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At least 379 records · Page 21

Expression of characteristics of ammonium nutrition as affected by pH of the root medium

To study the effect of root-zone pH on characteristic responses of NH4+ -fed plants, soybeans (Glycine max inverted question markL. inverted question mark Merr. cv. Ransom) were grown in flowing solution culture for 21 d on four sources of N (1.0 mol m-3 NO3-, 0.67 mol m-3 NO3- plus 0.33 mol m-3 NH4+, 0.33 mol m-3 NO3- plus 0.67 mol m-3 NH4+, and 1.0 mol m-3 NH4+) with nutrient solutions maintained at pH 6.0, 5.5, 5.0, and 4.5. Amino acid concentration increased in plants grown with NH4+ as the sole source of N at all pH levels. Total amino acid concentration in the roots of NH4+ -fed plants was 8 to 10 times higher than in NO3(-)-fed plants, with asparagine accounting for more than 70% of the total in the roots of these plants. The concentration of soluble carbohydrates in the leaves of NH4+ -fed plants was greater than that of NO3(-)-fed plants, but was lower in roots of NH4+ -fed plants, regardless of pH. Starch concentration was only slightly affected by N source or root-zone pH. At all levels of pH tested, organic acid concentration in leaves was much lower when NH4+ was the sole N source than when all or part of the N was supplied as NO3-. Plants grown with mixed NO3- plus NH4+ N sources were generally intermediate between NO3(-)- and NH4+ -fed plants. Thus, changes in tissue composition characteristic of NH4+ nutrition when root-zone pH was maintained at 4.5 and growth was reduced, still occurred when pH was maintained at 5.0 or above, where growth was not affected. The changes were slightly greater at pH 4.5 than at higher pH levels.

NASA Discipline Life Support Systems↗

Soluble carbohydrate allocation to roots, photosynthetic rate of leaves, and nitrate assimilation as affected by nitrogen stress and irradiance

Upon resupply of exogenous nitrogen to nitrogen-stressed plants, uptake rate of nitrogen is enhanced relative to nonstressed plants. Absorption of nitrogen presumably is dependent on availability of carbohydrates in the roots. A buildup in soluble carbohydrates thus should occur in roots of nitrogen-stressed plants, and upon resupply of exogenous nitrogen the increased uptake rate should be accompanied by a rapid decline in carbohydrates to prestress levels. To evaluate this relationship, three sets of tobacco plants growing in a complete hydroponic solution containing 1.0 mM NO3- were either continued in the complete solution for 21 d, transferred to a minus-nitrogen solution for 21 d, or transferred to a minus-nitrogen solution for 8-9 d and then returned to the 1.0 mM NO3- solution. These nitrogen treatments were imposed upon plants growing at photosynthetic photon flux densities of 700 and 350 micromoles m-2 s-1. Soluble carbohydrate levels in roots increased during onset of nitrogen stress to levels that were fourfold greater than in roots of non-stressed plants. Following resupply of external nitrogen, a rapid resumption of nitrogen uptake was accompanied by a decline in soluble carbohydrates in roots to levels characteristic of nonstressed plants. This pattern of soluble carbohydrate levels in roots during onset of and recovery from nitrogen stress occurred at both irradiance levels. The response of net photosynthetic rate to nitrogen stress could be expressed as a nonlinear function of concentration of reduced nitrogen in leaves. The net photosynthetic rate at a given concentration of reduced nitrogen, however, averaged 10% less at the lower than at the higher irradiance. The decline in net photosynthetic rate per unit of reduced nitrogen in leaves at the lower irradiance was accompanied by an increase in the nitrate fraction of total nitrogen in leaves from 20% at the higher irradiance to 38% at the lower irradiance.

NASA Discipline Number 61-10↗

Interaction of root gravitropism and phototropism in Arabidopsis wild-type and starchless mutants

Root gravitropism in wild-type Arabidopsis and in two starchless mutants, pgm1-1 and adg1-1, was evaluated as a function of light position to determine the relative strengths of negative phototropism and of gravitropism and how much phototropism affects gravitropic measurements. Gravitropism was stronger than phototropism in some but not all light positions in wild-type roots grown for an extended period, indicating that the relationship between the two tropisms is more complex than previously reported. Root phototropism significantly influenced the time course of gravitropic curvature and the two measures of sensitivity. Light from above during horizontal exposure overestimated all three parameters for all three genotypes except the wild-type perception time. At the irradiance used (80 micromol m(-2) s(-1)), the shortest periods of illumination found to exaggerate gravitropism were 45 min of continuous illumination and 2-min doses of intermittent illumination. By growing roots in circumlateral light or by gravistimulating in the dark, corrected values were obtained for each gravitropic parameter. Roots of both starchless mutants were determined to be about three times less sensitive than prior estimates. This study demonstrates the importance of accounting for phototropism in the design of root gravitropism experiments in Arabidopsis.

