Dynamics of stem water uptake among isohydric and anisohydric species experiencing a severe drought.

Predicting the impact of drought on forest ecosystem processes requires an understanding of trees' species-specific responses to drought, especially in the Eastern USA, where species composition is highly dynamic due to historical changes in land use and fire regime. Here, we adapted a framework that classifies trees' water-use strategy along the spectrum of isohydric to anisohydric behavior to determine the responses of three canopy-dominant species to drought. We used a collection of leaf-level gas exchange, tree-level sap flux and stand-level eddy covariance data collected in south-central Indiana from 2011 to 2013, which included an unusually severe drought in the summer of 2012. Our goal was to assess how patterns in the radial profile of sap flux and reliance on hydraulic capacitance differed among species of contrasting water-use strategies. In isohydric species, which included sugar maple (Acer saccharum Marsh.) and tulip poplar (Liriodendron tulipifera L.), we found that the sap flux in the outer xylem experienced dramatic declines during drought, but sap flux at inner xylem was buffered from reductions in water availability. In contrast, for anisohydric oak species (Quercus alba L. and Quercus rubra L.), we observed relatively smaller variations in sap flux during drought in both inner and outer xylem, and higher nighttime refilling when compared with isohydric species. This reliance on nocturnal refilling, which occurred coincident with a decoupling between leaf- and tree-level water-use dynamics, suggests that anisohydric species may benefit from a reliance on hydraulic capacitance to mitigate the risk of hydraulic failure associated with maintaining high transpiration rates during drought. In the case of both isohydric and anisohydric species, our work demonstrates that failure to account for shifts in the radial profile of sap flux during drought could introduce substantial bias in estimates of tree water use during both drought and non-drought periods.

Mycorrhizal type determines the magnitude and direction of root-induced changes in decomposition in a temperate forest.

Although it is increasingly being recognized that roots play a key role in soil carbon (C) dynamics, the magnitude and direction of these effects are unknown. Roots can accelerate soil C losses by provisioning microbes with energy to decompose organic matter or impede soil C losses by enhancing microbial competition for nutrients. We experimentally reduced belowground C supply to soils via tree girdling, and contrasted responses in control and girdled plots for three consecutive growing seasons. We hypothesized that decreases in belowground C supply would have stronger effects in plots dominated by ectomycorrhizal (ECM) trees rather than arbuscular mycorrhizal (AM) trees. In ECM-dominated plots, girdling decreased the activity of enzymes that break down soil organic matter (SOM) by c. 40%, indicating that, in control plots, C supply from ECM roots primes microbial decomposition. In AM-dominated plots, girdling had little effect on SOM-degrading enzymes, but increased the decomposition of AM leaf litter by c. 43%, suggesting that, in control plots, AM roots may intensify microbial competition for nutrients. Our findings indicate that root-induced changes in soil processes depend on forest composition, and that shifts in the distribution of AM and ECM trees owing to climate change may determine soil C gains and losses.

Chronic water stress reduces tree growth and the carbon sink of deciduous hardwood forests.

Predicted decreases in water availability across the temperate forest biome have the potential to offset gains in carbon (C) uptake from phenology trends, rising atmospheric CO2 , and nitrogen deposition. While it is well established that severe droughts reduce the C sink of forests by inducing tree mortality, the impacts of mild but chronic water stress on forest phenology and physiology are largely unknown. We quantified the C consequences of chronic water stress using a 13-year record of tree growth (n = 200 trees), soil moisture, and ecosystem C balance at the Morgan-Monroe State Forest (MMSF) in Indiana, and a regional 11-year record of tree growth (n > 300 000 trees) and water availability for the 20 most dominant deciduous broadleaf tree species across the eastern and midwestern USA. We show that despite ~26 more days of C assimilation by trees at the MMSF, increasing water stress decreased the number of days of wood production by ~42 days over the same period, reducing the annual accrual of C in woody biomass by 41%. Across the deciduous forest region, water stress induced similar declines in tree growth, particularly for water-demanding 'mesophytic' tree species. Given the current replacement of water-stress adapted 'xerophytic' tree species by mesophytic tree species, we estimate that chronic water stress has the potential to decrease the C sink of deciduous forests by up to 17% (0.04 Pg C yr(-1) ) in the coming decades. This reduction in the C sink due to mesophication and chronic water stress is equivalent to an additional 1-3 days of global C emissions from fossil fuel burning each year. Collectively, our results indicate that regional declines in water availability may offset the growth-enhancing effects of other global changes and reduce the extent to which forests ameliorate climate warming.

An improved approach for remotely sensing water stress impacts on forest C uptake.

