Throughfall Exclusion Research Articles

Acclimation of mature spruce and beech to five years of repeated summer drought – The role of stomatal conductance and leaf area adjustment for water use

Forests globally are experiencing severe droughts, leading to significant reductions in growth, crown dieback and even tree mortality. The ability of forest ecosystems to acclimate to prolonged and repeated droughts is critical for their survival with ongoing climate change. In a five-year throughfall exclusion experiment, we investigated the long-term physiological and morphological acclimation of mature Norway spruce (Picea abies [L.] KARST.) and European beech (Fagus sylvatica L.) to repeated summer drought at the leaf, shoot and whole tree level. Throughout the drought period, spruce reduced their total water use by 70 % to only 4–9 L per day and tree, while beech was less affected with about 30 % reduction of water use. During the first two summers, spruce achieved this by closing their stomata by up to 80 %. Additionally, from the second drought summer onwards, spruce produced shorter shoots and needles, resulting in a stepwise reduction of total leaf area of over 50 % by the end of the experiment. Surprisingly, no premature leaf loss was observed. This reduction in leaf area allowed a gradual increase in stomatal conductance. After the five-year drought experiment, water consumption per leaf area was the same as in the controls, while the total water consumption of spruce was still reduced. In contrast, beech showed no significant reduction in whole-tree leaf area, but nevertheless reduced water use by up to 50 % by stomatal closure. If the restriction of transpiration by stomatal closure is sufficient to ensure survival of Norway spruce during the first drought summers, then the slow but steady reduction in leaf area will ensure successful acclimation of water use, leading to reduced physiological drought stress and long-term survival. Neighboring beech appeared to benefit from the water-saving strategy of spruce by using the excess water.

Experimental warming and drying increase older carbon contributions to soil respiration in lowland tropical forests

Tropical forests account for over 50% of the global terrestrial carbon sink, but climate change threatens to alter the carbon balance of these ecosystems. We show that warming and drying of tropical forest soils may increase soil carbon vulnerability, by increasing degradation of older carbon. In situ whole-profile heating by 4 °C and 50% throughfall exclusion each increased the average radiocarbon age of soil CO2 efflux by ~2–3 years, but the mechanisms underlying this shift differed. Warming accelerated decomposition of older carbon as increased CO2 emissions depleted newer carbon. Drying suppressed decomposition of newer carbon inputs and decreased soil CO2 emissions, thereby increasing contributions of older carbon to CO2 efflux. These findings imply that both warming and drying, by accelerating the loss of older soil carbon or reducing the incorporation of fresh carbon inputs, will exacerbate soil carbon losses and negatively impact carbon storage in tropical forests under climate change.

Soil water consumption along the profile in response to soil water content variations in a black locust plantation with a rainfall exclusion in Loess Plateau

Soil drying is frequently observed in mature forests in water-limited regions, and poses a threat to sustainable development of the ecosystems. Herein, we assessed the effects of rainfall reduction on the water-use characteristics of a black locust (Robinia pseudoacacia) plantation in a sub-humid region of the Loess Plateau, China. Paired plots were established in the plantation, including a treatment with 30 % throughfall exclusion and a control. We comparatively investigated the dynamics of soil water content (SWC) throughout the profile in the two plots after four years of the treatment and determined the contribution of soil water storage (SWS) in each soil layer to water consumption according to the SWS budget. While SWC across the profile in the control plot showed general responses to replenishment and consumption under rainy and dry weather conditions, respectively, the treatment plot had smaller replenishment and consumption in deep layers, causing an aggravated deficiency in SWS and the formation of a temporary dried soil layer (DSL) at a depth of 55–100 cm. The cumulative decrement in SWS during each dry event (consecutive rainless days of ≥ 5-day duration) in the control plot was significantly correlated with SWC at 100–160 cm, suggesting a substantial contribution of water consumption from this layer. However, the treatment plot showed a major contribution to water consumption from the 0–100 cm soil layer during dry events. The vertical distribution of the root system supports these findings, which differed from our hypothesis that deep soil water would be used in response to reduced rainfall input. The distribution of fine root density in the treatment plot was relatively shallower (0–22 and 0–73.1 cm accounting for 50 % and 90 % of fine roots, respectively) than in the control plot (0–31.2 and 0–103.5 cm for corresponding values). These findings indicate that soil water deficiency resulted in the formation of DSL, which hindered water consumption and root development deeper in the soil profile. The severity and recovery of the DSL may be jointly determined by the vertical distribution of SWS and transpiration demand of the plantation. These findings provide insights for water resource management and sustainable development of the plantations in this region.

