Sap Flux Measurements Research Articles

Micromachined needle-like calorimetric flow sensor for sap flux density measurement in the xylem of plants

Miniaturized silicon thermal probes for plant’s sap flow measurement, or micro sap flow sensors, have advantages in minimum invasiveness, low power consumption, and fast responses. Practical applications in sap flow measurement has been demonstrated with the single-probe silicon micro sensors. However, the sensors could not detect flow directions and require estimating zero sap flow output that leads to significant source of uncertainty. Furthermore, silicon-needles would break easily during the insertion into plants. We present the first three-element micro thermal sap flow sensor packaged on a durable printed circuit board needle that can measure bidirectional flows with improved dynamics and precision. The performance of the newly designed calorimetric flow sensor was confirmed through precision calibration and field test on tomato stems. A calibration curve for a tomato stem was obtained with a sensitivity of 0.299 K/(µL mm−2 s−1) under the maximum temperature increase of 4.61 K. Results from the field test for one month revealed a correlation between the measured sap flux density and related conditions such as solar radiation, vapor pressure deficit, sunshade and irrigation. The developed sensor will contribute to practical long-term sap flow monitoring for small and delicate plants with minimal physical invasion.

The influence of climate and management on transpiration of residential trees during a bark beetle infestation

AbstractTrees in residential environments are affected by a unique combination of environmental and anthropogenic factors, including occasional insect outbreaks that are increasing in frequency and severity due to climate change. We studied loblolly pine trees infested by bark beetles in a residential backyard in a southeastern US city. We investigated the responses of tree and stand‐level transpiration to environmental factors (solar radiation, atmospheric vapor pressure deficit, and soil moisture), severe weather events (strong winds and heavy storms), bark beetle infestation, and human actions (insecticide treatments and tree removals). We used constant heat dissipation probes to make continuous sap flux measurements (J0) in tree boles. Over 22 months of the study, J0 of trees with confirmed infestation decreased from ~90 to ~60 g cm−2 day−1 and J0 of the rest of the trees increased from ~60 to ~80 g cm−2 day−1. One infested tree died, as its J0 steadily declined from 110 g cm−2 day−1 to zero over the course of 2 months, followed by a loss of foliage and visible signs of severe infestation 6 months later. J0 was sensitive to variations in incoming solar radiation and atmospheric vapor pressure deficit. In most trees, J0 linearly responded to soil water content during drought periods. Yet despite complex dynamics of J0 variations, plot‐level transpiration at the end of the study was the same as at the beginning due to compensatory increases in tree transpiration rates. This study highlights the intrinsic interplay of environmental, biotic, and anthropogenic factors in residential environments where human actions may directly mediate ecosystem responses to climate.

Modelling of sap flux density of oak in a humid region in China

AbstractTranspiration plays a vital role in determining the watershed water cycle. However, we still have little knowledge of the characteristics of tree transpiration in the Hanjiang River Basin, which is the water source for the middle route of South‐to‐North water diversion project. Here, we measured sap flux density of oak trees (Quercus, the dominant species here) at the 10‐min resolution for 2 years and explored its response to the environmental conditions. The incoming short‐wave radiation and vapour pressure deficit well explained the variation of daytime sap flux density, and a statistical model was then proposed to calculate the daytime sap flux density correspondingly; then a nighttime sap flux density module was proposed based on the daytime sap flux density calculation. Sap flux density showed clear counter‐clockwise hysteresis response to incoming short‐wave radiation, and clockwise hysteresis response to vapour pressure deficit. Our sap flux density model can well reproduce the corresponding hysteresis response to incoming short‐wave radiation and vapour pressure deficit. This study unravelled the environmental controls of sap flux density of the oak trees in the Hanjiang River Basin, proposed an efficient model for the sap flux density simulation, provided important knowledge for understanding the corresponding forests' water use, which is of critical significance in determining the water availability for the middle route of the South‐to‐North water diversion project.

