in riparian forest restoration projects that aim to improve their capacity for alleviating floods.

Expected Timber-Based Economic Impacts of a Wood-Boring Beetle (Acanthotomicus Sp.) That Kills American Sweetgum.

American sweetgum trees (Liquidambar styraciflua L. [Altingiaceae]) in China are being killed by a newly discovered wood-boring beetle "sweetgum inscriber" (Acanthotomicus sp.). It has not been detected in the United States yet, but given the extent of trade with Asian countries, eventual arrival of this beetle is a serious concern. The American sweetgum is one of the main hardwood species in the southern United States, and provides several economic and ecological benefits to society. We present the first economic analysis of the potential damage from sweetgum inscriber (SI) to timber-based land values in the southern United States. We modeled economic impacts for a range of feasible SI arrival rates that reflect policy interventions: 1) no efforts to prevent arrival (scenario A, once every 14 and 25 yr), 2) partial prevention by complying with ISPM 15 standards (scenario B, once every 33 and 100 yr), and 3) total prevention of arrival (scenario C, zero transmission of SI). Our results indicated much lower land values for sweetgum plantations without the prevention on SI establishment (scenario A, US$1,843-US$4,383 ha-1) compared with partial prevention (scenario B, US$5,426-US$8,050 ha-1) and total eradication of SI (scenario C, US$9,825). Across the region, upper bound timber-based economic losses to plantation owners is US$151.9 million (US$4.6 million annually)-an estimate that can help inform policy decisions.

Photosynthetic capacity of senescent leaves for a subtropical broadleaf deciduous tree species Liquidambar formosana Hance.

Photosynthetic capacity and leaf life span generally determine how much carbon a plant assimilates during the growing season. Leaves of deciduous tree species start senescence in late season, but whether the senescent leaves still retain capacity of carbon assimilation remains a question. In this study, we investigated leaf phenology and photosynthesis of a subtropical broadleaf deciduous tree species Liquidambar formosana Hance in the central southern continental China. The results show that L. formosana has extended leaf senescence (more than 2 months) with a substantial number of red leaves persisting on the tree. Leaf photosynthetic capacity decreases over season, but the senescent red leaves still maintain relatively high photosynthetic capacity at 42%, 66% and 66% of the mature leaves for net photosynthesis rate, apparent quantum yield, and quantum yield at the light compensation point, respectively. These results indicate that L. formosana may still contribute to carbon sink during leaf senescence.

Acanthotomicus sp. (Coleoptera: Curculionidae: Scolytinae), a New Destructive Insect Pest of North American Sweetgum Liquidambar styraciflua in China.

A previously unknown bark beetle species, Acanthotomicus sp., has emerged as a lethal pest of American sweetgum (Liquidambar styraciflua) in China. Our survey of nursery records from around Shanghai suggests that American sweetgum have been under heavy attack since at least 2013, resulting in the death of > 10,000 trees. Mass attacks of the apparently sweetgum-specific Acanthotomicus sp. can be diagnosed by accumulation of resinous exudates on the trunk, wilted foliage, and eventual numerous exit holes of the new generation. A Chinese native sweetgum Liquidambar formosana can also be colonized by Acanthotomicus sp. This pest is of concern not only as a killer of sweetgum in the Chinese nursery trade but also as a potentially destructive invasive pest of sweetgum in North America. This discovery suggests that global preinvasion assessment of pests is warranted.

Disentangling the visual cues used by a jumping spider to locate its microhabitat.

Many arthropod species have evolved to thrive only on the leaves of a particular species of plant, which they must be capable of finding in order to survive accidental displacement, developmental transitions or the changing of the seasons. A number of studies have tested whether such species select leaves to land or oviposit on based on their color, shape or size. Unfortunately, many studies did not control for correlates of these characters, such as the brightness of different colors, the areas of different shapes, and the level of ambient illumination in the vicinity of different sizes of leaves. In the present study, we tested for leaf color, shape and size preferences in a leaf-dwelling jumping spider (Lyssomanes viridis) with known summer and winter host plants, while controlling for these correlates. First, color preferences were tested outdoors under the natural illumination of their forest habitat. Lyssomanes viridis did not prefer to perch on a green substrate compared with various shades of gray, but did prefer the second darkest shade of gray we presented them with. Of the green and gray substrates, this shade of gray's integrated photon flux (350-700 nm), viewed from below, i.e. the spider's perspective in the arena, was the most similar to that of real leaves. This relationship also held when we weighted the transmitted photon flux by the jumping spiders' green photopigment spectral sensitivity. Spiders did not prefer the star-like leaf shape of their summer host plant, Liquidambar styraciflua, to a green circle of the same area. When given a choice between a L. styraciflua leaf-shaped stimulus that was half the area of an otherwise identical alternative, spiders preferred the larger stimulus. However, placing a neutral density filter over the side of the experimental arena with the smaller stimulus abolished this preference, with spiders then being more likely to choose the side of the arena with the smaller stimulus. In conclusion, L. viridis appears to use ambient illumination and possibly perceived leaf brightness but not leaf shape or color to locate its microhabitat. This calls for a careful re-examination of which visual cues a variety of arthropods are actually attending to when they search for their preferred host species or microhabitat.

