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Carbon dioxide sequestration from the atmosphere is commonly assessed using the eddy covariance method. Its net flux signal can be decomposed into gross primary production and ecosystem respiration components, but these have seldom been tested against independent methods. In addition, eddy covariance lacks the ability to partition carbon sequestration among individual trees or species within mixed forests. Therefore, we compared gross primary production from eddy covariance versus an independent method based on sap flow and water-use efficiency, as measured by the tissue heat balance method and δ13C of phloem contents, respectively. The latter measurements were conducted on individual trees throughout a growing season in a mixed broadleaf forest dominated by three tree species, namely English oak, narrow-leaved ash and common hornbeam (Quercus robur L., Fraxinus angustifolia Vahl, and Carpinus betulus L., respectively). In this context, we applied an alternative ecophysiological method aimed at verifying the accuracy of a state-of-the-art eddy covariance system while also offering a solution to the partitioning problem. We observed strong agreement in the ecosystem gross primary production estimates (R2 = 0.56; P < 0.0001), with correlation being especially high and nearly on the 1:1 line in the period before the end of July (R2 = 0.85; P < 0.0001). After this period, the estimates of gross primary production began to diverge. Possible reasons for the divergence are discussed, focusing especially on phenology and the limitation of the isotopic data. English oak showed the highest per-tree daily photosynthetic rates among tree species, but the smaller, more abundant common hornbeam contributed most to the stand-level summation, especially early in the spring. These findings provide a rigorous test of the methods and the species-level photosynthesis offers avenues for enhancing forest management aimed at carbon sequestration., (© The Author(s) 2024. Published by Oxford University Press.)

Large trees in plantations generally produce more wood per unit of resource use than small trees. Two processes may account for this pattern: greater photosynthetic resource use efficiency or greater partitioning of carbon to wood production. We estimated gross primary production (GPP) at the individual scale by combining transpiration with photosynthetic water-use efficiency of Eucalyptus trees. Aboveground production fluxes were estimated using allometric equations and modeled respiration; total belowground carbon fluxes (TBCF) were estimated by subtracting aboveground fluxes from GPP. Partitioning was estimated by dividing component fluxes by GPP. Dominant trees produced almost three times as much wood as suppressed trees. They used 25 ± 10% (mean ± SD) of their photosynthates for wood production, whereas suppressed trees only used 12 ± 2%. By contrast, dominant trees used 27 ± 19% of their photosynthate belowground, whereas suppressed trees used 58 ± 5%. Intermediate trees lay between these extremes. Photosynthetic water-use efficiency of dominant trees was c. 13% greater than the efficiency of suppressed trees. Suppressed trees used more than twice as much of their photosynthate belowground and less than half as much aboveground compared with dominant trees. Differences in carbon partitioning were much greater than differences in GPP or photosynthetic water-use efficiency., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

Rationale: New methods to measure stable isotopes of soil and tree water directly in the field enable us to increase the temporal resolution of obtained data and advance our knowledge on the dynamics of soil and plant water fluxes. Only few field applications exist. However, these are needed to further improve novel methods and hence exploit their full potential., Methods: We tested the borehole equilibration method in the field and collected in situ and destructive samples of stable isotopes of soil, trunk and root xylem water over a 2.5-month experiment in a tropical dry forest under natural abundance conditions and following labelled irrigation. Water from destructive samples was extracted using cryogenic vacuum extraction. Isotope ratios were determined with IRIS instruments using cavity ring-down spectroscopy both in the field and in the laboratory., Results: In general, timelines of both methods agreed well for both soil and xylem samples. Irrigation labelled with heavy hydrogen isotopes clearly impacted the isotope composition of soil water and one of the two studied tree species. Inter-method deviations increased in consequence of labelling, which revealed their different capabilities to cover spatial and temporal heterogeneities., Conclusions: We applied the novel borehole equilibration method in a remote field location. Our experiment reinforced the potential of this in situ method for measuring xylem water isotopes in both tree trunks and roots and confirmed the reliability of gas permeable soil probes. However, in situ xylem measurements should be further developed to reduce the uncertainty within the range of natural abundance and hence enable their full potential., (© 2021 The Authors. Rapid Communications in Mass Spectrometry published by John Wiley & Sons Ltd.)