Resin duct production of loblolly pine (Pinus taeda) in southeastern Oklahoma, USA is positively related to water availability, fertilization, and thinning

Bark beetle outbreaks frequently kill large areas of loblolly pine (Pinus taeda). Resin production is the main defense against these insects and resin duct metrics and areas are correlated with both resin flow and tree survival post bark beetle attack. Whether the number, area, or size of resin ducts is increase when trees grow faster or whether there is a tradeoff between tree growth and resin ducts is currently debated. We determined the impacts of water availability (ambient vs 30 % throughfall reduction), nutrient availability (fertilized vs non-fertilized), and thinning on vertical resin duct count (ducts per 5 mm strip of annual ring), density (ducts mm−2), area (duct area per 5 mm strip of annual ring), relative area (% duct area per ring), and average duct size (mm−2) in planted stands of loblolly pine (ages 6–13) in southeastern Oklahoma. Fertilization increased diameter growth (10 %) and metrics that scale to ring width, i.e., duct count (15 %) and duct area (22 %), but also duct relative area (8 %). Throughfall exclusion decreased diameter growth and duct count in some years, increased duct density, and reduced duct average size (8 %). The net effect was that fertilized stands with ambient water availability had the greatest resource availability among treatments in this study and had the greatest duct count, duct area, and duct relative area as well as average duct size greater than the throughfall exclusion treatments. During the post thinning period, i.e., ages 10–13, thinning increased diameter growth from 9.63 to 12.05 mm year−1, duct count from 9.69 to 12.22, duct area from 0.384 to 0.514 mm2, duct relative area from 1.67 % to 1.84 %, and average duct size from 0.0396 to 0.0428 mm2. Earlywood composed approximately 2/3 of the annual ring, but latewood had almost 5 times greater duct density such that there were more ducts and greater duct areas in the latewood. Our assessment of duct metrics provides a link between tree growth rate, resource availability, and duct metrics associated with resistance to bark beetles. Therefore, silvicultural practices used to increase tree growth likely also increase resistance to bark beetles.

Fungi and bacteria trade-off mediates drought-induced reduction in wood decomposition

Climate change has significantly increased the frequency and intensity of drought events in recent decades, which may affect the decomposition of organic matter such as deadwood. Previous studies have examined the impacts of microclimate and wood traits on deadwood decomposition, but how wood microbes regulate effects of drought intensity on deadwood decomposition remains unclear. In this study, a field drought experiment was conducted with three throughfall exclusion levels (i.e., control, −35% and −70% rainfall treatments) in a subtropical forest to probe relative importance of microclimate, wood traits, and microbial biomass on wood decomposition. Our results showed that the −35% and −70% rainfall treatments significantly decreased wood CO2 efflux by 28.27% and 47.49%, respectively. Drought-induced decreases in wood CO2 efflux were mainly mediated by wood microbial biomass, particularly wood fungi biomass. The structural equation modelling indicated a shift in the dominant wood microbial communities in regulating wood CO2 efflux from bacteria to fungi as drought intensities increased. Our findings highlight the crucial role of wood microbial community with the trade-off between fungi and bacteria on deadwood decomposition under drought, which should be taken into account to decode forest carbon cycle − climate feedback in the future research.

