Do 2 H and 18 O in leaf water reflect environmental drivers differently?

We compiled hydrogen and oxygen stable isotope compositions (δ2 H and δ18 O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ2 H was more closely correlated with δ2 H of xylem water or atmospheric vapour, whereas leaf water δ18 O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ2 H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ18 O. We next expressed leaf water as isotopic enrichment above xylem water (Δ2 H and Δ18 O) to remove the impact of xylem water isotopic variation. For Δ2 H, leaf water still correlated with atmospheric vapour, whereas Δ18 O showed no such correlation. This was explained by covariance between air relative humidity and the Δ18 O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that 2 H and 18 O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water.

Local groundwater decline exacerbates response of dryland riparian woodlands to climatic drought.

Dryland riparian woodlands are considered to be locally buffered from droughts by shallow and stable groundwater levels. However, climate change is causing more frequent and severe drought events, accompanied by warmer temperatures, collectively threatening the persistence of these groundwater dependent ecosystems through a combination of increasing evaporative demand and decreasing groundwater supply. We conducted a dendro-isotopic analysis of radial growth and seasonal (semi-annual) carbon isotope discrimination (Δ13 C) to investigate the response of riparian cottonwood stands to the unprecedented California-wide drought from 2012 to 2019, along the largest remaining free-flowing river in Southern California. Our goals were to identify principal drivers and indicators of drought stress for dryland riparian woodlands, determine their thresholds of tolerance to hydroclimatic stressors, and ultimately assess their vulnerability to climate change. Riparian trees were highly responsive to drought conditions along the river, exhibiting suppressed growth and strong stomatal closure (inferred from reduced Δ13 C) during peak drought years. However, patterns of radial growth and Δ13 C were quite variable among sites that differed in climatic conditions and rate of groundwater decline. We show that the rate of groundwater decline, as opposed to climate factors, was the primary driver of site differences in drought stress, and trees showed greater sensitivity to temperature at sites subjected to faster groundwater decline. Across sites, higher correlation between radial growth and Δ13 C for individual trees, and higher inter-correlation of Δ13 C among trees were indicative of greater drought stress. Trees showed a threshold of tolerance to groundwater decline at 0.5 m year-1 beyond which drought stress became increasingly evident and severe. For sites that exceeded this threshold, peak physiological stress occurred when total groundwater recession exceeded ~3 m. These findings indicate that drought-induced groundwater decline associated with more extreme droughts is a primary threat to dryland riparian woodlands and increases their susceptibility to projected warmer temperatures.

Changes in tree drought sensitivity provided early warning signals to the California drought and forest mortality event.

Climate warming in recent decades has negatively impacted forest health in the western United States. Here, we report on potential early warning signals (EWS) for drought-related mortality derived from measurements of tree-ring growth (ring width index; RWI) and carbon isotope discrimination (∆13 C), primarily focused on ponderosa pine (Pinus ponderosa). Sampling was conducted in the southern Sierra Nevada Mountains, near the epicenter of drought severity and mortality associated with the 2012-2015 California drought and concurrent outbreak of western pine beetle (Dendroctonus brevicomis). At this site, we found that widespread mortality was presaged by five decades of increasing sensitivity (i.e., increased explained variation) of both tree growth and ∆13 C to Palmer Drought Severity Index (PDSI). We hypothesized that increasing sensitivity of tree growth and ∆13 C to hydroclimate constitute EWS that indicate an increased likelihood of widespread forest mortality caused by direct and indirect effects of drought. We then tested these EWS in additional ponderosa pine-dominated forests that experienced varying mortality rates associated with the same California drought event. In general, drier sites showed increasing sensitivity of RWI to PDSI over the last century, as well as higher mortality following the California drought event compared to wetter sites. Two sites displayed evidence that thinning or fire events that reduced stand basal area effectively reversed the trend of increasing hydroclimate sensitivity. These comparisons indicate that reducing competition for soil water and/or decreasing bark beetle host tree density via forest management-particularly in drier regions-may buffer these forests against drought stress and associated mortality risk. EWS such as these could provide land managers more time to mitigate the extent or severity of forest mortality in advance of droughts. Substantial efforts at deploying additional dendrochronological research in concert with remote sensing and forest modeling will aid in forecasting of forest responses to continued climate warming.