Photosynthetic water-use efficiency (WUE) describes the link between terrestrial carbon (C) and water cycles. Estimates of intrinsic WUE (iWUE) from gas exchange and C isotopic composition (δ 13 C) differ due to an internal conductance in the leaf mesophyll (g m ) that is variable and seldom computed. We present the first direct estimates of whole-tree g m , together with iWUE from whole-tree gas exchange and δ 13 C of the phloem (δ 13 C ph ). We measured gas exchange, online 13 C-discrimination, and δ 13 C ph monthly throughout spring, summer, and autumn in Eucalyptus tereticornis grown in large whole-tree chambers. Six trees were grown at ambient temperatures and six at a 3°C warmer air temperature; a late-summer drought was also imposed. Drought reduced whole-tree g m . Warming had few direct effects, but amplified drought-induced reductions in whole-tree g m . Whole-tree g m was similar to leaf g m for these same trees. iWUE estimates from δ 13 C ph agreed with iWUE from gas exchange, but only after incorporating g m . δ 13 C ph was also correlated with whole-tree 13 C-discrimination, but offset by -2.5 ± 0.7‰, presumably due to post-photosynthetic fractionations. We conclude that δ 13 C ph is a good proxy for whole-tree iWUE, with the caveats that post-photosynthetic fractionations and intrinsic variability of g m should be incorporated to provide reliable estimates of this trait in response to abiotic stress., (© 2020 The Authors New Phytologist © 2020 New Phytologist Foundation.)

Woody plants vary in their adaptations to drought and shade. For a better prediction of vegetation responses to drought and shade within dynamic global vegetation models, it is critical to group species into functional types with similar adaptations. One of the key challenges is that the adaptations are generally determined by a large number of plant traits that may not be available for a large number of species. In this study, we present two heuristic woody plant groups that were separated using cluster analysis in a three-dimensional trait-environment space based on three key metrics for each species: mean xylem embolism resistance, shade tolerance and habitat aridity. The two heuristic groups separate these species into tolerators and avoiders. The tolerators either rely on their high embolism resistance to tolerate drought in arid habitats (e.g., Juniperus and Prunus) or rely on high shade tolerance to withstand shaded conditions in wet habitats (e.g., Picea, Abies and Acer). In contrast, all avoiders have low embolism resistance and low shade tolerance. In arid habitats, avoiders tend to minimize catastrophic embolism (e.g., most Pinus species) while in wet habitats, they may survive despite low shade tolerance (e.g., Betula, Populus, Alnus and Salix). Because our approach links traits to the environmental conditions, we expect it could be a promising framework for predicting changes in species composition, and therefore ecosystem function, under changing environmental conditions., (Published by Oxford University Press 2019.)

Rising atmospheric [CO2 ], ca , is expected to affect stomatal regulation of leaf gas-exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas-exchange that include maintaining a constant leaf internal [CO2 ], ci , a constant drawdown in CO2 (ca  - ci ), and a constant ci /ca . These strategies can result in drastically different consequences for leaf gas-exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas-exchange responses to varying ca . The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas-exchange responses to ca . To assess leaf gas-exchange regulation strategies, we analyzed patterns in ci inferred from studies reporting C stable isotope ratios (δ(13) C) or photosynthetic discrimination (∆) in woody angiosperms and gymnosperms that grew across a range of ca spanning at least 100 ppm. Our results suggest that much of the ca -induced changes in ci /ca occurred across ca spanning 200 to 400 ppm. These patterns imply that ca  - ci will eventually approach a constant level at high ca because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant ci . Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low ca , when additional water loss is small for each unit of C gain, and increasingly water-conservative at high ca , when photosystems are saturated and water loss is large for each unit C gain., (© 2015 John Wiley & Sons Ltd.)

It is well known that whole-plant water-use efficiency (transpiration efficiency of carbon gain, TE(C)) varies among plant species with different photosynthetic pathways. However, less is known of such variation among tree species within the C(3) group. Here we measured the TE(C) of seven C(3) tropical tree species. Isotopic analyses (delta(13)C, delta(18)O, and delta(15)N) and elemental analyses (carbon and nitrogen) were undertaken to provide insight into sources of variation in TE(C). Plants were grown over several months in approx. 80% full sunlight in individual 38-l containers in the Republic of Panama. Soil moisture content was nonlimiting. Significant variation was observed in TE(C) among the C(3) tree species. Values ranged from 1.6 mmol C mol(-1) H(2)O for teak (Tectona grandis) to 4.0 mmol C mol(-1) H(2)O for a legume, Platymiscium pinnatum. Variation in TE(C) was correlated with both leaf N concentration, a proxy for photosynthetic capacity, and oxygen-isotope enrichment, a proxy for stomatal conductance. The TE(C) varied with C-isotope discrimination within species, but the relationship broke down among species, reflecting the existence of species-specific offsets.