Physiological responses of a black locust plantation to drought stress based on a throughfall exclusion experiment in semi-arid northwestern China

Abstract Drought poses a significant threat on the ecosystem stability of extensive areas of black locust (Robinia pseudoacacia L.) plantations in northwestern China. However, limited understanding of the physiological responses of black locust to drought has impeded the development of proactive measures to alleviate potential adverse effects of drought. This study investigated the physiological impacts of varying drought intensities, manipulated by a throughfall exclusion experiment, on a 20-year-old black locust plantation in northwestern China. The experiment involved 40% throughfall exclusion for moderate drought, 80% exclusion for extreme drought, and no exclusion for control. One year after the implementation of the experiment, both predawn (Ψpd) and midday (Ψmd) leaf water potentials were significantly lower under drought treatments compared to those in control (P < .01). Stomatal conductance (gs) exhibited a strong reduction, leading to decreased leaf transpiration and photosynthesis under drought. However, the reduction in gs did not effectively prevent the decrease in Ψmd. Instead, both Ψpd and Ψmd became more negative with increasing drought stress, but their difference remaining relatively constant (being ~1.1 MPa) across treatments. These results suggest that black locust adopts a balanced water regulatory strategy between isohydry and anisohydry to cope with drought stress. These results contribute to an enhanced understanding of the crucial physiological responses of black locust under drought stress, offering valuable insights for future management strategies aimed at sustaining the ecosystem stability of black locust plantations in an increasingly arid climate.

Drought shortens subtropical understory growing season by advancing leaf senescence.

Subtropical forests, recognized for their intricate vertical canopy stratification, exhibit high resistance to extreme drought. However, the response of leaf phenology to drought in the species-rich understory remains poorly understood. In this study, we constructed a digital camera system, amassing over 360,000 images through a 70% throughfall exclusion experiment, to explore the drought response of understory leaf phenology. The results revealed a significant advancement in understory leaf senescence phenology under drought, with 11.75 and 15.76 days for the start and end of the leaf-falling event, respectively. Pre-season temperature primarily regulated leaf development phenology, whereas soil water dominated the variability in leaf senescence phenology. Under drought conditions, temperature sensitivities for the end of leaf emergence decreased from -13.72 to -11.06 days °C-1, with insignificance observed for the start of leaf emergence. Consequently, drought treatment shortened both the length of the growing season (15.69 days) and the peak growth season (9.80 days) for understory plants. Moreover, this study identified diverse responses among intraspecies and interspecies to drought, particularly during the leaf development phase. These findings underscore the pivotal role of water availability in shaping understory phenology patterns, especially in subtropical forests.

Effects of Rainfall Exclusion Treatment on Photosynthetic Characteristics of Black Locust in the Sub-Humid Region of the Loess Plateau, China.

The mesic-origin species Robinia pseudoacacia L. (black locust) is widely planted in the semiarid and sub-humid areas of the Loess Plateau for the reforestation of vegetation-degraded land. Under the scenario of changing precipitation patterns, exploring the response of photosynthesis to drought allows us to assess the risk to sustainable development of these plantations. In this study, paired plots were established including the control and a treatment of 30% exclusion of throughfall (since 2018). The photosynthetic characteristics were investigated using a portable photosynthesis system for four periods in the full-leaf growing season of 2021-2022, the fourth and fifth years, on both treated and controlled sampling trees. Leaf gas exchange parameters derived from diurnal changing patterns, light response curves, and CO2 response curves showed significant differences except for period II (9-11 September 2021) between the two plots. The photosynthetic midday depression was observed in 2022 in the treated plot. Meanwhile, the decline of net photosynthetic rate in the treated plot was converted from stomatal limitation to non-stomatal limitation. Furthermore, we observed that black locust adapted to long-term water deficiency by reducing stomatal conductance, increasing water use efficiency and intrinsic water use efficiency. The results demonstrate that reduction in precipitation would cause photosynthesis decrease, weaken the response sensitivity to light and CO2, and potentially impair photosynthetic resilience of the plantations. They also provide insights into the changes in photosynthetic functions under global climate change and a reference for management of plantations.