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In this study, the photosynthetic light response curves were measured for Liquidambar formosana during the leaf senescence from October to December in 2014. The measurements were simulated by a photosynthetic light response model (Ye model) and the conventional non-rectangular hyperbola model, in order to understand the photosynthetic capacity of senescing leaves of L. formosana. The results showed that the light sensitivity of the net photosynthetic rate decreased gra-dually during the leaf senescence. The measured maximum net photosynthetic rate was about 2.88 µmol CO2·m-2·s-1 when the leaf color just turned yellow, and dropped to 0.95 µmol CO2·m-2·s-1 in the later stage of leaf senescence (8th December). The two photosynthetic light-response models performed well in fitting the observation data, with Ye model being slightly better. Parameters estimated from the two models, such as the maximum net photosynthetic rate, the appa-rent quantum yield, the quantum yield at the light compensation point and the dark respiration rate, all gradually decreased with time, quantitatively describing the decrease in the photosynthetic capacity during the leaf senescence for L. formosana. The senescing leaves of L. formosana maintained positive net photosynthesis rates during the whole senescence, which had positive impact on carbon assimilation in the study area.

Chemical and anatomical changes in Liquidambar styraciflua L. xylem after long term exposure to elevated CO2.

The anatomical and chemical characteristics of sweetgum were studied after 11 years of elevated CO2 (544 ppm, ambient at 391 ppm) exposure. Anatomically, branch xylem cells were larger for elevated CO2 trees, and the cell wall thickness was thinner. Chemically, elevated CO2 exposure did not impact the structural components of the stem wood, but non-structural components were significantly affected. Principal component analysis (PCA) was employed to detect differences between the CO2 treatments by considering numerous structural and chemical variables, as well as tree size, and data from previously published sources (i.e., root biomass, production and turnover). The PCA results indicated a clear separation between trees exposed to ambient and elevated CO2 conditions. Correlation loadings plots of the PCA revealed that stem structural components, ash, Ca, Mg, total phenolics, root biomass, production and turnover were the major responses that contribute to the separation between the elevated and ambient CO2 treated trees.

Transcriptome analysis of a subtropical deciduous tree: autumn leaf senescence gene expression profile of Formosan gum.

Autumn leaf senescence is a spectacular natural phenomenon; however, the regulation networks controlling autumnal colors and the leaf senescence program remain largely unelucidated. Whether regulation of leaf senescence is similar in subtropical deciduous plants and temperate deciduous plants is also unknown. In this study, the gene expression of a subtropical deciduous tree, Formosan gum (Liquidambar formosana Hance), was profiled. The transcriptomes of April leaves (green leaves, 'G') and December leaves (red leaves, 'R') were investigated by next-generation gene sequencing. Out of 58,402 de novo assembled contigs, 32,637 were annotated as putative genes. Furthermore, the L. formosana-specific microarray designed based on total contigs was used to extend the observation period throughout the growing seasons of 2011-2013. Network analysis from the gene expression profile focused on the genes up-regulated when autumn leaf senescence occurred. LfWRKY70, LfWRKY75, LfWRKY65, LfNAC1, LfSPL14, LfNAC100 and LfMYB113 were shown to be key regulators of leaf senescnece, and the genes regulated by LfWRKY75, LfNAC1 and LfMYB113 are candidates to link chlorophyll degradation and anthocyanin biosynthesis to senescence. In summary, the gene expression profiles over the entire year of the developing leaf from subtropical deciduous trees were used for in silico analysis and the putative gene regulation in autumn coloration and leaf senescence is discussed in this study.

Effects of artificial defoliation on growth and biomass accumulation in short-rotation sweetgum (Liquidambar styraciflua) in North Carolina.