Increased tree water use with the development of a dense understory layer in a North American hardwood forest

AbstractThe formation of a single species, recalcitrant understory vegetation layer can limit tree regeneration and, in the long term, modify the composition of forests. Few studies have investigated how recalcitrant vegetation influences competition for water resources although the formation of a dense understory is likely to modify the forest water balance. In eastern North American hardwood forests, the development of a dense understory layer of American beech (Fagus grandifolia) has been observed in stands dominated by sugar maple (Acer saccharum), a phenomenon that shares many characteristics associated with recalcitrant vegetation. Given that water availability is generally negatively correlated with stand density, we hypothesized that the formation of a dense understory beech layer increased competition for water resources, thus leading to reduced water use by sugar maple trees in beech‐dominated stands. Using thermal dissipation sensors, we measured sap flux density (Fd) of two sugar maple trees at three beech‐dominated sites and three control sites. During the growing season, Fd of sugar maple trees was significantly larger at beech‐dominated sites compared to control sites, indicating a greater rate of water use by sugar maples in stands with a dense understory beech layer. We provide two hypotheses to explain our results at the tree scale: (i) reduced cover by forest floor vegetation could limit transpiration by this layer, thus allowing increased water availability to supply transpiration by overstory trees, or (ii) increased tree transpiration rate could be a mechanism to satisfy nutrient requirements in beech‐dominated stands often associated with lower soil fertility.

Complex canopy structures control tree transpiration: A study based on 3D modelling in a tropical rainforest

AbstractTropical rainforests are rich in tree species and comprise complex canopy structures. Transpiration by forest trees is a major hydrological flux which contributes to climate regulation. We explored the role of forest canopy structure on tree transpiration in a tropical rainforest on Sumatra, Indonesia. Drone‐based photogrammetry and the structure from motion technique were used to compute high‐resolution 3D point clouds and derive forest and tree structural variables. Transpiration of differently sized and vertically positioned trees was assessed with sap flux measurements. Per‐tree transpiration rates increased linearly with several variables related to crown dimension and were further enhanced by top‐heavy crown shape, dense leafage and increasing canopy openness as assessed with the gap light index. Under given environmental conditions, the two variables crown volume and top‐heaviness explained 74% of the observed variance in per‐tree transpiration. Transpiration rates per unit crown dimension were highest for small crowns, decreased non‐linearly with increasing crown dimension and were little affected by other analysed structural variables. Our study underlines the potential of 3D point cloud analyses for accurately determining the structure of complex forest canopies and thus better understanding and predicting transpiration rates of differently positioned trees.

Sap flow and leaf gas exchange response to a drought and heatwave in urban green spaces in a Nordic city

Abstract. Urban vegetation plays a role in offsetting urban CO2 emissions, mitigating heat through tree transpiration and shading, and acting as deposition surfaces for pollutants. The frequent occurrence of heatwaves and of concurrent drought conditions significantly disrupts the processes of urban trees, particularly their photosynthesis and transpiration rates. Despite the pivotal role of urban tree functioning in delivering essential ecosystem services, the precise nature of their response remains uncertain. We conducted sap flux density (Js) and leaf gas exchange measurements of four tree species (Tilia cordata, Tilia × europaea, Betula pendula, and Malus spp.) located in different urban green areas (Park, Street, Forest, and Orchard) in Helsinki, Finland. Measurements were made over two contrasting summers 2020 and 2021. Summer 2021 experienced a local heatwave and drought, whereas summer 2020 was more typical of Helsinki. In this study, we aimed to understand the responses of urban tree transpiration (measured with sap flux density) and leaf gas exchange to heatwave and drought conditions, and we examined the main environmental drivers controlling the tree transpiration rate during these periods. We observed varying responses of Js during the heatwave period at the four urban sites. When comparing the heatwave and no heatwave periods, a 35 %–67 % increase in Js was observed at the Park, Forest, and Orchard locations, whereas no significant change was seen at the Street site. Our results also showed that Js was higher (31 %–63 %) at all sites under drought conditions compared with non-dry periods. The higher Js values during the heatwave and dry periods were mainly driven by the high atmospheric demand for evapotranspiration, represented by the high vapor pressure deficit (VPD), suggesting that the trees were not experiencing severe enough heat or drought stress that stomatal control would have decreased transpiration. Accordingly, photosynthetic potential (Amax), stomatal conductance (gs), and transpiration (E) at the leaf level did not change during heatwave and drought periods, excluding the Park site where a significant reduction in gs was seen. VPD explained 55 %–69 % of the variation in the daily mean Js during heatwave and drought periods at all sites. At the Forest site, the increase in Js saturated after a certain VPD level, likely due to low soil water availability during these hot and dry periods. Overall, the heat and drought conditions were untypical of the region but not excessive enough to restrict stomatal control and transpiration, indicating that ecosystem services such as cooling were not at risk.