Fire deficits have increased drought sensitivity in dry conifer forests: Fire frequency and tree-ring carbon isotope evidence from Central Oregon.

A century of fire suppression across the Western United States has led to more crowded forests and increased competition for resources. Studies of forest thinning or stand conditions after mortality events have provided indirect evidence for how competition can promote drought stress and predispose forests to severe fire and/or bark beetle outbreaks. Here, we demonstrate linkages between fire deficits and increasing drought stress through analyses of annually resolved tree-ring growth, fire scars, and carbon isotope discrimination (Δ13 C) across a dry mixed-conifer forest landscape. Fire deficits across the study area have increased the sensitivity of leaf gas exchange to drought stress over the past >100 years. Since 1910, stand basal area in these forests has more than doubled and fire-return intervals have increased from 25 to 140 years. Meanwhile, the portion of interannual variation in tree-ring Δ13 C explained by the Palmer Drought Severity Index has more than doubled in ca. 300-500-year-old Pinus ponderosa as well as in fire-intolerant, ca. 90-190-year-old Abies grandis. Drought stress has increased in stands with a basal area of ≥25 m2 /ha in 1910, as indicated by negative temporal Δ13 C trends, whereas stands with basal area ≤25 m2 /ha in 1910, due to frequent or intense wildfire activity in decades beforehand, were initially buffered from increased drought stress and have benefited more from rising ambient carbon dioxide concentrations, [CO2 ], as demonstrated by positive temporal Δ13 C trends. Furthermore, the average Δ13 C response across all P. ponderosa since 1830 indicates that photosynthetic assimilation rates and stomatal conductance have been reduced by ~10% and ~20%, respectively, compared to expected trends due to increasing [CO2 ]. Although disturbance legacies contribute to local-scale intensity of drought stress, fire deficits have reduced drought resistance of mixed-conifer forests and made them more susceptible to challenges by pests and pathogens and other disturbances.

Millennial-scale tree-ring isotope chronologies from coast redwoods provide insights on controls over California hydroclimate variability.

To understand drivers of hydroclimate variability in north-coastal California, we obtained tree cross-sections from eleven coastal redwoods (mean age of 1232 years old) from the northern half of the species range. Tree rings from eight trees were cross-dated and sampled at sub-annual resolution for carbon isotope discrimination (Δ13C) and oxygen isotope composition (δ18O). Tree-ring Δ13C and δ18O, compared to modern climate data, demonstrate these signals primarily record summertime hydroclimate variability-primarily through variables associated with evaporative conditions and/or precipitation. Our 1100-year stable isotope chronologies showed that north-coastal California did not undergo the megadroughts observed elsewhere in California and the western United States. This result implicates extended periods of low winter precipitation, rather than growing season evaporation, as the primary driver of previous megadroughts across California and neighboring regions. Compared to cool conditions prevailing over the Northern Hemisphere during the Little Ice age (1301-1875 of the common era, CE), the frequency of isotopic events of a certain magnitude was greater during periods with warmer Northern Hemisphere temperatures such as the Medieval Climate Anomaly (900-1300 CE) and the modern period (1876 to present). This association between tree-ring isotopic variability and long-term shifts in temperatures is consistent with the expected patterns in mid-latitude hydroclimate variability expected from arctic amplification (i.e., shifts in equator-to-pole temperature differences that modify jet stream speed and amplitude) or amplified quasi-resonant wave activity (i.e., wave-patterns in high-altitude winds that become "trapped" within a certain pattern, thereby producing a longer-duration periods of drought or wetness) across mid-latitudes during the boreal summer.