Drought-induced changes in rare microbial community promoted contribution of microbial necromass C to SOC in a subtropical forest

Microbial necromass is considered as one of the primary drivers in soil organic carbon (SOC) formation and persistence. However, how microbial traits (e.g., diversity, composition) influence microbial necromass C and its contribution to SOC remains unclear, especially under persistent drought. Here, we took advantage of a 7-year Throughfall Exclusion Experiment (TEE) to examine effects of drought on microbial necromass C and its contribution to SOC at different soil depths in a subtropical forest. Microbial DNA sequencing and amino sugars biomarker analysis were integrated to probe regulation of microbial traits on microbial necromass C and its contribution to SOC storage under drought. Our results showed that persistent drought increased the contribution of microbial necromass C to SOC with increasing soil depth, ranging from 32.0% in 0–10 cm to 39.7% in 45–60 cm for fungal necromass and from 15.3% in 0–10 cm to 22.6% in 45–60 cm for bacterial one under drought. Interestingly, bacterial necromass C was more important for SOC accumulation than fungal necromass C in subsoil while vice versa in topsoil. Moreover, drought significantly altered the rare microbial diversity and community composition. Drought-induced shifts in rare microbial taxa largely explained the contributions of fungal and bacterial necromass C to SOC in both sub- and top-soils. Thus, microbial traits, especially rare microbial diversity and community composition, regulated the contribution of microbial necromass C to SOC at different soil depths. These findings highlight the crucial role of the rare microbial community on bacterial- and fungal-derived C in subsoil and topsoil, especially under persistent drought, which should be incorporated into land surface models to improve SOC prediction under future climate change.

Growth of European beech recovered faster than that of Norway spruce after a five-year experimental drought in a mixed forest stand

Key MessageBeech growth acclimated better during severe drought and recovered faster than spruce after drought ended. This was associated with a shift in performance along relative tree size towards small trees.The effects of several consecutive drought years and the recovery reaction of mature trees in particular after a long-term drought have been poorly studied so far. In this study, we demonstrate the growth reactions of mature trees during and after a five-year treatment of extended summer droughts, followed by controlled irrigation in a very productive mixed forest stand. We exposed 70-year-old Norway spruce (Picea abies [L.] Karst) and 90-year-old European beech (Fagus sylvatica [L.]) trees to reduced precipitation using automatic throughfall exclusion (TE) roofs during the growing seasons from 2014 to 2018, irrigated the trees in early summer 2019 and removed the roofs thereafter. From 2009 to 2022, we monitored annual tree growth and precipitation on 6 plots with throughfall exclusion and on 6 plots with ambient Control conditions (CO) of the KROOF canopy experiment. Norway spruce lost significant growth during drought, with some trees dying and others remaining at a low growth level without significant recovery from the effects of drought stress. European beech also significantly reduced growth at the beginning of the drought but emerged stronger in growth from the drought than the Control group. Spruce and beech showed a non-significant trend of increased inter-specific growth compared to intra-specific growth during drought. We found that spruce benefitted more from mixture than beech in the recovery phase after drought than during the drought phase itself. Most importantly, we observed a shift in growth performance along the relative tree size towards smaller trees in the TE plots for both species. This change in the relationship between diameter increment and tree size during and after drought is a major finding of our study and suggests a possible response mechanism to prolonged drought. This key observation requires further investigation and should be considered in future forest management strategies under changing climatic conditions.

