Soil Heating in Fire (SheFire): A model and measurement method for estimating soil heating and effects during wildland fires.

Fire has transformative effects on soil biological, chemical, and physical properties in terrestrial ecosystems around the world. While methods for estimating fire characteristics and associated effects aboveground have progressed in recent decades, there remain major challenges in characterizing soil heating and associated effects belowground. Overcoming these challenges is crucial for understanding how fire influences soil carbon storage, biogeochemical cycling, and ecosystem recovery. In this paper, we present a novel framework for characterizing belowground heating and effects. The framework includes (1) an open-source model to estimate fire-driven soil heating, cooling, and the biotic effects of heating across depths and over time (Soil Heating in Fire model; SheFire) and (2) a simple field method for recording soil temperatures at multiple depths using self-contained temperature sensor and data loggers (i.e., iButtons), installed along a wooden stake inserted into the soil (i.e., an iStake). The iStake overcomes many logistical challenges associated with obtaining temperature profiles using thermocouples. Heating measurements provide inputs to the SheFire model, and modeled soil heating can then be used to derive ecosystem response functions, such as heating effects on microorganisms and tissues. To validate SheFire estimates, we conducted a burn table experiment using iStakes to record temperatures that were in turn used to fit the SheFire model. We then compared SheFire predicted temperatures against measured temperatures at other soil depths. To benchmark iStake measurements against those recorded by thermocouples, we co-located both types of sensors in the burn table experiment. We found that SheFire demonstrated skill in interpolating and extrapolating soil temperatures, with the largest errors occurring at the shallowest depths. We also found that iButton sensors are comparable to thermocouples for recording soil temperatures during fires. Finally, we present a case study using iStakes and SheFire to estimate in situ soil heating during a prescribed fire and demonstrate how observed heating regimes would influence seed and tree root vascular cambium survival at different soil depths. This measurement-modeling framework provides a cutting-edge approach for describing soil temperature regimes (i.e., soil heating) through a soil profile and predicting biological responses.

Prosopis glandulosa persistence is facilitated by differential protection of buds during low- and high-energy fires.

Rangelands worldwide have experienced significant shifts from grass-dominated to woody-plant dominated states over the past century. In North America, these shifts are largely driven by overgrazing and landscape-scale fire suppression. Such shifts reduce productivity for livestock, can have broad-scale impacts to biodiversity, and are often difficult to reverse. Restoring grass dominance often involves restoring fire as an ecological process. However, many resprouting woody plants persist following disturbance, including fire, by resprouting from protected buds, rendering fire ineffective for reducing resprouting woody plant density. Recent research has shown that extreme fire (high-energy fires during periods of water stress) may reduce resprouting capacity. This previous research did not examine whether high-energy fires alone would be sufficient to cause mortality. We created an experimental framework for assessing the "buds-protection-resources" hypothesis of resprouting persistence under different fire energies. In July-August 2018 we exposed 48 individuals of a dominant resprouting woody plant in the region, honey mesquite (Prosopis glandulosa), to two levels of fire energy (high and low) and root crown exposure (exposed vs unexposed) and evaluated resprouting capacity. We censused basal and epicormic resprouts for two years following treatment. Water stress was moderate for several months leading up to fires but low in subsequent years. Epicormic and basal buds were somewhat protected from low- and high-energy fire. However, epicormic buds were protected in very few mesquites subjected to high-energy fires. High-energy fires decreased survival, caused loss of apical dominance, and left residual dead stems, which may increase chances of mortality from future fires. Basal resprout numbers were reduced by high-energy fires, which may have additional implications for long-term mesquite survival. While the buds, protection, and resources components of resprouter persistence all played a role in resprouting, high-energy fire decreased mesquite survival and reduced resprouting. This suggests that high-energy fires affect persistence mechanisms to different extents than low-energy fires. In addition, high-energy fires during normal rainfall can have negative impacts on resprouting capacity; water stress is not a necessary precursor to honey mesquite mortality from high-energy fire.

Grass bud responses to fire in a semiarid savanna system.

