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At least 91 records · Page 5

Multiseriate cortical sclerenchyma enhance root penetration in compacted soils

Mechanical impedance limits soil exploration and resource capture by plant roots. We examine the role of root anatomy in regulating plant adaptation to mechanical impedance and identify a root anatomical phene in maize (Zea mays) and wheat (Triticum aestivum) associated with penetration of hard soil: Multiseriate cortical sclerenchyma (MCS). We characterize this trait and evaluate the utility of MCS for root penetration in compacted soils. Roots with MCS had a greater cell wall-to-lumen ratio and a distinct UV emission spectrum in outer cortical cells. Genome-wide association mapping revealed that MCS is heritable and genetically controlled. We identified a candidate gene associated with MCS. Across all root classes and nodal positions, maize genotypes with MCS had 13% greater root lignin concentration compared to genotypes without MCS. Genotypes without MCS formed MCS upon exogenous ethylene exposure. Genotypes with MCS had greater lignin concentration and bending strength at the root tip. In controlled environments, MCS in maize and wheat was associated improved root tensile strength and increased penetration ability in compacted soils. Maize genotypes with MCS had root systems with 22% greater depth and 49% greater shoot biomass in compacted soils in the field compared to lines without MCS. Of the lines we assessed, MCS was present in 30 to 50% of modern maize, wheat, and barley cultivars but was absent in teosinte and wild and landrace accessions of wheat and barley. MCS merits investigation as a trait for improving plant performance in maize, wheat, and other grasses under edaphic stress.

54 ENVIRONMENTAL SCIENCES↗

Numerical Modeling of Air-Blast Suppression as a Function of Explosive-Charge Burial Depth

As a chemical explosion is buried, the mechanism for acoustic wave generation transitions from fully gas-generated at the surface to completely spall-induced at full containment depth. The fully gas-generated and completely spall-induced signals in the acoustic waveform are well described; however, the transition between these two end-members eludes numerical modeling because of the complex phenomena that are involved. The phenomena of crater formation and explosive cloud evolution are simulated using an Eulerian hydrocode that incorporates geomaterials with strength and porosity. Having accurately modeled these phenomena, we can confidently predict the propagation and relative strength of the gas-generated and spall-induced pulses in the recorded acoustic waveform. The numerical predictions agree with observations from the historical Stagecoach experiment as well as modern recordings from the Source Physics Experiment. In particular, the peak pressure p generated by an explosion is initially due to the gas-generated mechanism and decays with scaled depth of burial d s (depth d scaled by the cube-root of explosive yield w 1/3 ) as exp(-d s ) but then transitions near a scaled depth of 6 m/ton 1/3 to the spall-generated mechanism in which the decay is d$_{s}^{-7/4}$. This decay form is related to the strong ground-motion attenuation relationship that affects spall strength. So these results can improve seismoacoustic inverse models for the explosive source that need to account for the gas-generated and spall-induced signals and their effect on peak pressures and other acoustic signal features.

58 GEOSCIENCES↗

Alteration of the Cesium-137 soil profile by wild boar rooting after the Fukushima Daiichi Nuclear Power Plant accident

Large amounts of Cesium-137 ( 137 Cs, physical half-life ca. 30 years) were released into the environment from the 2011 Fukushima Daiichi Nuclear Power Plant (FDNPP) accident in Japan. Initially contaminants from the accident were deposited on the surface soil layers in terrestrial ecosystems, but over time vertical migration of 137 Cs has occurred, with most 137 Cs now distributed a few centimeters below the soil surface. Wild boar (Sus scrofa) rooting likely alters the distribution of contaminants in soil profiles. Such impacts may be particularly pronounced in areas contaminated from the FDNPP accident, given the increase in abundance of wild boar in these areas. Our objective was to determine whether rooting by wild boar alters the distribution of 137 Cs contamination within soil profiles through collection of soil cores from control sites and two different types of rooting samples, “new depressions” and “old depressions” within the difficult-to-return zone surrounding the FDNPP. The 137 Cs ratio [activity concentration of 137 Cs at each depth / the maximum activity concentration of 137 Cs in each core sample] rapidly decreased after reaching a peak at the control sites. However, the 137 Cs ratio at all rooted sites decreased more slowly, and 137 Cs was distributed deeper into the soil layers than control sites, across both new and old rooting sites. Moreover, the 137 Cs soil profiles varied greatly among core samples and sampling sites. Furthermore, these results demonstrate that soil disturbance from wild boar rooting can directly influence the distribution of contaminants within soil profiles. However, the extent to which deeper layers are affected by contamination depends on the unique characteristics (i.e., depth of rooting) of each wild boar rooting disturbance. Further studies are needed to examine the effects of long-term rooting on the alteration of 137 Cs in soil profiles.