Structural and compositional acclimation of forests to extended drought: results of the KROOF throughfall exclusion experiment in Norway spruce and European beech

Drought effects on tree growth and mortality are widely studied, but scant knowledge exists on its impact on stand density, size variation, or mixing proportions. Grasping drought's influence on structural and compositional diversity is crucial for stand dynamics, ecosystem services, and silvicultural adaptation. We relied on KROOF, a 5-year throughfall exclusion experiment in a mature Norway spruce (Picea abies) and European beech (Fagus sylvatica) stand, to analyze its impact on structural and compositional attributes, including Stand Density Index (SDI), Growth Dominance Coefficient (GDC), and species mixing proportion. Our study demonstrates that drought-induced growth reduction and tree loss decreased SDI by 27%, mixing proportion by 41% at Norway spruce’s expense, and homogenized stand structure. Furthermore, we reveal that stand density, mixing proportion, and structural diversity were more affected in Norway spruce, stabilizing growth at the stand level. Extended drought significantly altered growth partitioning in favor of smaller trees, with a 70% reduction in growth-size relationship slope and a 157% decrease in GDC. Species-level analysis indicated a stronger partitioning shift towards smaller trees, particularly in Norway spruce. We discuss that longer drought periods may trigger acclimation at tree and stand levels, potentially underestimated when based solely on individual drought years. Sustained stress could induce acclimation across various levels, from the stand to the species cohort, tree, and organ. Maintaining structural and compositional diversity may mitigate future drought stress effects on growth, mortality, and stand structure, as exemplified by the extended experimental drought. We suggest silvicultural approaches better attuned to natural processes amid climate change.

Throughfall exclusion and fertilization effects on tropical dry forest tree plantations, a large-scale experiment

Abstract. Across tropical ecosystems, global environmental change is causing drier climatic conditions and increased nutrient deposition. Such changes represent large uncertainties due to unknown interactions between drought and nutrient availability in controlling ecosystem net primary productivity (NPP). Using a large-scale manipulative experiment, we studied for 4 years whether nutrient availability affects the individual and integrated responses of aboveground and belowground ecosystem processes to throughfall exclusion in 30-year-old mixed plantations of tropical dry forest tree species in Guanacaste, Costa Rica. We used a factorial design with four treatments: control, fertilization (F), drought (D), and drought + fertilization (D + F). While we found that a 13 %–15 % reduction in soil moisture only led to weak effects in the studied ecosystem processes, NPP increased as a function of F and D + F. The relative contribution of each biomass flux to NPP varied depending on the treatment, with woody biomass being more important for F and root biomass for D + F and D. Moreover, the F treatment showed modest increases in maximum canopy cover. Plant functional type (i.e., N fixation or deciduousness) and not the experimental manipulations was the main source of variation in tree growth. Belowground processes also responded to experimental treatments, as we found a decrease in nodulation for F plots and an increase in microbial carbon use efficiency for F and D plots. Our results emphasize that nutrient availability, more so than modest reductions in soil moisture, limits ecosystem processes in tropical dry forests and that soil fertility interactions with other aspects of drought intensity (e.g., vapor pressure deficit) are yet to be explored.

Estimating the response of Himalayan old-growth mountain forests to decreased monsoon precipitation

Forests in the Himalayas are a major carbon store, but are under threat due to changes in precipitation regime. To simulate a precipitation decline, throughfall-exclusion (TFE) shelters were applied during three consecutive monsoon seasons in an oak forest (2.650 m a.s.l.) and a conifer-dominated forest (3.260 m a.s.l.) in central Bhutan. Leaf water potentials, tree mortality, stem increment, soil CO2 efflux, litterfall and fine root dynamics were assessed. TFE significantly and consistently decreased topsoil (0–30 cm) moisture and leaf water potentials of Quercus lanata and Quercus griffithii (lower elevation), and to a lesser extend those of Tsuga dumosa and Quercus semecarpifolia, (higher elevation). TFE did not impose tree mortality. Stem increment remained unaffected until the second TFE year, but showed reductions during the third year with Tsuga dumosa being most severely affected (-60%). Standing fine root biomass stocks were hardly affected by TFE. Increased root necromass and faster fine root growth in the lower elevation forest suggest that the oak trees increased C allocation below ground. Soil CO2 efflux sharply declined during all three TFE years in both forests. Above ground litter input was unaffected by TFE until the second treatment year. Overall, both forest ecosystems appeared highly resistant to the imposed soil drying, with no signs of tree mortality and stable living root biomass stocks.

