Microbial Drivers of Plant Performance during Drought Depend upon Community Composition and the Greater Soil Environment.

The increasing occurrence of drought is a global challenge that threatens food security through direct impacts to both plants and their interacting soil microorganisms. Plant growth promoting microbes are increasingly being harnessed to improve plant performance under stress. However, the magnitude of microbiome impacts on both structural and physiological plant traits under water limited and water replete conditions are not well-characterized. Using two microbiomes sourced from a ponderosa pine forest and an agricultural field, we performed a greenhouse experiment that used a crossed design to test the individual and combined effects of the water availability and the soil microbiome composition on plant performance. Specifically, we studied the structural and leaf functional traits of maize that are relevant to drought tolerance. We further examined how microbial relationships with plant phenotypes varied under different combinations of microbial composition and water availability. We found that water availability and microbial composition affected plant structural traits. Surprisingly, they did not alter leaf function. Maize grown in the forest-soil microbiome produced larger plants under well-watered and water-limited conditions, compared to an agricultural soil community. Although leaf functional traits were not significantly different between the watering and microbiome treatments, the bacterial composition and abundance explained significant variability in both plant structure and leaf function within individual treatments, especially water-limited plants. Our results suggest that bacteria-plant interactions that promote plant performance under stress depend upon the greater community composition and the abiotic environment. IMPORTANCE Globally, drought is an increasingly common and severe stress that causes significant damage to agricultural and wild plants, thereby threatening food security. Despite growing evidence of the potential benefits of soil microorganisms on plant performance under stress, decoupling the effects of the microbiome composition versus the water availability on plant growth and performance remains a challenge. We used a highly controlled and replicated greenhouse experiment to understand the impacts of microbial community composition and water limitation on corn growth and drought-relevant functions. We found that both factors affected corn growth, and, interestingly, that individual microbial relationships with corn growth and leaf function were unique to specific watering/microbiome treatment combinations. This finding may help explain the inconsistent success of previously identified microbial inocula in improving plant performance in the face of drought, outside controlled environments.

Thicketized oak woodlands reduce groundwater recharge.

Woodlands and pastures across the Post Oak Savannas (POS) in Texas have been undergoing thicketization over the last century via encroachment by understory shrubs such as Yaupon (Ilex decidua, Ilex vomitoria) and expansion of eastern redcedar (Juniperus virginiana). Because a large part of POS overlies the Carrizo-Wilcox (CW) aquifer - the third most important aquifer in Texas, there is a strong incentive to identify opportunities to increase groundwater recharge through land management. The purpose of this research is to evaluate the influence of thicketization of post oak (Quercus stellata) stands on deep drainage (DD) in POS. We achieved this by, a) applying chloride mass balance on soil cores, and b) simultaneously monitoring soil moisture in a woodland pasture setting in POS. Four sites representing different vegetation covers were identified for sampling: 1) a thicketized oak woodland paired with an adjacent open site, 2) a woodland mosaic, 3) a pasture and 4) a pine-oak stand paired with an adjacent open site. A total of 24 soil cores to the depth of 260 cm were collected and (soil) pore water chloride concentrations at multiple depths were measured. Soil moisture was monitored at 21 locations, to the depth of 140-260 cm using a neutron moisture meter. Negligible DD was estimated in the thicketized woodland, whereas most open locations recorded 3-18 cm/year and the woodland mosaic 0-1 cm of DD. Soil moisture data, collected from Jul-2020 to Jun-2021 also suggested higher deep drainage fluxes under open areas - with occurrence of sub-surface saturation only under the open areas and never under the woodlands. These results suggest that the thicketization in oak savannas is substantially reducing groundwater recharge. Given the extent of thicketized oak savannas across United States, this could be impacting water budgets and groundwater recharge rates on regional scales.

Dry Season Transpiration and Soil Water Dynamics in the Central Amazon.

With current observations and future projections of more intense and frequent droughts in the tropics, understanding the impact that extensive dry periods may have on tree and ecosystem-level transpiration and concurrent carbon uptake has become increasingly important. Here, we investigate paired soil and tree water extraction dynamics in an old-growth upland forest in central Amazonia during the 2018 dry season. Tree water use was assessed via radial patterns of sap flow in eight dominant canopy trees, each a different species with a range in diameter, height, and wood density. Paired multi-sensor soil moisture probes used to quantify volumetric water content dynamics and soil water extraction within the upper 100 cm were installed adjacent to six of those trees. To link depth-specific water extraction patterns to root distribution, fine root biomass was assessed through the soil profile to 235 cm. To scale tree water use to the plot level (stand transpiration), basal area was measured for all trees within a 5 m radius around each soil moisture probe. The sensitivity of tree transpiration to reduced precipitation varied by tree, with some increasing and some decreasing in water use during the dry period. Tree-level water use scaled with sapwood area, from 11 to 190 L per day. Stand level water use, based on multiple plots encompassing sap flow and adjacent trees, varied from ∼1.7 to 3.3 mm per day, increasing linearly with plot basal area. Soil water extraction was dependent on root biomass, which was dense at the surface (i.e., 45% in the upper 5 cm) and declined dramatically with depth. As the dry season progressed and the upper soil dried, soil water extraction shifted to deeper levels and model projections suggest that much of the water used during the month-long dry-down could be extracted from the upper 2-3 m. Results indicate variation in rates of soil water extraction across the research area and, temporally, through the soil profile. These results provide key information on whole-tree contributions to transpiration by canopy trees as water availability changes. In addition, information on simultaneous stand level dynamics of soil water extraction that can inform mechanistic models that project tropical forest response to drought.