Given that forests represent the primary terrestrial sink for atmospheric CO2 , projections of future carbon (C) storage hinge on forest responses to climate variation. Models of gross primary production (GPP) responses to water stress are commonly based on remotely sensed changes in canopy 'greenness' (e.g., normalized difference vegetation index; NDVI). However, many forests have low spectral sensitivity to water stress (SSWS) - defined here as drought-induced decline in GPP without a change in greenness. Current satellite-derived estimates of GPP use a vapor pressure deficit (VPD) scalar to account for the low SWSS of forests, but fail to capture their responses to water stress. Our objectives were to characterize differences in SSWS among forested and nonforested ecosystems, and to develop an improved framework for predicting the impacts of water stress on GPP in forests with low SSWS. First, we paired two independent drought indices with NDVI data for the conterminous US from 2000 to 2011, and examined the relationship between water stress and NDVI. We found that forests had lower SSWS than nonforests regardless of drought index or duration. We then compared satellite-derived estimates of GPP with eddy-covariance observations of GPP in two deciduous broadleaf forests with low SSWS: the Missouri Ozark (MO) and Morgan Monroe State Forest (MMSF) AmeriFlux sites. Model estimates of GPP that used VPD scalars were poorly correlated with observations of GPP at MO (r(2) = 0.09) and MMSF (r(2) = 0.38). When we included the NDVI responses to water stress of adjacent ecosystems with high SSWS into a model based solely on temperature and greenness, we substantially improved predictions of GPP at MO (r(2) = 0.83) and for a severe drought year at the MMSF (r(2) = 0.82). Collectively, our results suggest that large-scale estimates of GPP that capture variation in SSWS among ecosystems could improve predictions of C uptake by forests under drought.

Increase in forest water-use efficiency as atmospheric carbon dioxide concentrations rise.

Terrestrial plants remove CO2 from the atmosphere through photosynthesis, a process that is accompanied by the loss of water vapour from leaves. The ratio of water loss to carbon gain, or water-use efficiency, is a key characteristic of ecosystem function that is central to the global cycles of water, energy and carbon. Here we analyse direct, long-term measurements of whole-ecosystem carbon and water exchange. We find a substantial increase in water-use efficiency in temperate and boreal forests of the Northern Hemisphere over the past two decades. We systematically assess various competing hypotheses to explain this trend, and find that the observed increase is most consistent with a strong CO2 fertilization effect. The results suggest a partial closure of stomata-small pores on the leaf surface that regulate gas exchange-to maintain a near-constant concentration of CO2 inside the leaf even under continually increasing atmospheric CO2 levels. The observed increase in forest water-use efficiency is larger than that predicted by existing theory and 13 terrestrial biosphere models. The increase is associated with trends of increasing ecosystem-level photosynthesis and net carbon uptake, and decreasing evapotranspiration. Our findings suggest a shift in the carbon- and water-based economics of terrestrial vegetation, which may require a reassessment of the role of stomatal control in regulating interactions between forests and climate change, and a re-evaluation of coupled vegetation-climate models.

On the calibration of continuous, high-precision delta18O and delta2H measurements using an off-axis integrated cavity output spectrometer.

The (18)O and (2)H of water vapor serve as powerful tracers of hydrological processes. The typical method for determining water vapor delta(18)O and delta(2)H involves cryogenic trapping and isotope ratio mass spectrometry. Even with recent technical advances, these methods cannot resolve vapor composition at high temporal resolutions. In recent years, a few groups have developed continuous laser absorption spectroscopy (LAS) approaches for measuring delta(18)O and delta(2)H which achieve accuracy levels similar to those of lab-based mass spectrometry methods. Unfortunately, most LAS systems need cryogenic cooling and constant calibration to a reference gas, and have substantial power requirements, making them unsuitable for long-term field deployment at remote field sites. A new method called Off-Axis Integrated Cavity Output Spectroscopy (OA-ICOS) has been developed which requires extremely low-energy consumption and neither reference gas nor cryogenic cooling. In this report, we develop a relatively simple pumping system coupled to a dew point generator to calibrate an ICOS-based instrument (Los Gatos Research Water Vapor Isotope Analyzer (WVIA) DLT-100) under various pressures using liquid water with known isotopic signatures. Results show that the WVIA can be successfully calibrated using this customized system for different pressure settings, which ensure that this instrument can be combined with other gas-sampling systems. The precisions of this instrument and the associated calibration method can reach approximately 0.08 per thousand for delta(18)O and approximately 0.4 per thousand for delta(2)H. Compared with conventional mass spectrometry and other LAS-based methods, the OA-ICOS technique provides a promising alternative tool for continuous water vapor isotopic measurements in field deployments.