Soil Respiration Responses to Throughfall Exclusion Are Decoupled From Changes in Soil Moisture for Four Tropical Forests, Suggesting Processes for Ecosystem Models

AbstractClimatic drying is predicted for many tropical forests yet models remain poorly parameterized for these ecosystems, hampering predictions of forest‐climate interactions. We applied an integrated model–experiment approach, parameterizing an ecosystem model with tropical forest observational data and comparing model predictions to a field drying manipulation. We hypothesized that drying suppresses soil CO2 fluxes (i.e., respiration) in relatively dry tropical forests but increases CO2 fluxes in wetter tropical forests by alleviating anaerobiosis. We measured soil CO2 fluxes during wet‐dry cycles from 2015 to 2022 in four Panamanian forests that vary in rainfall and soil fertility. Measured soil CO2 fluxes declined in the dry season and peaked in the early wet season ahead of peak soil moisture, resulting in lower soil moisture optima for respiration than previously modeled. We then parameterized the model using field data and the new moisture‐respiration response functions. The updated model predicted increased soil CO2 fluxes with drying in wetter and fertile forests and suppressed fluxes in drier, infertile forests. In contrast to model predictions, a chronic throughfall exclusion experiment initially suppressed soil respiration across forests, with sustained suppression for four years in the wettest forest only (−28% ± 4% during the dry season). In the fertile forest, drying eventually elevated CO2 fluxes over this period (+75% ± 28% during the late wet season). The unexpected negative drying effect in the wettest, infertile forest could have resulted from reduced vertical flushing of nutrients into soils. Including hydro‐nutrient interactions in ecosystem models could improve predictions of tropical forest‐climate feedbacks.

Assessing the effects of a drought experiment on the reproductive phenology and ecophysiology of a wet tropical rainforest community.

Climate change is expected to increase the intensity and occurrence of drought in tropical regions, potentially affecting the phenology and physiology of tree species. Phenological activity may respond to a drying and warming environment by advancing reproductive timing and/or diminishing the production of flowers and fruits. These changes have the potential to disrupt important ecological processes, with potentially wide-ranging effects on tropical forest function. Here, we analysed the monthly flowering and fruiting phenology of a tree community (337 individuals from 30 species) over 7years in a lowland tropical rainforest in northeastern Australia and its response to a throughfall exclusion drought experiment (TFE) that was carried out from 2016 to 2018 (3years), excluding approximately 30% of rainfall. We further examined the ecophysiological effects of the TFE on the elemental (C:N) and stable isotope (δ13C and δ15N) composition of leaves, and on the stable isotope composition (δ13C and δ18O) of stem wood of four tree species. At the community level, there was no detectable effect of the TFE on flowering activity overall, but there was a significant effect recorded on fruiting and varying responses from the selected species. The reproductive phenology and physiology of the four species examined in detail were largely resistant to impacts of the TFE treatment. One canopy species in the TFE significantly increased in fruiting and flowering activity, whereas one understory species decreased significantly in both. There was a significant interaction between the TFE treatment and season on leaf C:N for two species. Stable isotope responses were also variable among species, indicating species-specific responses to the TFE. Thus, we did not observe consistent patterns in physiological and phenological changes in the tree community within the 3years of TFE treatment examined in this study.