Hydraulically-vulnerable trees survive on deep-water access during droughts in a tropical forest.

Deep-water access is arguably the most effective, but under-studied, mechanism that plants employ to survive during drought. Vulnerability to embolism and hydraulic safety margins can predict mortality risk at given levels of dehydration, but deep-water access may delay plant dehydration. Here, we tested the role of deep-water access in enabling survival within a diverse tropical forest community in Panama using a novel data-model approach. We inversely estimated the effective rooting depth (ERD, as the average depth of water extraction), for 29 canopy species by linking diameter growth dynamics (1990-2015) to vapor pressure deficit, water potentials in the whole-soil column, and leaf hydraulic vulnerability curves. We validated ERD estimates against existing isotopic data of potential water-access depths. Across species, deeper ERD was associated with higher maximum stem hydraulic conductivity, greater vulnerability to xylem embolism, narrower safety margins, and lower mortality rates during extreme droughts over 35 years (1981-2015) among evergreen species. Species exposure to water stress declined with deeper ERD indicating that trees compensate for water stress-related mortality risk through deep-water access. The role of deep-water access in mitigating mortality of hydraulically-vulnerable trees has important implications for our predictive understanding of forest dynamics under current and future climates.

Species-Specific Shifts in Diurnal Sap Velocity Dynamics and Hysteretic Behavior of Ecophysiological Variables During the 2015-2016 El Niño Event in the Amazon Forest.

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015-2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (Vs), stomatal conductance (gs) and leaf water potential (ΨL) of multiple canopy trees, and their correlations with leaf temperature (Tleaf) and environmental conditions [direct solar radiation, air temperature (Tair) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of gs (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in Tleaf. In the morning, Tleaf and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as Tleaf approached a daily peak, allowing ΨL to begin recovery, while sap velocity declined with an exponential relationship with Tleaf. In Manaus, hysteresis indices of the variables Tleaf-Tair and ΨL-Tleaf were calculated for different species and a significant difference (p < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season ("control scenario"). In some days during the 2015 ENSO event, Tleaf approached 40°C for all studied species and the differences between Tleaf and Tair reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, Tleaf was higher than Tair during the middle morning to early afternoon, and lower than Tair during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in ΨL during the afternoon period giving an observed counterclockwise hysteresis pattern between ΨL and Tleaf.

Warming combined with more extreme precipitation regimes modifies the water sources used by trees.

The persistence of vegetation under climate change will depend on a plant's capacity to exploit water resources. We analyzed water source dynamics in piñon pine and juniper trees subjected to precipitation reduction, atmospheric warming, and to both simultaneously. Piñon and juniper exhibited different and opposite shifts in water uptake depth in response to experimental stress and background climate over 3 yr. During a dry summer, juniper responded to warming with a shift to shallow water sources, whereas piñon pine responded to precipitation reduction with a shift to deeper sources in autumn. In normal and wet summers, both species responded to precipitation reduction, but juniper increased deep water uptake and piñon increased shallow water uptake. Shifts in the utilization of water sources were associated with reduced stomatal conductance and photosynthesis, suggesting that belowground compensation in response to warming and water reduction did not alleviate stress impacts for gas exchange. We have demonstrated that predicted climate change could modify water sources of trees. Warming impairs juniper uptake of deep sources during extended dry periods. Precipitation reduction alters the uptake of shallow sources following extended droughts for piñon. Shifts in water sources may not compensate for climate change impacts on tree physiology.

Dating of 'young' groundwaters using environmental tracers: advantages, applications, and research needs.

Many problems related to groundwater supply and quality, as well as groundwater-dependent ecosystems require some understanding of the timescales of flow and transport. For example, increased concern about the vulnerabilities of 'young' groundwaters (less than ~1000 years) to overexploitation, contamination, and land use/climate change effects are driving the need to understand flow and transport processes that occur over decadal, annual, or shorter timescales. Over the last few decades, a powerful suite of environmental tracers has emerged that can be used to interrogate a wide variety of young groundwater systems and provide information about groundwater ages/residence times appropriate to the timescales over which these systems respond. These tracer methods have distinct advantages over traditional approaches providing information about groundwater systems that would likely not be obtainable otherwise. The objective of this paper is to discuss how environmental tracers are used to characterise young groundwater systems so that more researchers, water managers, and policy-makers are aware of the value of environmental tracer approaches and can apply them in appropriate ways. We also discuss areas where additional research is required to improve ease of use and extend quantitative interpretations of tracer results.

A reservoir of nitrate beneath desert soils.

A large reservoir of bioavailable nitrogen (up to approximately 10(4) kilograms of nitrogen per hectare, as nitrate) has been previously overlooked in studies of global nitrogen distribution. The reservoir has been accumulating in subsoil zones of arid regions throughout the Holocene. Consideration of the subsoil reservoir raises estimates of vadose-zone nitrogen inventories by 14 to 71% for warm deserts and arid shrublands worldwide and by 3 to 16% globally. Subsoil nitrate accumulation indicates long-term leaching from desert soils, impelling further evaluation of nutrient dynamics in xeric ecosystems. Evidence that subsoil accumulations are readily mobilized raises concern about groundwater contamination after land-use or climate change.