Evidence that higher [CO2] increases tree growth sensitivity to temperature: a comparison of modern and paleo oaks.

To test tree growth sensitivity to temperature under different ambient CO2 concentrations, we determined stem radial growth rates as they relate to variation in temperature during the last deglacial period, and compare these to modern tree growth rates as they relate to spatial variation in temperature across the modern species distributional range. Paleo oaks were sampled from Northern Missouri, USA and compared to a pollen-based, high-resolution paleo temperature reconstruction from Northern Illinois, USA. Growth data were from 53 paleo bur oak log cross sections collected in Missouri. These oaks were preserved in river and stream sediments and were radiocarbon-dated to a period of rapid climate change during the last deglaciation (10.5 and 13.3 cal kyr BP). Growth data from modern bur oaks were obtained from increment core collections paired with USDA Forest Service Forest Inventory and Analysis data collected across the Great Plains, Midwest, and Upper Great Lakes regions. For modern oaks growing at an average [CO2] of 330 ppm, growth sensitivity to temperature (i.e., the slope of growth rate versus temperature) was about twice that of paleo oaks growing at an average [CO2] of 230 ppm. These data help to confirm that leaf-level predictions that photosynthesis and thus growth will be more sensitive to temperature at higher [CO2] in mature trees-suggesting that tree growth forest productivity will be increasingly sensitive to temperature under projected global warming and high-[CO2] conditions.

Basal area growth, carbon isotope discrimination, and intrinsic water use efficiency after fertilization of Douglas-fir in the Oregon Coast Range.

Many hectares of intensively managed Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands in western North America are fertilized with nitrogen (N) to increase growth rates, but only about ⅔ of all stands respond. Understanding the mechanisms of response facilitates prioritization of stands for treatment. The primary objective of this study was to test the hypothesis that the short-term basal area growth response to a single application of 224 kg N ha-1 as urea was associated with reduced stable carbon isotope discrimination (Δ13C) and increased intrinsic water use efficiency (iWUE) in a 20-yr-old plantation of Douglas-fir in the Oregon Coast Range, USA. Increment cores were measured to estimate earlywood, latewood, and total basal area increment over a time series from 1997 to 2015. Stable carbon isotope discrimination and iWUE were estimated using earlywood and latewood stable carbon isotope concentrations in tree-ring holocellulose starting seven years before fertilization in early 2009 and ending seven years after treatment. A highly significant (p<0.01) interaction effect between fertilization treatment and year was found for total basal area growth and earlywood basal area increment. Specifically, fertilized trees showed significant responses (p<0.05) in total basal area growth and earlywood basal area increment in the first (2009) and second (2010) growing seasons after fertilization in 2009. A marginally significant (p<0.10) fertilization effect was found for latewood basal area increment only in the first growing season after treatment. A significant treatment x year interaction was also found for Δ13C and iWUE in earlywood and latewood. Fertilization significantly reduced earlywood Δ13C and increased earlywood iWUE in the first and second growing seasons after fertilization. Only a marginally significant fertilization effect was detected for latewood Δ13C and iWUE in the second growing season after treatment. Previous studies of N fertilization of Douglas-fir forests have reported consistently increased growth and iWUE on low productivity sites treated with relatively high fertilization rates. This study suggested that these responses can also be observed on highly productive sites despite their lower frequency and apparently shorter duration. Other key mechanisms driving growth responses appear less important than iWUE, including an increase in LAI and shift from belowground to aboveground carbon allocation.

A dynamic leaf gas-exchange strategy is conserved in woody plants under changing ambient CO2 : evidence from carbon isotope discrimination in paleo and CO2 enrichment studies.

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.

Drivers of radial growth and carbon isotope discrimination of bur oak (Quercus macrocarpa†Michx.) across continental gradients in precipitation, vapour pressure deficit and irradiance.