Divergent responses of soil microorganisms to throughfall exclusion across tropical forest soils driven by soil fertility and climate history

Model projections predict tropical forests will experience longer periods of drought and more intense precipitation cycles under a changing climate. Such transitions have implications for structure-function relationships within microbial communities. We examine how throughfall exclusion might reshape prokaryotic and fungal communities across four lowland forests in Panama with a wide variation in mean annual precipitation and soil fertility. Four sites were established across a 1000 mm span in Mean Annual Precipitation (MAP: 2335–3421 mm). We expected microbial communities at sites with lower MAP to be less sensitive to throughfall exclusion than sites with higher MAP and fungal communities to be more resistant to disturbance than prokaryotes. At each location, partial throughfall exclusion structures were established over 10 × 10 m plots to reduce direct precipitation input. After short-term (∼3–9 months) throughfall exclusion, prokaryotic communities showed no change in composition. However, prolonged (12–18 months) throughfall exclusion resulted in divergent prokaryotic community responses, reflecting MAP and soil fertility. We observed the emergence of a “drought microbiome” within infertile sites, whereby the community structure of the experimental throughfall exclusion plots at the lower MAP sites diverged from their respective control sites and converged towards overlapping assemblages. Furthermore, under throughfall exclusion, taxa increasing in relative abundance at the wettest site reflected that endemic to control plots at the lowest MAP site, suggesting a shift toward communities with life-history traits selected for under a lower MAP. By contrast, fungal community composition across sites was resilient to throughfall exclusion; however, biomass diverged in response to throughfall exclusion, increasing at two sites while decreasing in the other two. Broadly, our results suggest that microbial communities' sensitivity to frequent drying and rewetting periods in tropical forest soils will depend on climate history and soil fertility, with infertile sites likely to respond readily to changes in precipitation.

Repeated summer drought changes the radial xylem sap flow profile in mature Norway spruce but not in European beech

Water consumption of trees is one of the most important processes connected to their survival under ongoing climate change and extreme events such as drought. Radial profiles of xylem sap flow density are an integral component to quantify the water transport for the level of an individual tree and that of ecosystems. However, knowledge of such radial profiles, in particular under stress, is very scarce. Here we show the radial profile of the xylem sap flow density in mature European beech (Fagus sylvatica L) and Norway spruce (Picea abies (L) Karst.) under repeated summer drought induced by throughfall exclusion (TE) and subsequent recovery compared to untreated control trees (CO). We measured xylem sap flow density (udaily in L dm−2 d−1) down to 8 cm sapwood depth at breast height using two different approaches, a thermal dissipation system and the heat field deformation method. In beech, repeated throughfall exclusion did not affect the radial xylem sap flow profile. However, in spruce, udaily was strongly reduced across the profile under repeated drought, changing the profile from a linear to a logarithmic regression. Even two years after drought release, the xylem sap flow profile did not fully recover in TE spruce. The reduction of udaily along the radial profile was accompanied by a reduction of the leaf area in TE spruce by c. 50%, while sapwood depth remained constant. The reduction of the xylem sap flow density along the profile reduced the calculated water consumption of TE spruce trees by more than 33% compared to CO trees, also after drought release. The impact of stressors such as repeated drought on the xylem sap flow density across the radial profile and its consequences for trees’ and stands’ water consumption needs to be addressed in more detail to minimize uncertainties in quantifying ecosystem water cycles.

Physiological recovery of tree water relations upon drought release-response of mature beech and spruce after five years of recurrent summer drought.