63 RADIATION, THERMAL, AND OTHER ENVIRON. POLLUTAN↗

Wildfire exacerbates high-latitude soil carbon losses from climate warming

Abstract Arctic and boreal permafrost soil organic carbon (SOC) decomposition has been slower than carbon inputs from plant growth since the last glaciation. Anthropogenic climate warming has threatened this historical trend by accelerating SOC decomposition and altering wildfire regimes. We accurately modeled observed plant biomass and carbon emissions from wildfires in Alaskan ecosystems under current climate conditions. In projections to 2300 under the RCP8.5 climate scenario, we found that warming and increased atmospheric CO 2 will result in plant biomass gains and higher litterfall. However, increased carbon losses from (a) wildfire combustion and (b) rapid SOC decomposition driven by increased deciduous litter production, root exudation, and active layer depth will lead to about 4.4 PgC of soil carbon losses from Alaska by 2300 and most (88%) of these loses will be from the top 1 m of soil. These SOC losses offset plant carbon gains, causing the ecosystem to transition to a net carbon source after 2200. Simulations excluding wildfire increases yielded about a factor of four lower SOC losses by 2300. Our results show that projected wildfire and its direct and indirect effects on plant and soil carbon may accelerate high-latitude soil carbon losses, resulting in a positive feedback to climate change.

54 ENVIRONMENTAL SCIENCES↗

Solution for Predictive Physical Modeling in CdTe and Other Thin-Film PV Technologies

The main goal of this work was to eliminate the ambiguity between the observed performance changes under stress and their physical root cause by enabling a depth of modeling that takes account of diffusion and drift at the atomistic level coupled to the electronic subsystem responsible for a PV device’s function. Development of advanced physical models describing capture/emission/ recombination phenomena relevant to multivalent dopants, amphoteric centers, and donor-acceptor pairs was also accomplished. These models, incorporating grain bulk, grain boundaries, and hetero-interface properties, were implemented in a self-contained simulation tool that drastically reduces interpretation ambiguity and, for the first time, allows for predictive design of thin-film PV devices.

14 SOLAR ENERGY↗

SPRUCE Measurements of Fine Root Production and Chemistry from Root Ingrowth Cores, Marcell Experimental Forest, Minnesota, 2022-2023

This dataset contains fine root production and tissue chemistry measurements from root ingrowth cores. Ingrowth cores were deployed in peat from June 28, 2022 to June 24, 2023 (2022-06-28 to 2023-06-24) inside SPRUCE Experiment plots at the Marcell Experimental Forest in northern Minnesota. The warming and elevated carbon dioxide (CO2) treatments in this dataset include +0 degrees Celsius (C) (+0 and +500 parts per million (ppm) elevated CO2), +4.5 degrees C (+0 and +500 ppm elevated CO2) and +9 degrees C (+0 and +500 ppm elevated CO2) for both hummocks and hollows, as well as +2.25 degrees C (+0 and +500 ppm) and +6.75 degrees C (+0 and +500 ppm) for hollows from minimum 10 cm depth from the peat surface. Measurements include root average diameter, root length, root biomass, and root tissue nitrogen (%N and δ15N) and carbon (%C and δ13C) concentration per plant functional type and microtopographical feature. Root length and biomass are standardized to 10 cm depth. These data were used to assess the warming and elevated CO2 response of fine roots across different peatland microtopographical features (hummocks and hollows) and plant functional types (shrub, spruce and larch). This dataset contains one data file in comma separate (.csv) format. Additional metadata are provided: one data dictionary and a file-level metadata file in comma separate (.csv) format and a user guide in PDF (*.pdf) format.