Non-NASA Center↗

A Dominant Arabidopsis Thaliana ACTIN7 Mutant for Studies of Cytoskeletal-Mediated Root Gravity and Spaceflight Stress Responses

A forward genetic screen for Arabidopsis thaliana mutants that exhibited differential sensitivity to the actin-disrupting compound, latrunculin B (LatB), was conducted to uncover new players involved in actin-mediated root gravity responses. This to the isolation of a mutant that exhibited robust primary root growth at 100 nM LatB, which is a concentration that severely inhibits wild-type root elongation. Phenotypic analysis revealed that hypocotyl elongation in the dark and root hair tip growth in the mutant could tolerate LatB concentrations that impaired these processes in wild type. A cross between the mutant and wild type resulted in progeny resistance to LatB, which indicated that the mutant is dominant (hereafter referred to as LBR1 for LatB Resistant1). Filamentous-actin (F-actin) organization in LBR1 primary roots remained intact at 100 nM LatB, while that of wild type exhibited fragmented F-actin. Next generation sequencing revealed that LBR1 had a single nucleotide polymorphism (SNP) in the AT5G09810 gene that changed cytosine to a thiamine at the first exon. AT5G09810 encodes ACTIN7 (ACT7), which is one of three vegetative actin isoforms in A. thaliana. The SNP in the ACT7 gene led to a change in a single amino acid from proline at position 32 to a serine. Transgenic complementation of LBR1 plants with wild-type ACT7 under the control of the ACT7 promoter (pACT7:ACT7) and wild-type plants with LBR1, which contained the proline to serine mutation, also under the ACT7 promoter (pACT7:LBR1) confirmed that LBR1 is ACT7. The pACT7:LBR1 construct was also able to confer LatB resistance to the act7-5 and act2-3 vegetative ACT mutants. Moreover, LBR1 exhibited partial tolerance to salt and low phosphate, and enhanced root skewing on a clinostat, suggesting that site-directed engineering of vegetative ACT presents a strategy for generating stress-tolerant plants for spaceflight applications and studies of actin-mediated gravity responses.

Plant Space Biology↗

A Dominant Arabidopsis Thaliana ACTIN7 Mutant for Studies of Cytoskeletal-Mediated Root Gravity and Spaceflight Stress Responses

A forward genetic screen for Arabidopsis thaliana mutants that exhibited differential sensitivity to the actin-disrupting compound, latrunculin B (LatB), was conducted to uncover new players involved in actin-mediated root gravity responses. This led to the isolation of a mutant that exhibited robust primary root growth at 100 nM LatB, which is a concentration that severely inhibits wild-type root elongation. Phenotypic analysis revealed that hypocotyl elongation in the dark and root hair tip growth in the mutant could tolerate LatB concentrations that impaired these processes in wild type. A cross between the mutant and wild type resulted in progeny resistance to LatB, which indicated that the mutant is dominant (hereafter referred to as LBR1 for LatB Resistant1). Filamentous-actin (F-actin) organization in LBR1 primary roots remained intact at 100 nM LatB, while that of wild type exhibited fragmented F-actin. Next generation sequencing revealed that LBR1 had a single nucleotide polymorphism (SNP) in the AT5G09810 gene that changed cytosine to a thiamine at the first exon. AT5G09810 encodes ACTIN7 (ACT7), which is one of three vegetative actin isoforms in A. thaliana. The SNP in the ACT7 gene led to a change in a single amino acid from proline at position 32 to a serine. Transgenic complementation of LBR1 plants with wild-type ACT7 under the control of the ACT7 promoter (pACT7:ACT7) and wild-type plants with LBR1, which contained the proline to serine mutation, also under the ACT7 promoter (pACT7:LBR1) confirmed that LBR1 is ACT7. The pACT7:LBR1 construct was also able to confer LatB resistance to the act7-5 and act2-3 vegetative ACT mutants. Moreover, LBR1 exhibited partial tolerance to salt and low phosphate, and enhanced root skewing on a clinostat, suggesting that site-directed engineering of vegetative ACT presents a strategy for generating stress-tolerant plants for spaceflight applications and studies of actin-mediated gravity responses.