The stable oxygen isotope ratio (delta(18)O) of tree-ring cellulose is jointly determined by the delta(18)O of xylem water, the delta(18)O of atmospheric water vapor, the humidity of the atmosphere and perhaps by species-specific differences in leaf structure and function. Atmospheric humidity and the delta(18)O of water vapor vary seasonally and annually, but if the canopy atmosphere is well mixed, atmospheric characteristics should be uniform among co-occurring trees. In contrast, xylem water delta(18)O is determined by the delta(18)O of water being drawn from the soil, which varies with depth. If co-occurring trees draw water from different soil depths, this soil-water delta(18)O signal would be manifest as differences in delta(18)O among the trees. We examined the variation in tree ring delta(18)O, over eight decades during the 20th Century, among three species co-occurring in natural forest stands of the northern Rocky Mountains in the USA. We sampled 10 Douglas-firs (Pseudotsuga menziesii (Mirb.) Franco var. glauca), 10 ponderosa pines (Pinus ponderosa Laws.) and seven western white pines (Pinus monticola Dougl.). As expected, variation in atmospheric conditions was recorded in the delta(18)O of the cellulose produced in a given year, but observed climatic correlations with delta(18)O were weak. Significant correlations with June climate data included: daily maximum temperature (r = 0.29), daily minimum temperature (r = -0.25), mean temperature (r = 0.20), mean daily precipitation (r = -0.54), vapor pressure deficit (r = 0.32) and solar radiation (r = 0.44). Lagged effects were observed in Douglas-fir and western white pine. In these species, the delta(18)O of a given annual ring was correlated with the delta(18)O of the previous ring. Ponderosa pine showed no significant autocorrelation. Although the species means were correlated among years (r = 0.67 to 0.76), ponderosa pine was consistently enriched in delta(18)O relative to the other species; differences were close to 2 per thousand and they are steadily increasing. Relative to the mean for the three species, ponderosa pine is becoming steadily more enriched (-1.0 per thousand). In contrast, Douglas-fir is being steadily depleted and western pine is intermediate, with an enrichment of 0.5 per thousand. Because all trees were exposed to the same atmospheric conditions, the differences in delta(18)O observed between species are likely due either to differences in the depth of water extraction or leaf function. If the former, presumably ponderosa pine has steadily taken up more water from near the soil surface and Douglas-fir has shifted uptake to a greater depth. If the latter, we suggest the pronounced changes in leaf-water delta(18)O are a result of changes in leaf structure and function with tree size and age.

Ecosystem process models are often used to predict carbon flux on a landscape or on a global scale. Such models must be aggregate and canopies are often treated as a uniform unit of foliage. Parameters that are known to vary within the canopy, e.g., nitrogen content and leaf mass per area, are often estimated by a mean value for the canopy. Estimating appropriate means is complicated, especially in mixed-species stands and in complex terrain. We analyzed sources of variation in specific parameters with the goal of testing various simplifying assumptions. The measurements came from mixed-species forests in the northern Rocky Mountains. We found that, for three important parameters (nitrogen concentration and content, and leaf mass per area), a sample taken near the vertical center of the crown provided a good estimate of the mean values for the crown. Altitude (700-1700 m), solar insolation (4200-5400 MJ m(-2) year(-1)) and leaf area index (1-11) had negligible effects on the parameters; only species differences were consistently detected. The correlation between mass-based photosynthetic rates and mass-based nitrogen concentrations was much weaker than the correlation between area-based photosynthetic rates and area-based nitrogen concentration. Comparison of photosynthesis-nitrogen relationships for a wide variety of conifer species and sites revealed a broad general trend that can be used in models. These results suggest important potential simplifications in model parameterization, most notably that canopy means can be estimated with ease, that complex terrain is a minor source of variation in these parameters and that use of one photosynthesis-nitrogen relationship for conifer species does not result in large errors. Species-to-species variation, however, was large and needs to be accounted for when parameterizing process models.

Leaf longevity determines the annual cost of replacing foliage biomass and influences water and nutrient budgets. Longevity is readily estimated in most evergreen species by annual bud scars along the shoot. However, some species with indeterminate growth do not show these annual markers, making estimation of longevity difficult. One of these species is the widespread and economically valuable western red cedar (Thuja plicata J. Donn ex D. Don), for which no dependable estimates of leaf longevity exist. In this study, we estimated leaf longevity for western red cedar by counting growth rings in shoots at the point of leaf abscission. Estimates were obtained on 26 dominant or codominant trees growing in natural stands in a montane forest in northern Idaho, USA. Leaf longevity averaged 8.9 (SE = 0.2) years, but it strongly increased with depth in the canopy (0.3 year m(-1); mean crown depth was 15 m), increasing from a mean of 6.8 years in the upper third of the canopy to 10.6 years in the lower third. The increase in longevity with depth in the crown is consistent with many reports showing that longevity increases in resource-limiting environments. Longevity did not vary significantly with altitude or solar insolation in these montane forests. Among stand-level variables, longevity was correlated only with leaf area index: it increased slightly in stands with high leaf area indices. This approach to longevity estimation may be useful for any species that produces annual rings but no obvious bud scars, including many Cupressaceae species.