ESS-DIVE CSV File Formatting Guidelines Reporting ↗

Experimental Soil Warming Impacts Soil Moisture and Plant Water Stress and Thereby Ecosystem Carbon Dynamics (Blodgett, CA)

This dataset contains data on daily soil temperature, moisture and flux, and soil carbon stock and root biomass across a soil profile down to 100 cm depth at Blodgett Forest Research Station, CA, USA. These data were generated to determine if modeling of an experimental soil warming of 4C showed increased soil CO2 emissions and changes in bulk soil carbon stocks with depth consistent with field observations, as part of the study: Riley et al. (2025) Experimental Soil Warming Impacts Soil Moisture and Plant Water Stress and Thereby Ecosystem Carbon Dynamics in Journal of Advances in Modeling Earth Systems. This research was performed within the framework of the TES Belowground Biogeochemistry SFA project, in particular association with a 1 m-deep experimental soil heating experiment at the University of California Blodgett Forest Research Station, California (120 ° 39′40′′W; 38 ° 54′43′′N). Continuous data were collected at the plot level, and bulk soil carbon and root biomass were sampled once a year from each plot from 0-100 cm, in 10 cm intervals. Measurements relevant to the current study include soil temperature and soil volumetric water content measured continuously at multiple depths in the top meter; fine root biomass and SOC stocks measured from annual soil cores. Soil flux was continuously monitored using a LI-8100 Automated CO2 Flux System in conjunction with the LI-8150 Multiplexer (Licor, Nebraska, USA). Soil flux was determined using SoilFluxPro software, with flux values showing an R² fit of less than 0.9 being excluded from the analysis. Data were collected from each paired plot (1-3): one control (C) and one heated (H).

54 ENVIRONMENTAL SCIENCES↗

Bioenergy sorghum’s deep roots: A key to sustainable biomass production on annual cropland

Abstract Bioenergy sorghum has high biomass yield potential, drought resilience, good nitrogen use efficiency, and a root system that contributes to the accumulation of soil organic carbon. In this study, field grown bioenergy sorghum root systems were analyzed during the growing season to characterize their depth, biomass, morphology, anatomy, and gene expression profiles. Bioenergy sorghum roots grew continuously during a 155‐day growing season producing ~175 nodal roots, accumulating ~7 Mg of dry biomass per hectare, and reaching >2 m deep in the soil profile. Nodal roots within 20 cm of the stem were 1–5 mm in diameter, whereas roots deeper in soil profiles were enriched in lateral roots with small diameters (~30–500 µm) enabling growth through soil macropores. In older field‐grown plants, roots with intact endodermal, vascular and inner root tissues were surrounded by degraded or aerenchyma‐filled epidermal and cortical cell layers. Transcriptome analysis of nodal, surface, and deep roots identified >2,500 differentially expressed genes involved in root growth, transport, adaptation, defense, and AMF–root interaction. Deep roots (180–240 cm) differentially expressed genes that regulate lateral root growth. Surface roots (0–20 cm) located mid‐row differentially expressed genes involved in nitrate transport, whereas ammonium transport genes were expressed in surface and deep roots and genes involved in phosphate transport were expressed in nodal, surface, and deep roots. Overall, bioenergy sorghum's long growing season enables root systems to grow deeper and accumulate more biomass than annual grain crops such as maize, attributes that could help restore annual cropland soil organic carbon levels and improve soil productivity. Deep roots active in nutrient transport are positioned to take‐up fertilizer leached deep into soil profiles mitigating potential nutrient run‐off. Bioenergy sorghum's large and deep root system is a key to sustainable production of biofuels, biopower, and bioproducts on annual cropland.

59 BASIC BIOLOGICAL SCIENCES↗

Daily water stable isotopes, transpiration, and matrix potential data for an aspen and engelmann stand in the East River Watershed (version 2)