As climate change progresses, the frequency and duration of drought stress events are increasing. While the mechanisms of drought acclimation of trees has received considerable attention in recent years, the recovery processes remain critically understudied. We used a unique throughfall exclusion experiment in a mature temperate mixed forest consisting of the more isohydric Norway spruce and more anisohydric European beech, to study the recovery and resilience after drought release. We hypothesized that pre-dawn water potential (ΨPD) of both species will increase within 1 day after watering, while the recovery of stomatal conductance (gs) and the reversal of osmoregulation will be significantly delayed in the more isohydric spruce. Furthermore, we hypothesized that the xylem sap flow density (udaily) will not fully recover within the growing season due to the strong drought impact. After 5 years of summer drought, trees showed significantly reduced ΨPD, udaily and increased osmoregulation in leaves, but only isohydric spruce displayed increased leaf abscisic acid concentrations. In line with our hypothesis, ΨPD and gs recovered within 1 day in beech. Conversely, isohydric spruce showed delayed increases in ΨPD and gs. The delay in recovery of spruce was partially related to the replenishment of the stem water reservoir, as indicated by the missing response of udaily at the crown base compared with DBH level upon watering. However, udaily fully recovered only in the next growing season for beech and was still reduced in spruce. Nevertheless, in both species, osmotic acclimations of leaves were reversed within several weeks. While both species displayed full resilience to drought stress in water-related physiology, the recovery time was in several cases, e.g., udaily, ΨPD and gs, shorter for beech than for spruce. With future increases in the frequency of drought events under ongoing climate change, tree species that recover more quickly will be favored.

Forest fluxes and mortality response to drought: model description (ORCHIDEE-CAN-NHA r7236) and evaluation at the Caxiuanã drought experiment

Abstract. Extreme drought events in Amazon forests are expected to become more frequent and more intense with climate change, threatening ecosystem function and carbon balance. Yet large uncertainties exist on the resilience of this ecosystem to drought. A better quantification of tree hydraulics and mortality processes is needed to anticipate future drought effects on Amazon forests. Most state-of-the-art dynamic global vegetation models are relatively poor in their mechanistic description of these complex processes. Here, we implement a mechanistic plant hydraulic module within the ORCHIDEE-CAN-NHA r7236 land surface model to simulate the percentage loss of conductance (PLC) and changes in water storage among organs via a representation of the water potentials and vertical water flows along the continuum from soil to roots, stems and leaves. The model was evaluated against observed seasonal variability in stand-scale sap flow, soil moisture and productivity under both control and drought setups at the Caxiuanã throughfall exclusion field experiment in eastern Amazonia between 2001 and 2008. A relationship between PLC and tree mortality is built in the model from two empirical parameters, the cumulated duration of drought exposure that triggers mortality, and the mortality fraction in each day exceeding the exposure. Our model captures the large biomass drop in the year 2005 observed 4 years after throughfall reduction, and produces comparable annual tree mortality rates with observation over the study period. Our hydraulic architecture module provides promising avenues for future research in assimilating experimental data to parameterize mortality due to drought-induced xylem dysfunction. We also highlight that species-based (isohydric or anisohydric) hydraulic traits should be further tested to generalize the model performance in predicting the drought risks.

Sap flow velocities of Acer saccharum and Quercus velutina during drought: Insights and implications from a throughfall exclusion experiment in West Virginia, USA

Forest species composition mediates evapotranspiration and the amount of water available to human-use downstream. In the last century, the heavily forested Appalachian region has been undergoing forest mesophication which is the progressive replacement of xeric species (e.g. black oak (Quercus velutina)) by mesic species (e.g. sugar maple (Acer saccharum)). Given differences between xeric and mesic species in water use efficiency and rainfall interception losses, investigating the consequences of these species shifts on water cycles is critical to improving predictions of ecosystem responses to climate change. To meet this need, we quantified the degree to which the sap velocities of two dominant broadleaved species (sugar maple and black oak) in West Virginia, responded to ambient and experimentally altered soil moisture conditions using a throughfall exclusion experiment. We then used these data to explore how predictions of future climate under two emissions scenarios could affect forest evapotranspiration rates. Overall, we found that the maples had higher sap velocity rates than the oaks. Sap velocity in maples showed a stronger sensitivity to vapor pressure deficit (VPD), particularly at high levels of VPD, than sap velocity in oaks. Experimentally induced reductions in shallow soil moisture did not have a relevant impact on sap velocity. In response to future climate scenarios of increased vapor pressure deficits in the Central Appalachian Mountains, our results highlight the different degrees to which two important tree species will increase transpiration, and potentially reduce the water available to the heavily populated areas downstream.