Sap flux and stable isotopes of water show contrasting tree water uptake strategies in two co‐occurring tropical rainforest tree species

AbstractThe short‐term dynamics of tree water use strategies for neighbouring co‐occurring species are poorly understood. Here, we quantify the high frequency changes in water sources and sap flux patterns of two commonly co‐occurring tropical rainforest tree species: Dendrocnide photinophylla (Kunth; Chew) and Argyrodendron peralatum (F.M. Bailey; Edlin ex J.H. Boas). A combination of continuous sap flux measurements and hourly sampling of xylem water stable isotope composition (δD and δ18O) were used to observe water use strategies during a 24‐h transpiration cycle. Sap flux ranged between 2.82 and 28.50 L day−1, with A. peralatum recording a 67% higher rate than D. photinophylla. For both tree species, sap flux increased with tree size and diurnal sap flux increase resulted in more isotopically enriched xylem water. A Bayesian Mixing Model analysis, which used sampled soil water isotopic composition from five soil depths ranging from of 0 to 1 m, revealed that D. photinophylla primarily used water from very shallow depth or soil surface layer (2–60 cm), while A. peralatum sourced its water mostly from deeper layers (60–100 cm). We propose that these differences in species' water consumption patterns are related to plant water storage capacity and wood anatomical features. This research demonstrates that combining xylem isotope composition and sap flux measurements can help reveal species‐specific water use strategies, which can be beneficial for improved process understanding in ecohydrological modelling.

UAV-based thermography reveals spatial and temporal variability of evapotranspiration from a tropical rainforest

Evapotranspiration (ET) from tropical forests plays a significant role in regulating the climate system. Forests are diverse ecosystems, encompass heterogeneous site conditions and experience seasonal fluctuations of rainfall. Our objectives were to quantify ET from a tropical rainforest using high-resolution thermal images and a simple modeling framework. In lowland Sumatra, thermal infrared (TIR) images were taken from an uncrewed aerial vehicle (UAV) of upland and riparian sites during both dry and wet seasons. We predicted ET from land surface temperature data retrieved from the TIR images by applying the DATTUTDUT energy balance model. We further compared the ET estimates to ground-based sap flux measurements for selected trees and assessed the plot-level spatial and temporal variability of ET across sites and seasons. Average ET across sites and seasons was 0.48 mm h–1, which is comparable to ET from a nearby commercial oil palm plantation where this method has been validated against eddy covariance measurements. For given trees, a positive correlation was found between UAV-based ET and tree transpiration derived from ground-based sap flux measurements, thereby corroborating the observed spatial patterns. Evapotranspiration at upland sites was 11% higher than at riparian sites across all seasons. The heterogeneity of ET was lower at upland sites than at riparian sites, and increased from the dry season to the wet season. This seasonally enhanced ET variability can be an effect of local site conditions including partial flooding and diverse responses of tree species to moisture conditions. These results improve our understanding of forest-water interactions in tropical forests and can aid the further development of vegetation-atmosphere models. Further, we found that UAV-based thermography using a simple, energy balance modeling scheme is a promising method for ET assessments of natural (forest) ecosystems, notably in data scarce regions of the world.