Increasingly, land managers have attempted to use extreme prescribed fire as a method to address woody plant encroachment in savanna ecosystems. The effect that these fires have on herbaceous vegetation is poorly understood. We experimentally examined immediate (<24 hr) bud response of two dominant graminoids, a C3 caespitose grass, Nassella leucotricha, and a C4 stoloniferous grass, Hilaria belangeri, following fires of varying energy (J/m2) in a semiarid savanna in the Edwards Plateau ecoregion of Texas. Treatments included high- and low-energy fires determined by contrasting fuel loading and a no burn (control) treatment. Belowground axillary buds were counted and their activities classified to determine immediate effects of fire energy on bud activity, dormancy, and mortality. High-energy burns resulted in immediate mortality of N. leucotricha and H. belangeri buds (p < .05). Active buds decreased following high-energy and low-energy burns for both species (p < .05). In contrast, bud activity, dormancy, and mortality remained constant in the control. In the high-energy treatment, 100% (n = 24) of N. leucotricha individuals resprouted while only 25% (n = 24) of H. belangeri individuals resprouted (p < .0001) 3 weeks following treatment application. Bud depths differed between species and may account for this divergence, with average bud depths for N. leucotricha 1.3 cm deeper than H. belangeri (p < .0001). Synthesis and applications: Our results suggest that fire energy directly affects bud activity and mortality through soil heating for these two species. It is imperative to understand how fire energy impacts the bud banks of grasses to better predict grass response to increased use of extreme prescribed fire in land management.

Comparing the influence of wildfire and prescribed burns on watershed nitrogen biogeochemistry using 15N natural abundance in terrestrial and aquatic ecosystem components.

We evaluated differences in the effects of three low-severity spring prescribed burns and four wildfires on nitrogen (N) biogeochemistry in Rocky Mountain headwater watersheds. We compared paired (burned/unburned) watersheds of four wildfires and three spring prescribed burns for three growing seasons post-fire. To better understand fire effects on the entire watershed ecosystem, we measured N concentrations and δ15N in both the terrestrial and aquatic ecosystems components, i.e., soil, understory plants in upland and riparian areas, streamwater, and in-stream moss. In addition, we measured nitrate reductase activity in foliage of Spiraea betulifolia, a dominant understory species. We found increases of δ15N and N concentrations in both terrestrial and aquatic ecosystem N pools after wildfire, but responses were limited to terrestrial N pools after prescribed burns indicating that N transfer from terrestrial to aquatic ecosystem components did not occur in low-severity prescribed burns. Foliar δ15N differed between wildfire and prescribed burn sites; the δ15N of foliage of upland plants was enriched by 2.9 ‰ (difference between burned and unburned watersheds) in the first two years after wildfire, but only 1.3 ‰ after prescribed burns. In-stream moss δ15N in wildfire-burned watersheds was enriched by 1.3 ‰, but there was no response by moss in prescription-burned watersheds, mirroring patterns of streamwater nitrate concentrations. S. betulifolia showed significantly higher nitrate reductase activity two years after wildfires relative to corresponding unburned watersheds, but no such difference was found after prescribed burns. These responses are consistent with less altered N biogeochemistry after prescribed burns relative to wildfire. We concluded that δ15N values in terrestrial and aquatic plants and streamwater nitrate concentrations after fire can be useful indicators of the magnitude and duration of fire effects and the fate of post-fire available N.

Constraining 3-PG with a new δ13C submodel: a test using the δ13C of tree rings.

A semi-mechanistic forest growth model, 3-PG (Physiological Principles Predicting Growth), was extended to calculate δ(13)C in tree rings. The δ(13)C estimates were based on the model's existing description of carbon assimilation and canopy conductance. The model was tested in two ~80-year-old natural stands of Abies grandis (grand fir) in northern Idaho. We used as many independent measurements as possible to parameterize the model. Measured parameters included quantum yield, specific leaf area, soil water content and litterfall rate. Predictions were compared with measurements of transpiration by sap flux, stem biomass, tree diameter growth, leaf area index and δ(13)C. Sensitivity analysis showed that the model's predictions of δ(13)C were sensitive to key parameters controlling carbon assimilation and canopy conductance, which would have allowed it to fail had the model been parameterized or programmed incorrectly. Instead, the simulated δ(13)C of tree rings was no different from measurements (P > 0.05). The δ(13)C submodel provides a convenient means of constraining parameter space and avoiding model artefacts. This δ(13)C test may be applied to any forest growth model that includes realistic simulations of carbon assimilation and transpiration.

Nocturnal transpiration causing disequilibrium between soil and stem predawn water potential in mixed conifer forests of Idaho.