The nutrient balance of streams and adjacent riparian ecosystems may be modified by the elimination of anadromous fish runs and perhaps by forest fertilization. To better understand nitrogen (N) dynamics within stream and riparian ecosystems we fertilized two streams and their adjacent riparian corridors in central Idaho. On each stream two nitrogen doses were applied to a swathe approximately 35 m wide centered on the stream. The fertilizer N was enriched in 15N to 18 per thousand. This enrichment is light relative to many previous labeling studies, yet sufficient to yield a traceable signal in riparian and stream biota. This paper reports pre-treatment differences in delta15N and the first-year N response to fertilizer within the riparian woody plant community. Future papers will describe the transfer of allochthonous litter N to the stream and its subsequent processing by stream biota. Pre-treatment delta15N differed between the two creeks (P=0.0002), possibly due to residual salmon nitrogen in one of the creeks. Pre-treatment delta15N of current-year needles was enriched compared to leaf litter, which was in turn enriched compared to needles aged 4 years and older. We conclude that fractionation due to retranslocation occurs in at least two phases. The first phase, which optimizes allocation of N in younger needle age classes, is distinctly different from the second, which conserves N prior to abscission. The delta15N difference between creeks was eliminated by the fertilization (P=0.42). In the two dominant conifer species, Abies lasiocarpa and Picea engelmannii, most fertilizer N was found in the current-year foliage; little was found in older needles and none was detected in litter (P=0.53). The only N-fixing shrub species, Alnus incana, took up only a small amount of fertilizer N [mean percent N derived from fertilizer (%Ndff) 5.0+/-1.6% (SE)]. Far more fertilizer N was taken up by other deciduous shrubs (mean %Ndff=33.9+/-4.5%). Fertilizer N made up 25% (+/-4.2%) of the N in deciduous shrub litter. These results demonstrate the feasibility of light labeling with 15N and the potential influence of riparian plant species composition on stream nutrient dynamics via allochthonous leaf litter inputs.

Forest process models predict ecosystem responses from climate variables and physiological parameters. The parameters describe key ecosystem attributes, often as lumped averages. However, the sources and magnitude of variation in these physiological parameters are unknown, which complicates sampling if models are to be parameterized with field measurements. We measured several key parameters, which had been identified by sensitivity analyses of three models, in Abies grandis (Dougl.) Lindl. and Pseudotsuga menziesii var. glauca (Beissn.) Franco trees throughout the growing season. Trees were sampled at eight sites across the interior northwest of the USA. At each site, fertilized and control plots were sampled. The design provided statistical replication for the analysis of variance within a site, allowing us to draw inferences about a regional population of stands. Specific leaf area (SLA) varied by canopy position and treatment (P = 0.0003), by date of sampling (P < 0.0001) and by species (P = 0.0188). Mass-based foliar nitrogen concentration (%N) increased during the summer in both species (P = 0.0019), but at a faster rate in P. menziesii var. glauca than in A. grandis. Sun foliage had a higher mean %N (1.00, SE = 0.02%) than shade foliage (0.92 +/- 0.01%). Apparent quantum yield (Phi) varied among treatments, between species and between canopy positions; each of these variables interacted with date of sampling (P = 0.0207, P < 0.0001 and P = 0.0344, respectively). In A. grandis, mean Phi values (+/- SE) were 0.048 +/- 0.006 mol CO2 (mol incident photon)-1 for sun foliage and 0.067 +/- 0.007 mol CO2 (mol incident photon)(-1) for shade foliage. In P. menziesii, the corresponding mean Phi values were 0.032 +/- 0.003 and 0.047 +/- 0.004 mol CO2 (mol incident photon)-1. Parameters SLA, %N and Phi were all influenced by date, fertilizer treatment, species and crown position. We discuss methods of inferring quantum yields from light response curves and their utility for parameterizing process models. Parameter mean values are presented for each site; these tables provide a documented data set for the parameterization of models describing the western interior forests of the USA.