Role Of Hydraulic Traits On Stomatal Regulation Of Transpiration Under Different Vapor Pressure Deficits Across Five Mediterranean Tree Crops.

The differential stomatal regulation of transpiration among plant species in response to water deficit is not fully understood, although several hydraulic traits have been reported to influence it. This knowledge gap is partly due to a lack of direct and concomitant experimental data of transpiration, stomatal conductance and hydraulic traits. We measured sap flux density (Js), stomatal conductance (gs) and different hydraulic traits in five crop species. We aim to contribute to establish the causal relationship between water consumption and its regulation using a hydraulic trait-based approach. The results showed that the species-specific regulation of Js by gs was overall coordinated with the functional hydraulic traits analyzed. Especially relevant was the negative and significant relationship found between the Huber value (Hv) and its functional analogue ratio between maximum Js and gs (Jsmax/gsmax) which can be understood as a compensation to maintain the hydraulic supply to the leaves. Hv was also significantly related to the slope of the relationship between gs and Js response to vapor pressure deficit and explained most of its variability, adding up to evidence recognizing Hv as a major trait in plant water relations. Thus, hydraulic basis for the regulation of tree water use should be considered.

Trees with anisohydric behavior as main drivers of nocturnal evapotranspiration in a tropical mountain rainforest.

This study addresses transpiration in a tropical evergreen mountain forest in the Ecuadorian Andes from the leaf to the stand level, with emphasis on nocturnal plant-water relations. The stand level: Evapotranspiration (ET) measured over 12 months with the Eddy-Covariance (ECov) technique proved as the major share (79%) of water received from precipitation. Irrespective of the humid climate, the vegetation transpired day and night. On average, 15.3% of the total daily ET were due to nocturnal transpiration. Short spells of drought increased daily ET, mainly by enhanced nighttime transpiration. Following leaf transpiration rather than air temperature and atmospheric water vapor deficit, ET showed its maximum already in the morning hours. The tree level: Due to the humid climate, the total water consumption of trees was generally low. Nevertheless, xylem sap flux measurements separated the investigated tree species into a group showing relatively high and another one with low sap flux rates. The leaf level: Transpiration rates of Tapirira guianensis, a member of the high-flux-rate group, were more than twice those of Ocotea aciphylla, a representative of the group showing low sap flux rates. Representatives of the Tapirira group operated at a relatively high leaf water potential but with a considerable diurnal amplitude, while the leaves of the Ocotea group showed low water potential and small diurnal fluctuations. Overall, the Tapirira group performed anisohydrically and the Ocotea group isohydrically. Grouping of the tree species by their water relations complied with the extents of the diurnal stem circumference fluctuations. Nighttime transpiration and hydrological type: In contrast to the isohydrically performing trees of the Ocotea group, the anisohydric trees showed considerable water vapour pressure deficit (VPD)-dependent nocturnal transpiration. Therefore, we conclude that nighttime ET at the forest level is mainly sourced by the tree species with anisohydric performance.

Tree variability limits the detection of nutrient treatment effects on sap flux density in a northern hardwood forest.

The influence of nutrient availability on transpiration is not well understood, in spite of the importance of transpiration to forest water budgets. Soil nutrients have the potential to affect tree water use through indirect effects on leaf area or stomatal conductance. For example, following addition of calcium silicate to a watershed at Hubbard Brook, in New Hampshire, streamflow was reduced for 3 years, which was attributed to a 25% increase in evapotranspiration associated with increased foliar production. The first objective of this study was to quantify the effect of nutrient availability on sap flux density in a nitrogen, phosphorus, and calcium addition experiment in New Hampshire in which tree diameter growth, foliar chemistry, and soil nutrient availability had responded to treatments. We measured sap flux density in American beech (Fagus grandifolia, Ehr.), red maple (Acer rubrum L.), sugar maple (Acer saccharum Marsh.), white birch (Betula papyrifera Marsh.), or yellow birch (Betula alleghaniensis Britton.) trees, over five years of experiments in five stands distributed across three sites. In 2018, 3 years after a calcium silicate addition, sap flux density averaged 36% higher in trees in the treatment than the control plot, but this effect was not very significant (p = 0.07). Our second objective was to determine whether this failure to detect effects with greater statistical confidence was due to small effect sizes or high variability among trees. We found that tree-to-tree variability was high, with coefficients of variation averaging 39% within treatment plots. Depending on the species and year of the study, the minimum difference in sap flux density detectable with our observed variability ranged from 46% to 352%, for a simple ANOVA. We analyzed other studies reported in the literature that compared tree water use among species or treatments and found detectable differences ranging from 16% to 78%. Future sap flux density studies could benefit from power analyses to guide sampling intensity. Including pretreatment data, in the case of manipulative studies, would also increase statistical power.