Tree-ring characteristics are commonly used to reconstruct climate variables, but divergence from the assumption of a single biophysical control may reduce the accuracy of these reconstructions. Here, we present data from bur oaks (Quercus macrocarpa Michx.) sampled within and beyond the current species bioclimatic envelope to identify the primary environmental controls on ring-width indices (RWIs) and carbon stable isotope discrimination (Δ(13) C) in tree-ring cellulose. Variation in Δ(13) C and RWI was more strongly related to leaf-to-air vapour pressure deficit (VPD) at the centre and western edge of the range compared with the northern and wettest regions. Among regions, Δ(13) C of tree-ring cellulose was closely predicted by VPD and light responses of canopy-level Δ(13) C estimated using a model driven by eddy flux and meteorological measurements (R(2) = 0.96, P = 0.003). RWI and Δ(13) C were positively correlated in the drier regions, while they were negatively correlated in the wettest region. The strength and direction of the correlations scaled with regional VPD or the ratio of precipitation to evapotranspiration. Therefore, the correlation strength between RWI and Δ(13) C may be used to infer past wetness or aridity from paleo wood by determining the degree to which carbon gain and growth have been more limited by moisture or light.

Tree-ring stable isotopes record the impact of a foliar fungal pathogen on CO(2) assimilation and growth in Douglas-fir.

Swiss needle cast (SNC) is a fungal disease of Douglas-fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree-rings of Douglas-fir and a non-susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ(13)C) of treated Douglas-fir tree-rings was greater than that of untreated Douglas-fir tree-rings during the fungicide treatment period. Both annual growth and tree-ring Δ(13)C increased with treatment such that treated Douglas-fir had values similar to co-occurring western hemlock during the treatment period. There was no difference in the tree-ring oxygen stable isotope ratio between treated and untreated Douglas-fir. Tree-ring Δ(13)C of diseased Douglas-fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years.

Reconstructing relative humidity from plant delta18O and deltaD as deuterium deviations from the global meteoric water line.

Cellulose delta18O and deltaD can provide insights on climates and hydrological cycling in the distant past and how these factors differ spatially. However, most studies of plant cellulose have used only one isotope, most commonly delta18O, resulting in difficulties partitioning variation in delta18O of precipitation vs. evaporative conditions that affect leaf water isotopic enrichment. Moreover, observations of pronounced diurnal differences from conventional steady-state model predictions of leaf water isotopic fractionation have cast some doubt on single isotope modeling approaches for separating precipitation and evaporation drivers of cellulose delta18O or deltaD. We explore a dual isotope approach akin to the concept of deuterium-excess (d), to establish deuterium deviations from the global meteoric water line in leaf water (deltad(l)) as driven by relative humidity (RH). To demonstrate this concept, we survey studies of leaf water delta18O and deltaD in hardwood vs. conifer trees. We then apply the concept to cellulose delta18O and deltaD using a mechanistic model of cellulose delta18O and deltaD to reconstruct deuterium deviations from the global meteoric water line (deltad(c)) in Quercus macrocarpa, Q. robur, and Pseudotsuga menziesii. For each species, deltad(c) showed strong correlations with RH across sites. deltad(c) agreed well with steady-state predictions for Q. macrocarpa, while for Q. robur, the relationship with RH was steeper than expected. The slope of deltad(c) vs. RH of P. menziesii was also close to steady-state predictions, but deltad(c) were more enriched than predicted. This is in agreement with our leaf water survey showing conifer deltad(l) was more enriched than predicted. Our data reveal that applications of this method should be appropriate for reconstructing RH from cellulose delta18O and deltaD after accounting for differences between hardwoods and conifers. Hence, deltad(c) should be useful for understanding variability in RH associated with past climatic cycles, across regional climates, or across complex terrain where climate modeling is challenging. Furthermore, deltad(c) and inferred RH values should help in constraining variation in source water delta18O.