Plant Space Biology↗

Root phenotypes for improved nitrogen capture

Abstract Background Suboptimal nitrogen availability is a primary constraint for crop production in low-input agroecosystems, while nitrogen fertilization is a primary contributor to the energy, economic, and environmental costs of crop production in high-input agroecosystems. In this article we consider avenues to develop crops with improved nitrogen capture and reduced requirement for nitrogen fertilizer. Scope Intraspecific variation for an array of root phenotypes has been associated with improved nitrogen capture in cereal crops, including architectural phenotypes that colocalize root foraging with nitrogen availability in the soil; anatomical phenotypes that reduce the metabolic costs of soil exploration, improve penetration of hard soil, and exploit the rhizosphere; subcellular phenotypes that reduce the nitrogen requirement of plant tissue; molecular phenotypes exhibiting optimized nitrate uptake kinetics; and rhizosphere phenotypes that optimize associations with the rhizosphere microbiome. For each of these topics we provide examples of root phenotypes which merit attention as potential selection targets for crop improvement. Several cross-cutting issues are addressed including the importance of soil hydrology and impedance, phenotypic plasticity, integrated phenotypes, in silico modeling, and breeding strategies using high throughput phenotyping for co-optimization of multiple phenes. Conclusions Substantial phenotypic variation exists in crop germplasm for an array of root phenotypes that improve nitrogen capture. Although this topic merits greater research attention than it currently receives, we have adequate understanding and tools to develop crops with improved nitrogen capture. Root phenotypes are underutilized yet attractive breeding targets for the development of the nitrogen efficient crops urgently needed in global agriculture.

60 APPLIED LIFE SCIENCES↗

SPRUCE High-Resolution Minirhizotrons in an Experimentally-Warmed Peatland Provide an Unprecedented Glimpse at Fine Roots and their Fungal Partners: Supporting Data

Images were collected using first of their kind, non-destructive, high-resolution automated minirhizotrons (RhizoSystems, LLC) to assess the response of plant fine-root and fungal mycelium dynamics to elevated temperatures after 4-6 years of whole-ecosystem warming and exposure to elevated carbon dioxide concentrations (e[CO2]) in a peat bog where the SPRUCE (Spruce and Peatland Responses Under Changing Environments) experiment is located. We focused on two SPRUCE experimental plots: Plot 10 has elevated temperature (+9°C) and plot 19 is a control (+0°C). Both have elevated carbon dioxide (e[CO2]). Changes in root and fungal abundance with warming were estimated from a timeseries of landscape-level mosaiced images for each plot by measuring the proportional abundance of five belowground classes: fine roots of vascular plants, ectomycorrhizas, fungal hyphae, fungal rhizomorphs, and fungal sporocarps. To examine root and fungal phenology responses to warming, the length per individual root or fungal structure (except fungal hyphae that did not grow linearly but rather increased in areal coverage) were measured per image area of a set of timeseries patch-level mosaiced images for each plot. The experimental work was conducted in a Picea mariana [black spruce] – Sphagnum spp. bog forest in northern Minnesota, 40 km north of Grand Rapids, in the USDA Forest Service Marcell Experimental Forest (MEF). This ecosystem, which is located at the southern margin of the boreal forest, is considered especially vulnerable to climate change and anticipated to be near its tipping point. These data were used in analyses published in Defrenne et al (2021). This dataset contains 5 data files in comma-separate values (*.csv) format and a compressed folder (*.zip) containing 383 JPEG (*.jpg) images. Data files contain landscape-level assessment of belowground class abundance, patch-level growth phenology, and environmental variables (originally published in Hanson et al., 2016 and Hanson et al 2020). Images were collected with automated minirhizotrons and analyzed for phenology. Additional metadata are provided: 5 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↗

Influence of PRE-emergence herbicides on soybean development, root nodulation and symbiotic nitrogen fixation

We report widespread occurrence of herbicide-resistant weeds has further limited effective POST-emergence herbicide options in soybean (Glycine max (L.) Merr.) leading to an increased adoption of PRE-emergence herbicides. The objective of this study was to investigate the influence of 11 commonly used PRE herbicides on soybean development, root nodulation, and symbiotic N fixation. Soybean plants were grown under greenhouse conditions in pots (10 L; 4 plants per pot) filled with silt loam soil and treated one day after planting with a labeled field rate of imazethapyr, chlorimuron-ethyl, cloransulam-methyl, metribuzin, sulfentrazone, flumioxazin, saflufenacil, acetochlor, S-metolachlor, dimethenamid-P, pyroxasulfone and no herbicide (nontreated control). Sulfentrazone reduced soybean canopy at the VC growth stage but no canopy reduction was observed at the V2 growth stage from any of the herbicide treatments. At the R2 growth stage, herbicides had no effect on soybean development (root and shoot biomass), root nodulation (# nodule per plant, nodule diameter, and nodule biomass) and symbiotic N fixation (acetylene reduction assay and 15 N natural abundance). According to our findings, although PRE herbicides may slightly affect early-season soybean development, the impacts on plant growth, root nodulation, and symbiotic N fixation were negligible. Thus, when sprayed according to the label, the benefits of PRE herbicides for weed control likely outweigh any potential concern regarding soybean development, root nodulation, and N fixation.