Apple tree transpiration during cycles of progressive drought as assessed via continuous gravimetric and xylem sap flux measurements

Apple tree transpiration during cycles of progressive drought as assessed via continuous gravimetric and xylem sap flux measurements

A low cost, low power sap flux device for distributed and intensive monitoring of tree transpiration

Accurate estimation of transpiration in individual trees is important for understanding plant responses to environmental drivers, closing the water balance in forest stands and catchments, and calibrating earth system models, among other applications. However, the cost and power consumption of commercial systems based on sap flow methods still limit their usage. We developed and tested a cost-effective (<$150), simple to construct, and energy efficient sap flux device based on the heat pulse method. Energy savings were achieved by reducing the voltage of heat pulses and using an internal clock to completely shut down the device between pulses. Device accuracy was confirmed by laboratory estimates of sap flow made on excised branches of Acer saccharum and Tsuga canadensis (adjusted R2 = 0.96). In a 174-d field installation of 12 devices, batteries (eight rechargeable Ni-MH AA) needed to be replaced every 14 days. Sap flux measurements in the field tracked expected variations in vapor pressure deficit and tree phenology. The low cost, compact design, reliability, and power consumption of this device enable sap flux studies to operate with more replication and in more diverse ecological settings than has been practical in the past.

An in situ real time probe spacing correction method for multi-needle heat pulse sap flow sensors

Probe spacing of sap flow sensors affects the quantification of sap flow in plants. The assumption of nocturnal zero flow conditions, required by most non-destructive methods for probe spacing calibration, is often invalid in natural biological systems and extremely hard to validate with in situ measurements. Our study aims to present a novel method for accurate in situ determination of probe spacing. The method uses nonlinear curve fitting with two mathematical models applied during low flow conditions. Numerical simulations were conducted with three types of probe misalignment and demonstrated that errors in heat pulse velocity (Vh) (< 2.0%) is less than the error associated with commonly-used probe spacing correction methods. Field experiment on European beech (Fagus sylvatica L.) comparisons with corrected probe spacing applied to the heat ratio (HR) and Sapflow+ method produced improved sap flux densities, with both methods agreeing very well (root mean square deviation < 1 cm3 cm−2 h−1). The proposed method allows real time probe spacing measurements at low flow rates, alleviating some uncertainties associated with zero flow assumption. The new method can hence successfully be used to estimate in situ probe spacing for operational measurements of sap flux density.

Assessing water requirement of orange trees using sap flow measurements in Narkhed-Pandhurna critical zone observatory (CZO) in central India

&#x0D; We used thermal dissipation method for sap flux measurements in orange trees to assess its water requirement in Narkhed-Pandhurna region. Thermal Dissipation Probe (TDP) sensors were installed in 5-year old (young) and 15-year old (mature) orange trees to measure the diurnal sap flux variations in trees during November 21, 2019, to January 31, 2020 (71 days). The results show that the maximum daily water uptake by the 5-year old tree was 1.1 L observed on 39th day of measurement (December 29, 2019) and in the 15-year old tree it was 5.0 L, and it observed on 38th day (December 28, 2019) of measurement. The cumulative water uptake during the study period by the 5-year old tree was 49.0 L, and the 15-year old tree consumed 257.4 L of water. The results were compared with the recommended irrigational values of Indian Horticulture Board (IHB), Government of India (GoI) and Groundwater Survey and Development Agency (GSDA), Government of Maharashtra (GoMH) for orange orchards. The initial investigation shows that recommended guidelines for irrigation of orange trees are exorbitantly high and needs to be revised. The sap flow methods are more precise that can measure sap flow at a very short interval and can generate a time series of data. It can be used to revise the guidelines with the aim to conserve water and propose precision water irrigation for the study area in particular and different agro-climatic zones of the country in general.&#x0D;