Sweetgum, Liquidambar styraciflua L. (Hamamelidales: Hamamelidaceae), is a species of interest for short-rotation plantation forestry in the southeastern United States. Despite its high levels of resistance to many native insects and pathogens, the species is susceptible to generalist defoliators during outbreak epidemics. The objective of this field study was to evaluate the potential impact of defoliation on sweetgum growth and productivity within the context of an operational plantation. Over three growing seasons, trees were subjected to artificial defoliation treatments of various intensity (control = 0% defoliation; low intensity = 33% defoliation; moderate intensity = 67% defoliation; high intensity = 99% defoliation) and frequency (not defoliated; defoliated once in April of the first growing season; defoliated twice, once in April of the first growing season and again in April of the second growing season). The responses of stem height, stem diameter, stem volume, crown volume, total biomass accumulation, and branch growth were measured in November of each growing season. At the end of the first growing season, when trees had received single defoliations, significant reductions in all growth traits followed the most severe (99%) defoliation treatment only. After the second and third growing seasons, when trees had received one or two defoliations of varying intensity, stem diameter and volume and total tree biomass were reduced significantly by 67 and 99% defoliation, while reductions in stem height and crown volume followed the 99% treatment only. All growth traits other than crown volume were reduced significantly by two defoliations but not one defoliation. Results indicate that sweetgum is highly resilient to single defoliations of low, moderate, and high intensity. However, during the three-year period of the study, repeated high-intensity defoliation caused significant reductions in growth and productivity that could have lasting impacts on yield throughout a harvest rotation.

Stored carbon partly fuels fine-root respiration but is not used for production of new fine roots.

The relative use of new photosynthate compared to stored carbon (C) for the production and maintenance of fine roots, and the rate of C turnover in heterogeneous fine-root populations, are poorly understood. We followed the relaxation of a (13)C tracer in fine roots in a Liquidambar styraciflua plantation at the conclusion of a free-air CO(2) enrichment experiment. Goals included quantifying the relative fractions of new photosynthate vs stored C used in root growth and root respiration, as well as the turnover rate of fine-root C fixed during [CO(2)] fumigation. New fine-root growth was largely from recent photosynthate, while nearly one-quarter of respired C was from a storage pool. Changes in the isotopic composition of the fine-root population over two full growing seasons indicated heterogeneous C pools; < 10% of root C had a residence time < 3 months, while a majority of root C had a residence time > 2 yr. Compared to a one-pool model, a two-pool model for C turnover in fine roots (with 5 and 0.37 yr(-1) turnover times) doubles the fine-root contribution to forest NPP (9-13%) and supports the 50% root-to-soil transfer rate often used in models.

Photosynthetic and antioxidant responses of Liquidambar formosana and Schima superba seedlings to sulfuric-rich and nitric-rich simulated acid rain.

To study whether differential responses occur in photosynthesis and antioxidant system for seedlings of Liquidambar formosana, an acid rain (AR)-sensitive tree species and Schima superba, an AR-tolerant tree species treated with three types of pH 3.0 simulated AR (SiAR) including sulfuric-rich (S-SiAR), nitric-rich (N-SiAR), sulfate and nitrate mixed (SN-SiAR), we investigated the changes of leaf necrosis, chlorophyll content, soluble protein and proline content, photosynthesis and chlorophyll fluorescence characteristics, reactive oxygen species production, membrane lipid peroxidation, small molecular antioxidant content, antioxidant enzyme activities and related protein expressions. Our results showed that SiAR significantly caused leaf necrosis, inhibited photosynthesis, induced superoxide radical and hydrogen peroxide generation, aggravated membrane lipid peroxidation, changed antioxidant enzyme activities, modified related protein expressions such as Cu/Zn superoxide dismutase (SOD), l-ascorbate peroxidase (APX, EC 1. 11. 1. 11), glutathione S transferase (GST, EC 2. 5. 1. 18) and Rubisco large subunit (RuBISCO LSU), altered non-protein thiols (NPT) and glutathione (GSH) content in leaves of L. formosana and S. superba. Taken together, we concluded that the damages caused by SiAR in L. formosana were more severe and suffered from more negative impacts than in S. superba. S-SiAR induced more serious damages for the plants than did SN-SiAR and N-SiAR.

The effects of elevated CO2 and nitrogen fertilization on stomatal conductance estimated from 11 years of scaled sap flux measurements at Duke FACE.