Hydraulic architecture of two species differing in wood density: opposing strategies in co-occurring tropical pioneer trees.

Co-occurring species often have different strategies for tolerating daily cycles of water stress. One underlying parameter that can link together the suite of traits that enables a given strategy is wood density. Here we compare hydraulic traits of two pioneer species from a tropical forest in Panama that differ in wood density: Miconia argentea and Anacardium excelsum. As hypothesized, the higher wood density of Miconia was associated with smaller diameter vessels and fibres, more water stress-resistant leaves and stems, and roughly half the capacitance of the lower wood density Anacardium. However, the scaling of hydraulic parameters such as the increases in leaf area and measures of hydraulic conductivity with stem diameter was remarkably similar between the two species. The collection of traits exhibited by Miconia allowed it to tolerate more water stress than Anacardium, which relied more heavily on its capacitance to buffer daily water potential fluctuations. This work demonstrates the importance of examining a range of hydraulic traits throughout the plant and highlights the spectrum of possible strategies for coping with daily and seasonal water stress cycles.

Cross-biome synthesis of source versus sink limits to tree growth.

Uncertainties surrounding tree carbon allocation to growth are a major limitation to projections of forest carbon sequestration and response to climate change. The prevalence and extent to which carbon assimilation (source) or cambial activity (sink) mediate wood production are fundamentally important and remain elusive. We quantified source-sink relations across biomes by combining eddy-covariance gross primary production with extensive on-site and regional tree ring observations. We found widespread temporal decoupling between carbon assimilation and tree growth, underpinned by contrasting climatic sensitivities of these two processes. Substantial differences in assimilation-growth decoupling between angiosperms and gymnosperms were determined, as well as stronger decoupling with canopy closure, aridity, and decreasing temperatures. Our results reveal pervasive sink control over tree growth that is likely to be increasingly prominent under global climate change.

Reduced wood stiffness and strength, and altered stem form, in young antisense 4CL transgenic poplars with reduced lignin contents.

• Reduced lignin content in perennial crops has been sought as a means to improve biomass processability for paper and biofuels production, but it is unclear how this could affect wood properties and tree form. • Here, we studied a nontransgenic control and 14 transgenic events containing an antisense 4-coumarate:coenzyme A ligase (4CL) to discern the consequences of lignin reduction in poplar (Populus sp.). During the second year of growth, trees were grown either free-standing in a field trial or affixed to stakes in a glasshouse. • Reductions in lignin of up to 40% gave comparable losses in wood strength and stiffness. This occurred despite the fact that low-lignin trees had a similar wood density and up to three-fold more tension wood. In free-standing and staked trees, the control line had twice the height for a given diameter as did low-lignin trees. Staked trees had twice the height for a given diameter as free-standing trees in the field, but did not differ in wood stiffness. • Variation in tree morphogenesis appears to be governed by lignin x environment interactions mediated by stresses exerted on developing cells. Therefore our results underline the importance of field studies for assessing the performance of transgenic trees with modified wood properties.

Tyloses and phenolic deposits in xylem vessels impede water transport in low-lignin transgenic poplars: a study by cryo-fluorescence microscopy.