We provide daily stable isotope (2H & 18O) ratios in soil water and xylem (plant stem) water, as well as the sap flow (transpiration) and the soil's matric potential at a forested site near Gothic, Colorado, in the East River catchment. We measured the stable isotopic composition of the transpiration and the daily transpiration flux sum of three aspen and three engelmann spruce. In both forest stands, we installed a soil profile and measured the soil matric potential at 15, 30, and 60 cm depth as well as the stable isotopes of soil pore water at 5, 10, 30, 60, and 90 cm depths. All isotope measurements were done in situ via vapor probes connected to a cavity ring down spectrometer (Picarro L1240i).We further report the daily meteorological data observed at billy barr near our study site. We also provide for each tree the relative share of root water uptake derived from the isotope measurements via a Bayesian mixing model (MixSIAR).The daily data is provided as a time series in "Iso_MP_Sap_DataDaily_ESSDiveUpload.csv" and the units are provided in "dd.csv"; the location of the instrumented trees and soil profiles are given as latitude and longitude coordinates saved as CSV and KMZ files; and a file-level metadata (flmd.csv) file that lists each file contained in the dataset with associated metadata.The data was gathered to investigate the short-term changes of the water sources (i.e., variation of root water uptake from different soil depths) of the studied subalpine trees.Update 07/16/2025: The relative and absolute plant water uptake depths were grouped to ensure that the MixSIAR model was applied with endmembers that differed in their d2H value by at least 3 permill and at least 1 permill for d18O. Whenever the difference between observed d2H values for two or more probes at neighboring depths was less than 3 permill, we used the average value for the source water endmember. For days at which probes that were not next to each other measurements did not differ at least 3 permill, the average of all probes between these two depths was used as the water source endmember representing the depth range between these two probes.

54 ENVIRONMENTAL SCIENCES↗

Low-Cost X-Ray CT System for Imaging of Roots

The goal of this project was to develop and demonstrate an innovative, low cost, field deployable, stationary 3D x-ray computed tomography (CT) system that will image total root phenotypes with a micron size resolution at a throughput of hundreds of plants per cycle. This system is based on UHV’s unique low cost linear x-ray tube technology and sophisticated reconstruction & image segmentation algorithms developed at University of Massachusetts, Lowel and University of Nottingham; and was tested for several types of soils at University of Wisconsin and Texas A&M University. Currently, no technologies exist that have been designed to image roots in complex media such as agricultural field conditions. Due to its small size, high resolution & fast imaging of fine roots, low power consumption, large penetration depth (i.e. ability to see through several feet of soil) and ease of field deployability, this CT system will increase the speed and efficacy of discovery, field translation, and deployment of improved crops and systems that improve soil carbon accumulation and storage, decrease N2O emissions, and improve water efficiency leading towards advancements that could mitigate 10% of the total US Greenhouse gases. This degree of imaging in the field has never been available and would be invaluable to scientists in understanding how environmental conditions and phenotypic variations contribute to carbon deposition through root development.

54 ENVIRONMENTAL SCIENCES↗

Persistence and Plasticity in Conifer Water–Use Strategies

The selective use of seasonal precipitation by vegetation is critical to understanding the residence time and flow path of water in watersheds, yet there are limited datasets to test how climate alters these dynamics. Here, we use measurements of the seasonal cycle of tree ring δ 18 O for two widespread conifer species in the Rocky Mountains of North America to provide a multi-decadal depiction of the seasonal origins of forest water use. The results show that while the conifer tree stands had a dominant preference for use of snowmelt, there were multi-annual periods over the last four decades when use of summer precipitation was preferential. Utilization of summer rain emerged during years with increased snowfall and tree growth, suggesting that summer rain enhanced the transpiration stream only during the periods of highest water use. We hypothesize this could be explained through shallowing of the root profile during wetter periods and/or through the influence of changing water table depths on the residence time of summer precipitation in the root zone. We suggest the tree ring proxy approach used here could be applied in other watersheds to provide critical insight into the temporal dynamics of plant water use that could not be inferred from short measurement campaigns. Furthermore, these data on the seasonal origins of forest water are critical for understanding forest vulnerability to drought, the processes that affect precipitation pathways and residence time in watersheds and the interpretation of tree ring proxy data.

59 BASIC BIOLOGICAL SCIENCES↗

SPRUCE S1 Bog Production of Newly-Grown Fine Roots Assessed Using Root Ingrowth Cores in 2013

This dataset contains data from root ingrowth cores from the Spruce and Peatland Responses Under Changing Environments (SPRUCE) experimental site within the Marcell Experimental Forest in northern Minnesota, USA. Root ingrowth cores were used to capture newly-grown fine roots during the 2013 growing season (June - October) at 20 locations at the southern end of the S1-Bog. Rates of fine root production were measured with root ingrowth cores and fine root traits were measured on roots that grew into ingrowth cores. Traits are reported by plant species and root orders and include nitrogen (N) content, root tissue density, specific root length, average diameter. Ingrowth cores were used to compare species-specific depth distribution patterns with minirhizotron analyses and to obtain newly produced roots for chemical analysis and measurement of morphological traits prior to the construction of the SPRUCE experimental plots. This dataset contains two data files in comma-separate values (*.csv) format. Additional metadata are provided: two data dictionaries and a file-level metadata file in comma-separate values (.csv) format and a user guide in PDF (*.pdf) format.