In this study, we employ a network of thermal dissipation probes (TDPs) monitoring sap flux density to estimate leaf-specific transpiration (E(L)) and stomatal conductance (G(S)) in Pinus taeda (L.) and Liquidambar styraciflua L. exposed to +200 ppm atmospheric CO(2) levels (eCO(2)) and nitrogen fertilization. Scaling half-hourly measurements from hundreds of sensors over 11 years, we found that P. taeda in eCO(2) intermittently (49% of monthly values) decreased stomatal conductance (G(S)) relative to the control, with a mean reduction of 13% in both total E(L) and mean daytime G(S). This intermittent response was related to changes in a hydraulic allometry index (A(H)), defined as sapwood area per unit leaf area per unit canopy height, which decreased a mean of 15% with eCO(2) over the course of the study, due mostly to a mean 19% increase in leaf area (A(L)). In contrast, L. styraciflua showed a consistent (76% of monthly values) reduction in G(S) with eCO(2) with a total reduction of 32% E(L), 31% G(S) and 23% A(H) (due to increased A(L) per sapwood area). For L. styraciflua, like P. taeda, the relationship between A(H) and G(S) at reference conditions suggested a decrease in G(S) across the range of A(H). Our findings suggest an indirect structural effect of eCO(2) on G(S) in P. taeda and a direct leaf level effect in L. styraciflua. In the initial year of fertilization, P. taeda in both CO(2) treatments, as well as L. styraciflua in eCO(2), exhibited higher G(S) with N(F) than expected from shifts in A(H), suggesting a transient direct effect on G(S). Whether treatment effects on mean leaf-specific G(S) are direct or indirect, this paper highlights that long-term treatment effects on G(S) are generally reflected in A(H) as well.

Plasticity in bundle sheath extensions of heterobaric leaves.

PREMISE OF THE STUDY: Leaf venation is linked to physiological performance, playing a critical role in ecosystem function. Despite the importance of leaf venation, associated bundle sheath extensions (BSEs) remain largely unstudied. Here, we quantify plasticity in the spacing of BSEs over irradiance and precipitation gradients. Because physiological function(s) of BSEs remain uncertain, we additionally explored a link between BSEs and water use efficiency (WUE). METHODS: We sampled leaves of heterobaric trees along intracrown irradiance gradients in natural environments and growth chambers and correlated BSE spacing to incident irradiance. Additionally, we sampled leaves along a precipitation gradient and correlated BSE spacing to precipitation and bulk δ(13)C, a proxy for intrinsic WUE. BSE spacing was quantified using a novel semiautomatic method on fresh leaf tissue. KEY RESULTS: With increased irradiance or decreased precipitation, Liquidambar styraciflua decreased BSE spacing, while Acer saccharum showed little variation in BSE spacing. Two additional species, Quercus robur and Platanus occidentalis, decreased BSE spacing with increased irradiance in growth chambers. BSE spacing correlated with bulk δ(13)C, a proxy for WUE in L. styraciflua, Q. robur, and P. occidentalis leaves but not in leaves of A. saccharum. CONCLUSIONS: We demonstrated that BSE spacing is plastic with respect to irradiance or precipitation and independent from veins, indicating BSE involvement in leaf adaptation to a microenvironment. Plasticity in BSE spacing was correlated with WUE only in some species, not supporting a function in water relations. We discuss a possible link between BSE plasticity and life history, particularly canopy position.

Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species.

Juvenile leaves in high-light environments commonly appear red as a result of anthocyanin pigments, which play a photoprotective role during light-sensitive ontogenetic stages. The loss of anthocyanin during leaf development presumably corresponds to a decreased need for photoprotection, as photosynthetic maturation allows leaves to utilize higher light intensities. However, the relationship between photosynthetic development and anthocyanin decline has yet to be quantitatively described. In this study, anthocyanin concentration was measured against photopigment content, lamina thickness, anatomical development, and photosynthetic CO(2) exchange in developing leaves of three deciduous tree species. In all species, anthocyanin disappearance corresponded with development of c. 50% mature photopigment concentrations, c. 80% lamina thickness, and differentiation of the mesophyll into palisade and spongy layers. Photosynthetic gas exchange correlated positively with leaf thickness and chlorophyll content, and negatively with anthocyanin concentration. Species with more rapid photosynthetic maturation lost anthocyanin earliest in development. Chlorophyll a/b ratios increased with leaf age, and were lower than those of acyanic species, consistent with a shading effect of anthocyanin. These results suggest that anthocyanin reassimilation is linked closely with chloroplast and whole-leaf developmental processes, supporting the idea that anthocyanins protect tissues until light processing and carbon fixation have matured to balance energy capture with utilization.