Of 14 transgenic poplar genotypes (Populus tremula × Populus alba) with antisense 4-coumarate:coenzyme A ligase that were grown in the field for 2 years, five that had substantial lignin reductions also had greatly reduced xylem-specific conductivity compared with that of control trees and those transgenic events with small reductions in lignin. For the two events with the lowest xylem lignin contents (greater than 40% reduction), we used light microscopy methods and acid fuchsin dye ascent studies to clarify what caused their reduced transport efficiency. A novel protocol involving dye stabilization and cryo-fluorescence microscopy enabled us to visualize the dye at the cellular level and to identify water-conducting pathways in the xylem. Cryo-fixed branch segments were planed in the frozen state on a sliding cryo-microtome and observed with an epifluorescence microscope equipped with a cryo-stage. We could then distinguish clearly between phenolic-occluded vessels, conductive (stain-filled) vessels, and nonconductive (water- or gas-filled) vessels. Low-lignin trees contained areas of nonconductive, brown xylem with patches of collapsed cells and patches of noncollapsed cells filled with phenolics. In contrast, phenolics and nonconductive vessels were rarely observed in normal colored wood of the low-lignin events. The results of cryo-fluorescence light microscopy were supported by observations with a confocal microscope after freeze drying of cryo-planed samples. Moreover, after extraction of the phenolics, confocal microscopy revealed that many of the vessels in the nonconductive xylem were blocked with tyloses. We conclude that reduced transport efficiency of the transgenic low-lignin xylem was largely caused by blockages from tyloses and phenolic deposits within vessels rather than by xylem collapse.

Antisense down-regulation of 4CL expression alters lignification, tree growth, and saccharification potential of field-grown poplar.

Transgenic down-regulation of the Pt4CL1 gene family encoding 4-coumarate:coenzyme A ligase (4CL) has been reported as a means for reducing lignin content in cell walls and increasing overall growth rates, thereby improving feedstock quality for paper and bioethanol production. Using hybrid poplar (Populus tremula × Populus alba), we applied this strategy and examined field-grown transformants for both effects on wood biochemistry and tree productivity. The reductions in lignin contents obtained correlated well with 4CL RNA expression, with a sharp decrease in lignin amount being observed for RNA expression below approximately 50% of the nontransgenic control. Relatively small lignin reductions of approximately 10% were associated with reduced productivity, decreased wood syringyl/guaiacyl lignin monomer ratios, and a small increase in the level of incorporation of H-monomers (p-hydroxyphenyl) into cell walls. Transgenic events with less than approximately 50% 4CL RNA expression were characterized by patches of reddish-brown discolored wood that had approximately twice the extractive content of controls (largely complex polyphenolics). There was no evidence that substantially reduced lignin contents increased growth rates or saccharification potential. Our results suggest that the capacity for lignin reduction is limited; below a threshold, large changes in wood chemistry and plant metabolism were observed that adversely affected productivity and potential ethanol yield. They also underline the importance of field studies to obtain physiologically meaningful results and to support technology development with transgenic trees.

Transgenic poplars with reduced lignin show impaired xylem conductivity, growth efficiency and survival.

We studied xylem anatomy and hydraulic architecture in 14 transgenic insertion events and a control line of hybrid poplar (Populus spp.) that varied in lignin content. Transgenic events had different levels of down-regulation of two genes encoding 4-coumarate:coenzyme A ligase (4CL). Two-year-old trees were characterized after growing either as free-standing trees in the field or as supported by stakes in a greenhouse. In free-standing trees, a 20 to 40% reduction in lignin content was associated with increased xylem vulnerability to embolism, shoot dieback and mortality. In staked trees, the decreased biomechanical demands on the xylem was associated with increases in the leaf area to sapwood area ratio and wood specific conductivity (k(s)), and with decreased leaf-specific conductivity (k(l)). These shifts in hydraulic architecture suggest that the bending stresses perceived during growth can affect traits important for xylem water transport. Severe 4CL-downregulation resulted in the patchy formation of discoloured, brown wood with irregular vessels in which water transport was strongly impeded. These severely 4CL-downregulated trees had significantly lower growth efficiency (biomass/leaf area). These results underscore the necessity of adequate lignification for mechanical support of the stem, water transport, tree growth and survival.

Comparative hydraulic architecture of tropical tree species representing a range of successional stages and wood density.