Soil water potential (Psi(s)) is often estimated by measuring leaf water potential before dawn (Psi(pd)), based on the assumption that the plant water status has come into equilibrium with that of the soil. However, it has been documented for a number of plant species that stomata do not close completely at night, allowing for nocturnal transpiration and thus preventing nocturnal soil-plant water potential equilibration. The potential for nighttime transpiration necessitates testing the assumption of nocturnal equilibration before accepting Psi(pd) as a valid estimate of Psi(s). We determined the magnitude of disequilibrium between Psi(pd) and Psi(s) in four temperate conifer species across three height classes through a replicated study in northern Idaho. Based on both stomatal conductance and sap flux measurements, we confirmed that the combination of open stomata and high nocturnal atmospheric vapor pressure deficit (D) resulted in nocturnal transpiration in all four species. Nocturnal stomatal conductance (g(s-noc)) averaged about 33% of mid-morning conductance values. We used species-specific estimates of g(s-noc) and leaf specific conductance to correct Psi(pd) values for nocturnal transpiration at the time the samples were collected. Compared with the unadjusted values, corrected values reflected a significantly higher Psi(pd) (when D > 0.12 kPa). These results demonstrate that comparisons of Psi(pd) among species, canopy height classes and sites, and across growing seasons can be influenced by differential amounts of nocturnal transpiration, leading to flawed results. Consequently, it is important to account for the presence of nocturnal transpiration, either through a properly parameterized model or by making Psi(pd) measurements when D is sufficiently low that it cannot drive nocturnal transpiration. Violating these conditions will likely result in underestimation of Psi(s).

Cold air drainage and modeled nocturnal leaf water potential in complex forested terrain.

Spatial variation in microclimate caused by air temperature inversions plays an important role in determining the timing and rate of many physical and biophysical processes. Such phenomena are of particular interest in mountainous regions where complex physiographic terrain can greatly complicate these processes. Recent work has demonstrated that, in some plants, stomata do not close completely at night, resulting in nocturnal transpiration. The following work was undertaken to develop a better understanding of nocturnal cold air drainage and its subsequent impact on the reliability of predawn leaf water potential (Psi(pd)) as a surrogate for soil water potential (Psi(s)). Eight temperature data loggers were installed on a transect spanning a vertical distance of 155 m along a north facing slope in the Mica Creek Experimental Watershed (MCEW) in northern Idaho during July and August 2004. Results indicated strong nocturnal temperature inversions occurring from the low- to upper-mid-slope, typically spanning the lower 88 m of the vertical distance. Based on mean temperatures for both months, inversions resulted in lapse rates of 29.0, 27.0 and 25.0 degrees C km(-1) at 0000, 0400 and 2000 h, respectively. At this scale (i.e., < 1 km), the observed lapse rates resulted in highly variable nighttime vapor pressure deficits (D) over the length of the slope, with variable impacts on modeled disequilibrium between soil and leaf water potential. As a result of cold air drainage, modeled Psi(pd) became consistently more negative (up to -0.3 MPa) at higher elevations during the night based on mean temperatures. Nocturnal inversions on the lower- and mid-slopes resulted in leaf water potentials that were at least 30 and 50% more negative over the lower 88 m of the inversion layer, based on mean and maximum temperatures, respectively. However, on a cloudy night, with low D, the maximum decrease in Psi(pd) was -0.04 MPa. Our results indicate that, given persistent cold air drainage and nighttime stomatal opening, serious errors will result if Psi(s) is estimated from Psi(pd).

Growth response of Douglas-fir seedlings to nitrogen fertilization: importance of Rubisco activation state and respiration rates.