54 ENVIRONMENTAL SCIENCES↗

Fine-root functional trait responses to experimental warming: a global meta-analysis

Whether and how warming alters functional traits of absorptive plant roots remains to be answered across the globe. Tackling this question is crucial to better understanding terrestrial responses to climate change as fine-root traits drive many ecosystem processes. In this work, we carried out a detailed synthesis of fine-root trait responses to experimental warming by performing a meta-analysis of 964 paired observations from 177 publications. Warming increased fine-root biomass, production, respiration and nitrogen concentration as well as decreased root carbon : nitrogen ratio and nonstructural carbohydrates. Warming effects on fine-root biomass decreased with greater warming magnitude, especially in short-term experiments. Furthermore, the positive effect of warming on fine-root biomass was strongest in deeper soil horizons and in colder and drier regions. Total fine-root length, morphology, mortality, life span and turnover were unresponsive to warming. Our results highlight the significant changes in fine-root traits in response to warming as well as the importance of warming magnitude and duration in understanding fine-root responses. These changes have strong implications for global soil carbon stocks in a warmer world associated with increased root-derived carbon inputs into deeper soil horizons and increases in fine-root respiration.

59 BASIC BIOLOGICAL SCIENCES↗

Root system architecture and environmental flux analysis in mature crops using 3D root mesocosms

Current methods of root sampling typically only obtain small or incomplete sections of root systems and do not capture their true complexity. To facilitate the visualization and analysis of full-sized plant root systems in 3-dimensions, we developed customized mesocosm growth containers. While highly scalable, the design presented here uses an internal volume of 45 ft 3 (1.27 m 3 ), suitable for large crop and bioenergy grass root systems to grow largely unconstrained. Furthermore, they allow for the excavation and preservation of 3-dimensional root system architecture (RSA), and facilitate the collection of time-resolved subterranean environmental data. Sensor arrays monitoring matric potential, temperature and CO 2 levels are buried in a grid formation at various depths to assess environmental fluxes at regular intervals. Methods of 3D data visualization of fluxes were developed to allow for comparison with root system architectural traits. Following harvest, the recovered root system can be digitally reconstructed in 3D through photogrammetry, which is an inexpensive method requiring only an appropriate studio space and a digital camera. We developed a pipeline to extract features from the 3D point clouds, or from derived skeletons that include point cloud voxel number as a proxy for biomass, total root system length, volume, depth, convex hull volume and solidity as a function of depth. Ground-truthing these features with biomass measurements from manually dissected root systems showed a high correlation. We evaluated switchgrass, maize, and sorghum root systems to highlight the capability for species wide comparisons. We focused on two switchgrass ecotypes, upland (VS16) and lowland (WBC3), in identical environments to demonstrate widely different root system architectures that may be indicative of core differences in their rhizoeconomic foraging strategies. Finally, we imposed a strong physiological water stress and manipulated the growth medium to demonstrate whole root system plasticity in response to environmental stimuli. Hence, these new “3D Root Mesocosms” and accompanying computational analysis provides a new paradigm for study of mature crop systems and the environmental fluxes that shape them.

59 BASIC BIOLOGICAL SCIENCES↗

Drivers for the cracking of multilayer polyamide‐based backsheets in field photovoltaic modules: In‐depth degradation mapping analysis

Abstract There is a lack of understanding on the root cause of cracking of photovoltaic (PV) backsheets due to the challenge of multilayer characterization and the complicated failure modes at the submodule level. In this work, the in‐depth degradation mapping of field‐exposed polyamide‐based (PA‐based) PV module backsheets was studied, with the major focus on the identification of underlying drivers for the through cracking (in between the solar cells). PV modules were retrieved from five different locations, comprising a variety of climates, including humid subtropical, hot‐summer Mediterranean, tropical savanna climate, and hot arid. A suite of microscale cross‐sectional characterizations, including chemical changes, fluorescence intensity, and modulus as a function of distance from the air surface of the backsheet, was performed. Results showed more advanced signs of degradation of the inner layer than the outer layer in the cracking region. Increases in the modulus were identified as the major indicator for the cracking. Moreover, a rudimentary test by immersion in acetic acid, which forms during photodegradation of the ethylene‐vinyl acetate copolymer (EVA) encapsulant, showed the first‐time direct evidence that acetic acid can largely accelerate the chemical degradation and facilitate the cracking of PA inner layer. This study suggests that the field cracking of PA‐based backsheet can be attributed to the combined effects of chemical degradation and physical reorganization (chemi‐crystallization) under cyclic thermomechanical stresses. Therefore, it is important to consider effects of local microclimates and interlayer infection to understand the heterogeneous nature of backsheet failures.