54 ENVIRONMENTAL SCIENCES↗

Root exudate composition from different plant species influences the growth of rhizosphere bacteria

Plant roots release exudates that fuel microbial activities and can structure rhizosphere microbial communities, but how different plant species use their root exudate to potentially select for different soil microbes in the rhizosphere is not well understood. Here, we investigated how root exudate from plants of three diverging lineages, Lactuca sativa (lettuce), Brassica juncea (mustard cabbage), and Zea mays (maize) influence the growth of their own rhizosphere bacteria (host) and those from other plant species (non-host) in growth bioassays. We found that on the community level, lettuce rhizosphere bacteria grew better in non-host exudate, but mustard cabbage and maize rhizosphere bacteria grew similarly well in both host and non-host exudate. However, individual bacteria taxa showed strong preferences for exudate from different plant species. The bacterial growth patterns were independent of C and N quantity, suggesting that certain exometabolic compounds may drive the growth patterns. Our results demonstrate that root exudate from a given plant species have the potential to stimulate or suppress soil bacteria and hint at a mechanism that different plant species use to select for their specific suite of rhizosphere bacteria. Furthermore, these findings contribute to our broader understanding of how root exudate quality could be a mechanism that plants use to select for distinct microbial communities in the rhizosphere.

54 ENVIRONMENTAL SCIENCES↗

Specific modulation of the root immune system by a community of commensal bacteria

Significance In natural environments, plants establish intimate interactions with a wide diversity of microbes. It is unknown, however, how microbiota composed of commensal bacteria colonize roots in the face of a sophisticated plant immune system that evolved to recognize microbial-associated molecular patterns. We investigate the interaction between plant immune system function and the root microbiota. We report that root-associated commensal bacteria actively suppress the host immune response in the context of a community. Suppressors and nonsuppressors co-occur in the root microbiome and the presence of the former can enhance the colonization ability of the latter. We highlight the role of a specific sector of the plant immune system and its suppression in gating microbial access to the roots.

59 BASIC BIOLOGICAL SCIENCES↗

Phosphate Availability Modulates Root Exudate Composition and Rhizosphere Microbial Community in a Teosinte and a Modern Maize Cultivar

Domestication and breeding have affected interactions between plants and their microbiomes in ways that are only beginning to be understood but may have important implications for recruitment of rhizosphere microorganisms, particularly under stress conditions. We investigated the responses of a modern maize (Zea mays subsp. mays) cultivar and its wild relative, teosinte (Z. mays subsp. parviglumis), to different phosphate availabilities. We appraised responses of the plant-microbial holobiont to phosphate stresses by profiling root exudate metabolomes, and microbial communities in the root endosphere and rhizosphere. We also performed plate assays to quantify phosphate-solubilizing microorganisms from the rhizosphere. Although root exudate metabolite profiles were distinct between the teosinte and modern maize under high phosphate, both plants shifted exudate compositions in response to phosphate stress toward a common metabolite profile. Root and rhizosphere microbial communities also responded significantly to both plant type and the phosphate availability. A subset of bacterial and fungal taxa were differentially abundant under the different phosphate conditions, with each of the three conditions favoring different taxa. Both teosinte and maize rhizospheres harbored phosphate-solubilizing microorganisms under all growth conditions. These results suggest that the root exudation response to phosphate stress was conserved through the domestication of maize from teosinte, shifting exudation levels of specific metabolites. Although microbial communities also shifted, plate-based assays did not detect selective recruitment of phosphate solubilizers in response to phosphate availability.

59 BASIC BIOLOGICAL SCIENCES↗

Roots selectively decompose litter to mine nitrogen and build new soil carbon

Abstract Plant–microbe interactions in the rhizosphere shape carbon and nitrogen cycling in soil organic matter (SOM). However, there is conflicting evidence on whether these interactions lead to a net loss or increase of SOM. In part, this conflict is driven by uncertainty in how living roots and microbes alter SOM formation or loss in the field. To address these uncertainties, we traced the fate of isotopically labelled litter into SOM using root and fungal ingrowth cores incubated in a Miscanthus x giganteus field. Roots stimulated litter decomposition, but balanced this loss by transferring carbon into aggregate associated SOM. Further, roots selectively mobilized nitrogen from litter without additional carbon release. Overall, our findings suggest that roots mine litter nitrogen and protect soil carbon.