Measurement of low sap flux density in plants using the single needle heat pulse probe

Heat pulse probes are used worldwide to quantify sap flow in plants. Probes consisting of a single needle inserted into a plant have a simple physical configuration relative to multi-probe methods. The single probe also minimizes physical damage to plant tissue. Single probe methods are therefore increasingly used for sap flux density measurements. However, at low sap flux density values, existing single probe heat pulse (SPHP) methods cannot be used for accurate measurements. To increase the sensitivity of SPHP to low sap flux densities, this paper describes and validates a simplified expression of sap flow and heat transport under various heat inputs and finite heat pulse time intervals. To test this theoretical development, measurements were collected from SPHP probes inserted into mature poplar forest trees (Populus tomentosa Carrière) during the 2020 growing season. The results show that the new method can successfully monitor low sap flux density: the lowest measured heat pulse velocity was 3 cm h−1, which is an improvement from the original SPHP method (20 cm h−1). The improved SPHP method had a root mean square deviation (RMSD) of 1.63 cm h−1 in comparison with the heat ratio (HR) method. The proposed SPHP method is therefore suitable for quantifying both low and high sap flux densities.

Disentangling the Effects of Vapor Pressure Deficit and Soil Water Availability on Canopy Conductance in a Seasonal Tropical Forest During the 2015 El Niño Drought

AbstractWater deficit in the atmosphere and soil are two key interactive factors that constrain transpiration and vegetation productivity. It is not clear which of these two factors is more important for the water and carbon flux response to drought stress in ecosystems. In this study, field data and numerical modeling were used to isolate their impact on evapotranspiration (ET) and gross primary productivity (GPP) at a tropical forest site in Barro Colorado Island (BCI), Panama, focusing on their response to the drought induced by the El Niño event of 2015–2016. Numerical simulations were performed using a plant hydrodynamic scheme (HYDRO) and a heuristic approach that ignores stomatal sensitivity to leaf water potential in the Energy Exascale Earth System Model (E3SM) Land Model (ELM). The sensitivity of canopy conductance (Gs) to vapor pressure deficit (VPD) obtained from eddy‐covariance fluxes and measured sap flux shows that, at both ecosystem and plant scale, soil water stress is more important in limiting Gs than VPD at BCI during the El Niño event. The model simulations confirmed the importance of water stress limitation on Gs, but overestimated the VPD impact on Gs compared to that estimated from the observations. We also found that the predicted soil moisture is less sensitive to the diversity of plant hydraulic traits than ET and GPP. During the dry season at BCI, seasonal ET, especially soil evaporation at VPD &gt; 0.42 kPa, simulated using HYDRO and ELM, were too strong and will require alternative parameterizations.

Radial variations in xylem sap flux in a temperate red pine plantation forest

BackgroundScaling sap flux measurements to whole-tree water use or stand-level transpiration is often done using measurements conducted at a single point in the sapwood of the tree and has the potential to cause significant errors. Previous studies have shown that much of this uncertainty is related to (i) measurement of sapwood area and (ii) variations in sap flow at different depths within the tree sapwood.ResultsThis study measured sap flux density at three depth intervals in the sapwood of 88-year-old red pine (Pinus resinosa) trees to more accurately estimate water-use at the tree- and stand-level in a plantation forest near Lake Erie in Southern Ontario, Canada. Results showed that most of the water transport (65%) occurred in the outermost sapwood, while only 26% and 9% of water was transported in the middle and innermost depths of sapwood, respectively.ConclusionsThese results suggest that failing to consider radial variations in sap flux density within trees can lead to an overestimation of transpiration by as much as 81%, which may cause large uncertainties in water budgets at the ecosystem and catchment scale. This study will help to improve our understanding of water use dynamics and reduce uncertainties in sap flow measurements in the temperate pine forest ecosystems in the Great Lakes region and help in protecting these forests in the face of climate change.