We tested the hypothesis that branch hydraulic conductivity partly controls foliar stable carbon isotope ratio (delta13C) by its influence on stomatal conductance in Pinus monticola Dougl. Notching and phloem-girdling treatments were applied to reduce branch conductivity over the course of a growing season. Notching and phloem girdling reduced leaf-specific conductivity (LSC) by about 30 and 90%, respectively. The 90% reduction in LSC increased foliar delta13C by about 1 per thousand (P < 0.0001, n = 65), whereas the 30% reduction in LSC had no effect on foliar delta13C (P = 0.90, n = 65). Variation in the delta13C of dark respiration was similar to that of whole-tissues when compared among treatments. These isotopic measurements, in addition to instantaneous gas exchange measurements, suggested only minor adjustments in the ratio of intercellular to atmospheric CO2 partial pressures (ci/ca) in response to experimentally reduced hydraulic conductivity. A strong correlation was observed between stomatal conductance (gs) and photosynthetic demand over a tenfold range in gs. Although ci/ca and delta13C appeared to be relatively homeostatic, current-year leaf area varied linearly as a function of branch hydraulic conductivity (r2 = 0.69, P < 0.0001, n = 18). These results suggest that, for Pinus monticola, adjustment of leaf area is a more important response to reduced branch conductivity than adjustment of ci/ca.

Univariate time-series analyses were conducted on stable carbon isotope ratios obtained from tree-ring cellulose. We looked for the presence and structure of autocorrelation. Significant autocorrelation violates the statistical independence assumption and biases hypothesis tests. Its presence would indicate the existence of lagged physiological effects that persist for longer than the current year. We analyzed data from 28 trees (60-85 years old; mean = 73 years) of western white pine (Pinus monticola Dougl.), ponderosa pine (Pinus ponderosa Laws.), and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca) growing in northern Idaho. Material was obtained by the stem analysis method from rings laid down in the upper portion of the crown throughout each tree's life. The sampling protocol minimized variation caused by changing light regimes within each tree. Autoregressive moving average (ARMA) models were used to describe the autocorrelation structure over time. Three time series were analyzed for each tree: the stable carbon isotope ratio (delta(13)C); discrimination (delta); and the difference between ambient and internal CO(2) concentrations (c(a) - c(i)). The effect of converting from ring cellulose to whole-leaf tissue did not affect the analysis because it was almost completely removed by the detrending that precedes time-series analysis. A simple linear or quadratic model adequately described the time trend. The residuals from the trend had a constant mean and variance, thus ensuring stationarity, a requirement for autocorrelation analysis. The trend over time for c(a) - c(i) was particularly strong (R(2) = 0.29-0.84). Autoregressive moving average analyses of the residuals from these trends indicated that two-thirds of the individual tree series contained significant autocorrelation, whereas the remaining third were random (white noise) over time. We were unable to distinguish between individuals with and without significant autocorrelation beforehand. Significant ARMA models were all of low order, with either first- or second-order (i.e., lagged 1 or 2 years, respectively) models performing well. A simple autoregressive (AR(1)), model was the most common. The most useful generalization was that the same ARMA model holds for each of the three series (delta(13)C, delta, c(a) - c(i)) for an individual tree, if the time trend has been properly removed for each series. The mean series for the two pine species were described by first-order ARMA models (1-year lags), whereas the Douglas-fir mean series were described by second-order models (2-year lags) with negligible first-order effects. Apparently, the process of constructing a mean time series for a species preserves an underlying signal related to delta(13)C while canceling some of the random individual tree variation. Furthermore, the best model for the overall mean series (e.g., for a species) cannot be inferred from a consensus of the individual tree model forms, nor can its parameters be estimated reliably from the mean of the individual tree parameters. Because two-thirds of the individual tree time series contained significant autocorrelation, the normal assumption of a random structure over time is unwarranted, even after accounting for the time trend. The residuals of an appropriate ARMA model satisfy the independence assumption, and can be used to make hypothesis tests.

We investigated variation in height growth, gas exchange, chlorophyll fluorescence and leaf stable carbon isotope ratio among wind-pollinated progenies of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca), ponderosa pine (Pinus ponderosa Dougl. ex Laws.) and western white pine (Pinus monticola Dougl. ex D. Don) from a small group of contiguous stands on the Priest River Experimental Forest in northern Idaho. Photosynthetic variables differed between height classes in the pines, but not in Douglas-fir. Among species and families, tall families of ponderosa pine regained photosynthetic capacity earliest in the spring and maintained it latest in the growing season. Tall families of western white pine had higher instantaneous water-use efficiencies and lower photosynthetic rates than short families on warm days in August.