Plant hydraulic architecture (PHA) has been linked to water transport sufficiency, photosynthetic rates, growth form and attendant carbon allocation. Despite its influence on traits central to conferring an overall competitive advantage in a given environment, few studies have examined whether key aspects of PHA are indicative of successional stage, especially within mature individuals. While it is well established that wood density (WD) tends to be lower in early versus late successional tree species, and that WD can influence other aspects of PHA, the interaction of WD, successional stage and the consequent implications for PHA have not been sufficiently explored. Here, we studied differences in PHA at the scales of wood anatomy to whole-tree hydraulic conductance in species in early versus late successional Panamanian tropical forests. Although the trunk WD was indistinguishable between the successional groups, the branch WD was lower in the early successional species. Across all species, WD correlated negatively with vessel diameter and positively with vessel packing density. The ratio of branch:trunk vessel diameter, branch sap flux and whole-tree leaf-specific conductance scaled negatively with branch WD across species. Pioneer species showed greater sap flux in branches than in trunks and a greater leaf-specific hydraulic conductance, suggesting that pioneer species can move greater quantities of water at a given tension gradient. In combination with the greater water storage capacitance associated with lower WD, these results suggest these pioneer species can save on the carbon expenditure needed to build safer xylem and instead allow more carbon to be allocated to rapid growth.

Tree-ring isotopes adjacent to Lake Superior reveal cold winter anomalies for the Great Lakes region of North America.

Tree-ring carbon isotope discrimination (Δ13C) and oxygen isotopes (δ18O) collected from white pine (Pinus strobus) trees adjacent to Lake Superior show potential to produce the first winter-specific paleoclimate reconstruction with inter-annual resolution for this region. Isotopic signatures from 1976 to 2015 were strongly linked to antecedent winter minimum temperatures (Tmin), Lake Superior peak ice cover, and regional to continental-scale atmospheric winter pressure variability including the North American Dipole. The immense thermal inertia of Lake Superior underlies the unique connection between winter conditions and tree-ring Δ13C and δ18O signals from the following growing season in trees located near the lake. By combining these signals, we demonstrate feasibility to reconstruct variability in Tmin, ice cover, and continental-scale atmospheric circulation patterns (r ≥ 0.65, P < 0.001).

Impacts of dwarf mistletoe on the physiology of host Tsuga heterophylla trees as recorded in tree-ring C and O stable isotopes.

Dwarf mistletoes, obligate, parasitic plants with diminutive aerial shoots, have long-term effects on host tree water relations, hydraulic architecture and photosynthetic gas exchange and can eventually induce tree death. To investigate the long-term (1886-2010) impacts of dwarf mistletoe on the growth and gas exchange characteristics of host western hemlock, we compared the diameter growth and tree-ring cellulose stable carbon (C) and oxygen (O) isotope ratios (δ(13)Ccell, δ(18)Ocell) of heavily infected and uninfected trees. The relative basal area growth of infected trees was significantly greater than that of uninfected trees in 1886-90, but declined more rapidly in infected than uninfected trees through time and became significantly lower in infected than uninfected trees in 2006-10. Infected trees had significantly lower δ(13)Ccell and δ(18)Ocell than uninfected trees. Differences in δ(18)Ocell between infected and uninfected trees were unexpected given that stomatal conductance and environmental variables that were expected to influence the δ(18)O values of leaf water were similar for both groups. However, estimates of mesophyll conductance (gm) were significantly lower and estimates of effective path length for water movement (L) were significantly higher in leaves of infected trees, consistent with their lower values of δ(18)Ocell. This study reconstructs the long-term physiological responses of western hemlock to dwarf mistletoe infection. The long-term diameter growth and δ(13)Ccell trajectories suggested that infected trees were growing faster than uninfected trees prior to becoming infected and subsequently declined in growth and leaf-level photosynthetic capacity compared with uninfected trees as the dwarf mistletoe infection became severe. This study further points to limitations of the dual-isotope approach for identifying sources of variation in δ(13)Ccell and indicates that changes in leaf internal properties such as gm and L that affect δ(18)Ocell must be considered.