Lyu, Yadong↗

Castanha et al. 2018: Hopland Lysimeter 13C-Labeled Root Litter Decomposition Study.

The breakdown and decomposition of plant inputs are critical for nutrient cycling, soil development, and climate-ecosystem feedbacks, but uncertainties persist in how the rates and products of litter decomposition are affected by soil temperature, rhizosphere, and depth of input. To elucidate these questions we measured the effects of soil warming (+ 4 °C), rhizosphere, and depth of litter placement on the decomposition of Avena fatua (wild oat grass) root litter in a Mediterranean grassland ecosystem. Field lysimeters were subjected to three environmental treatments (heating, control, and plant removal) and three 13C-labeled root litter addition treatments (to A horizon, to B horizon, and no-addition disturbance control) for each of two harvest time points. We buried root litter in February 2014 and measured loss of 13C in CO2 from the soil surface and in leachate as dissolved organic carbon (DOC) over two growing seasons. At the end of each of the 2014 and 2015 growing seasons we recovered the 13C remaining in the soil. Loss of root litter C occurred almost entirely via heterotrophic respiration, with an estimated < 2% lost as DOC during the initial decay period. The added roots were broken down and incorporated into bulk soil material very quickly; only ~ 30% of added root was visible after 6 months. In the first growing season, decomposition occurred faster in the B than in the A horizon, the latter having greater moisture limitation. Subsequently, there was almost no further decomposition in the B horizon. After two growing seasons, less than 20% of the added root litter C remained in the A or B horizons of all environmental treatments. Heating did not stimulate decomposition, likely because it exacerbated the moisture limitation. However, while plots without plants dried down more slowly than plots with plants, their decomposition rate was not significantly greater, possibly due to the lack of rhizosphere processes such as priming. We conclude that in this Mediterranean grassland ecosystem, soil moisture, which is affected by season, depth, heating, and rhizosphere, plays a dominant role in mediating the effect of those factors on root litter decomposition, which after two seasons did not differ by depth or by treatment.The files in this dataset comprise (1) 13C-labeled litter additions made to each plot, (2) litter recovery after each of 2 growing seasons, (3) periodical soil flux measurements, (4) leachate measurements, and continuously monitored soil (5) temperature and (6) moisture.

54 ENVIRONMENTAL SCIENCES↗

Hydraulic redistribution by deeply rooted grasses and its ecohydrologic implications in the southern Great Plains of North America

Perennial bioenergy crops with deep (>1 m) rooting systems, such as switchgrass (Panicum virgatum L.), are hypothesized to increase carbon storage in deep soil. Deeply rooted plants may also affect soil hydrology by accessing deep soil water for transpiration, which can affect soil water content and infiltration in deep soil layers, thereby affecting groundwater recharge. Using stable H and O isotope (δ 2 H and δ 18 O) and 3 H values, we studied the soil water conditions at 20–30 cm intervals to depths of 2.4–3.6 m in paired fields of switchgrass and shallow rooted crops at three sites in the southern Great Plains of North America. We found that soil under switchgrass had consistently higher soil water content than nearby soil under shallow-rooted annual crops by a margin of 15%–100%. Soil water content and isotopic depth profiles indicated that hydraulic redistribution of deep soil water by switchgrass roots explained these observed soil water differences. To our knowledge, these are the first observations of hydraulic redistribution in deeply rooted grasses, and complement earlier observations of dynamic soil water fluxes under shallow-rooted grasses. Hydraulic redistribution by switchgrass may be a strategy for drought avoidance, wherein the plant may actively prevent water limitation. Furthermore, this raises the possibility that deeply rooted grasses may be used to passively subsidize soil water to more shallow-rooted species in inter-cropping arrangements.

3H↗