Seed dispersal in heterogeneous environments: bridging the gap between mechanistic dispersal and forest dynamics models.

Seed dispersal is an important determinant of vegetation composition. We present a mechanistic model of seed dispersal by wind that incorporates heterogeneous vegetation structure. Vegetation affects wind speeds, a primary determinant of dispersal distance. Existing models combine wind speed and fall velocity of seeds. We expand on them by allowing vegetation, and thus wind profiles, to vary along seed trajectories, making the model applicable to any wind-dispersed plant in any community. Using seed trap data on seeds dispersing from forests into adjacent sites of two distinct vegetation structures, we show that our model was unbiased and accurate, even though dispersal patterns differed greatly between the two structures. Our spatially heterogeneous model performed better than models that assumed homogeneous vegetation for the same system. Its sensitivity to vegetation structure and ability to predict seed arrival when vegetation structure was incorporated demonstrates the model's utility for providing realistic estimates of seed arrival in realistic landscapes. Thus, we begin to bridge mechanistic seed dispersal and forest dynamics models. We discuss the merits of our model for incorporation into forest simulators, applications where such incorporation has been or is likely to be especially fruitful, and future model refinements to increase understanding of seed dispersal by wind.

Radiation-use efficiency and gas exchange responses to water and nutrient availability in irrigated and fertilized stands of sweetgum and sycamore.

We investigated how water and nutrient availability affect radiation-use efficiency (epsilon) and assessed leaf gas exchange as a possible mechanism for shifts in epsilon. We measured aboveground net primary production (ANPP) and annual photosynthetically active radiation (PAR) capture to calculate epsilon as well as leaf-level physiological variables (light-saturated net photosynthesis, Asat; stomatal conductance, gs; leaf internal CO2 concentration, Ci; foliar nitrogen concentration, foliar [N]; and midday leaf water potential, Psileaf) during the second (2001) and third (2002) growing seasons in sweetgum (Liquidambar styraciflua L.) and sycamore (Platanus occidentalis L.) stands receiving a factorial combination of irrigation and fertilization at the Savannah River Site, South Carolina. Irrigation and fertilization increased PAR capture (maximum increase 60%) in 2001 and 2002 for both species and annual PAR capture was well correlated with ANPP (mean r2 = 0.77). A decreasing trend in epsilon was observed in non-irrigated stands for sweetgum in 2001 and for sycamore in both years, although this was only significant for sycamore in 2002. Irrigated stands maintained higher gas exchange rates than non-irrigated stands for sweetgum in 2001 and for sycamore in both years, although foliar [N] and Psileaf were generally unaffected. Because Ci decreased in proportion to gs in non-irrigated stands, it appeared that greater stomatal limitation of photosynthesis was associated with decreased Asat. On several measurement dates for sweetgum in 2001 and for sycamore in both years, epsilon was positively correlated with gas exchange variables (Asat, gs, Ci) (r ranged from 0.600 to 0.857). These results indicate that PAR capture is well correlated with ANPP and that gas exchange rates modified by irrigation can influence the conversion of captured light energy to biomass.

Estimating stem respiration in trees by a mass balance approach that accounts for internal and external fluxes of CO2.

The respiration rate of a tree stem has commonly been estimated from measurements of CO2 efflux to the atmosphere. These estimates assume that all CO2 efflux originates from respiration of local tissues and that all CO2 produced by local tissues escapes to the atmosphere through the bark. However, dissolved CO2 can be transported in the xylem stream, and CO2 concentration ([CO2]) in xylem can be up to three orders of magnitude greater than that of the atmosphere, suggesting that measurements of CO2 efflux do not account for all CO2 produced by respiration. Here, we propose a new mass balance approach for estimating the respiration rate of tree stems that accounts for both external and internal fluxes of CO2. We demonstrate this approach using measurements of CO2 efflux, sap flux and internal [CO(2)] to calculate the rate of CO2 production of a segment of stem tissue in situ. At different times of the day, CO2 produced by respiration of stem tissues followed different flux pathways. During daylight hours when sap was flowing, a large proportion of respired CO2 was carried away in the xylem stream, whereas at night, most respiratory CO2 escaped to the atmosphere through the bark. Our calculations showed errors in efflux-based estimates of respiration of up to 76% compared with estimates that include both internal and external fluxes.