High foliar nitrogen concentration ([N]) is associated with high rates of photosynthesis and thus high tree productivity; however, at excessive [N], tree productivity is reduced. Reports of excessive [N] in the Douglas-fir forests of the Oregon Coast Range prompted this investigation of growth and needle physiological responses to increasing foliar N concentrations in 1-year-old Douglas-fir seedlings. After 1 year of N fertilization, total seedling biomass increased with each successive increase in N fertilizer concentration, except in the highest N fertilization treatment. Of the many physiological responses that were analyzed, only photosynthetic capacity (i.e., Vcmax), respiration rates and leaf specific conductance (KL) differed significantly between N treatments. Photosynthetic capacity showed a curvilinear relationship with foliar [N], reaching an apparent maximum rate when needle N concentrations exceeded about 12 mg g(-1). In vitro measurements of ribulose-1,5-bisphosphate carboxylase (Rubisco) activity suggested that photosynthetic capacity was best related to activated, not total, Rubisco content. Rubisco activation state declined as foliar [N] increased, and based on its significant correlation (r2= 0.63) with foliar Mn:Mg ratios, it may be related to Mn inactivation of Rubisco. Respiration rates increased linearly as foliar N concentration increased (r2= 0.84). The value of K(L) also increased as foliar [N] increased, reaching a maximum when foliar [N] exceeded about 10 mg g(-1). Changes in K(L) were unrelated to changes in leaf area or sapwood area because leaf area to sapwood area ratios remained constant. Cumulative effects of the observed physiological responses to N fertilization were analyzed by modeling annual net CO2 assimilation (Anet) based on treatment specific values of Vcmax, dark respiration (Rdark) and KL. Estimates of Anet were highly correlated with measured total seedling biomass (r2= 0.992), suggesting that long-term, cumulative effects of maximum Rubisco carboxylation, Rdark and KL responses to N fertilization may limit seedling production when foliar N exceeds about 13 mg g(-1) or is reduced to less than about 11 mg g(-1).

Pseudothecia of Swiss needle cast fungus, Phaeocryptopus gaeumannii, physically block stomata of Douglas fir, reducing CO2 assimilation.

The following study investigates the timing and mechanism of impact of Swiss needle cast on Douglas-fir (Pseudotsuga menziesii) needle physiology (i.e. gas exchange). Swiss needle cast is a foliar disease caused by the fungus Phaeocryptopus gaeumannii, which occurs throughout the range of Douglas fir and until recently has been considered unimportant. However, recent surveys show the Swiss needle cast currently affects > 52611 ha of forested lands in western Oregon, USA, causing a reduction in growth of c. 23% or an implied growth loss of c. 3.2 m3 ha-1 yr-1 for 1996 alone. Gas exchange of artificially inoculated 2-yr-old Douglas-fir seedlings was monitored on a monthly basis using A/Ci curve analysis. No effect of fungal presence on gas exchange was noted until the emergence of fungal fruiting structures (pseudothecia) from needle stomata. However, once present, maximum stomatal conductance and CO2 assimilation rates were inversely proportional to the presence of pseudothecia. A/Ci curve analysis showed that declines in CO2 assimilation appeared to be due to both stomatal and nonstomatal limitations. Stomatal limitations to CO2 assimilation were the direct result of reduced CO2 diffusion through blocked stomata. Nonstomatal limitations arose, in part, from an indirect effect of pseudothecia development on Rubisco activation. For example, in both Swiss needle cast-infected foliage and foliage with artificially blocked stomata (by external application of petroleum jelly), the amount of Rubisco activation showed a strong, positive relationship with daily maximum stomatal conductance. A mechanism is proposed that outlines the impact of pseudothecia development on stomatal conductance and CO2 assimilation rates.

Stomatal behavior of four woody species in relation to leaf-specific hydraulic conductance and threshold water potential.

Midday stomatal closure is mediated by the availability of water in the soil, leaf and atmosphere, but the response to these environmental and internal variables is highly species specific. We tested the hypothesis that species differences in stomatal response to humidity and soil water availability can be explained by two parameters: leaf-specific hydraulic conductance (K(L)) and a threshold leaf water potential (Psi(threshold)). We used a combination of original and published data to estimate characteristic values of K(L) and Psi(threshold) for four common tree species that have distinctly different stomatal behaviors: black cottonwood (Populus trichocarpa Torr. & Gray.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), red alder (Alnus rubra Bong.) and western hemlock (Tsuga heterophylla (Raf.) Sarg.). We used the values to parameterize a simple, nonelastic model that predicts stomatal conductance by linking hydraulic flux to transpirational flux and maintaining Psi(leaf) above Psi(threshold). The model successfully predicted fundamental features of stomatal behavior that have been reported in the literature for these species. We conclude that much of the variation among the species in stomatal response to soil and atmospheric water deficits can be explained by K(L) and Psi(threshold). The relationship between Psi(threshold) and xylem vulnerability to cavitation differed among these species.