Water Use by Chinese Pine Is Less Conservative but More Closely Regulated Than in Mongolian Scots Pine in a Plantation Forest, on Sandy Soil, in a Semi-Arid Climate.

The diversity of plant water use patterns among species and ecosystems is a matter of widespread debate. In this study, Chinese pine (Pinus tabuliformis, CP) and Mongolian Scots pine (Pinus sylvestris var. mongolica, MP), which is co-exist in the shelterbelt plantations in the Horqin Sandyland in northern China, were chosen for comparison of water use traits by monitoring xylem sap flow alongside recordings of the associated environmental factors over four growing seasons. Continuous sap flux density measurements were converted into crown projected area transpiration intensity (Tr) and canopy stomatal conductance (Gs). The results indicated that MP showed a higher canopy transpiration intensity than in CP, with Tr daily means (±standard deviation) of 0.84 ± 0.36 and 0.79 ± 0.43 mm⋅d–1, respectively (p = 0.07). However, the inter-annual variability of daily Tr in MP was not significant, varying only approximately a 1.1-fold (p = 0.29), while inter-annual variation was significant for CP, with 1.24-fold variation (p < 0.01). In particular, the daily mean Tr value for CP was approximately 1.7-times higher than that of MP under favorable soil moisture conditions, with values for relative extractable soil water within the 0–1.0 m soil layer (REW) being above 0.4. However, as the soil dried out, the value of Tr for CP decreased more sharply, falling to only approximately 0.5-times the value for MP when REW fell to < 0.2. The stronger sensitivity of Tr and/or Gs to REW, together with the more sensitive response of Gs to VPD in CP, confirms that CP exhibits less conservation of soil water utilization but features a stronger ability to regulate water use. Compared with MP, CP can better adapt to the dry conditions associated with climate change.

Comparative drought sensitivity of co‐occurring white spruce and paper birch in interior Alaska

Abstract Numerous recent studies have argued that moisture limitation is leading to growth declines and mortality of black spruce (Picea mariana) and white spruce (Picea glauca) in the North American boreal forest. A parallel line of research suggests that increasingly common severe wildfires are altering successional pathways and leading to long‐term replacement of spruce forests with those dominated by paper birch (Betula papyrifera, Betula neoalaskana) and trembling aspen (Populus tremuloides). When both conifers and deciduous trees establish after fire, this biome shift hypothesis implicitly assumes that deciduous species will outcompete the conifers, owing to their more rapid vertical growth and because they might be less sensitive to warm and dry conditions. We established a research site in a white spruce‐paper birch forest on an east‐facing slope in interior Alaska and tested the hypothesis that Alaska paper birch are better adapted to warm and dry conditions than white spruce. Over 6 years (2013–2018), we made hourly measurements of microclimate and xylem sap flux of both species. We also collected increment cores and conducted climate‐growth analyses for both species. During our 6‐year study, growing seasons with low volumetric soil water content (SWC) were those that followed shallow winter snowpacks and had limited summer rainfall, not necessarily those with warm air temperature. Both species were sensitive to moisture limitation. The tree‐ring data revealed significant positive effects of cumulative water year precipitation on radial growth, with a steeper slope for paper birch than for white spruce. Radial growth of both species was also positively related to mean water year air temperature. Sap flux density declined progressively for white spruce over the range of observed SWC and abruptly for paper birch when SWC fell below ~15%. Synthesis. Our results show that, while paper birch might be less sensitive to mild drought than white spruce, it may be more sensitive to severe drought, raising questions about the ability of paper birch to outcompete co‐occurring white spruce in a drier climate.