54 ENVIRONMENTAL SCIENCES↗

Embracing fine-root system complexity in terrestrial ecosystem modeling

Projecting the dynamics and functioning of the biosphere requires a holistic consideration of whole-ecosystem processes. However, biases toward leaf, canopy, and soil modeling since the 1970s have constantly left fine-root systems being rudimentarily treated. As accelerated empirical advances in the last two decades establish clearly functional differentiation conferred by the hierarchical structure of fine-root orders and associations with mycorrhizal fungi, a need emerges to embrace this complexity to bridge the data-model gap in still extremely uncertain models. In this work, we propose a three-pool structure comprising transport and absorptive fine roots with mycorrhizal fungi (TAM) to model vertically resolved fine-root systems across organizational and spatial–temporal scales. Emerging from a conceptual shift away from arbitrary homogenization, TAM builds upon theoretical and empirical foundations as an effective and efficient approximation that balances realism and simplicity. A proof-of-concept demonstration of TAM in a big-leaf model both conservatively and radically shows robust impacts of differentiation within fine-root systems on simulating carbon cycling in temperate forests. Theoretical and quantitative support warrants exploiting its rich potentials across ecosystems and models to confront uncertainties and challenges for a predictive understanding of the biosphere. Echoing a broad trend of embracing ecological complexity in integrative ecosystem modeling, TAM may offer a consistent framework where modelers and empiricists can work together toward this grand goal.

59 BASIC BIOLOGICAL SCIENCES↗

Endophyte‐induced systemic spatial reprogramming of metabolism in Populus trichocarpa roots under drought

Beneficial, facultative endophytes help plants thrive in challenging environments by altering their host's metabolism, but how these cellular scale metabolic changes propagate to the systems biology scale is unknown. In this work, we employed a high-resolution chemical imaging approach to map metabolic changes at the Populus trichocarpa root-zone and cell-type levels combined with machine learning (ML) models to identify root metabolites and exudates that have predictive power over treatment class. We found that a nine-strain consortium of beneficial endophytes differentially altered the metabolome of droughted root tissues in a manner specific to cell type and root zone, with endophyte abundance showing a clear correlation to individual metabolites. Our study demonstrates that integrating spatial metabolomics with ML can reveal localized metabolic patterns linked to root–microbe interactions and generate novel hypotheses about underlying biological mechanisms.

Drought↗

Variation in Root Exudate Composition Influences Soil Microbiome Membership and Function

Root exudation is one of the primary processes that mediate interactions between plant roots, microorganisms, and the soil matrix, yet the mechanisms by which exudation alters microbial metabolism in soils have been challenging to unravel. Here, utilizing distinct sorghum genotypes, we characterized the chemical heterogeneity between root exudates and the effects of that variability on soil microbial membership and metabolism. Distinct exudate chemical profiles were quantified and used to formulate synthetic root exudate treatments: a high-organic-acid treatment (HOT) and a high-sugar treatment (HST). To parse the response of the soil microbiome to different exudate regimens, laboratory soil reactors were amended with these root exudate treatments as well as a nonexudate control. Amplicon sequencing of the 16S rRNA gene illustrated distinct microbial diversity patterns and membership in response to HST, HOT, or control amendments. Exometabolite changes reflected these microbial community changes, and we observed enrichment of organic and amino acids, as well as possible phytohormones in the HST relative to the HOT and control. Linking the metabolic capacity of metagenome-assembled genomes in the HST to the exometabolite patterns, we identified microorganisms that could produce these phytohormones. Our findings emphasize the tractability of high-resolution multiomics tools to investigate soil microbiomes, opening the possibility of manipulating native microbial communities to improve specific soil microbial functions and enhance crop production.

59 BASIC BIOLOGICAL SCIENCES↗

Carbon dioxide, water vapor and methane soil efflux (soil respiration) in a Pinus palustris root exclusion in Georgetown, SC

This dataset contains processed data from a combination of survey flux chambers and long-term automated flux chambers. Soil flux measurements were conducted from June 2023 through December 2025 in a mature longleaf pine forest in Georgetown, SC. Soil respiration measurements were conducted approximately biweekly for two and a half years, before and after a root exclusion that took place on May 5, 2024. Processed, QAQC’d data for the treatment (root exclusion) and control (roots intact) before and after the root exclusion can be found in the file: 1_DATA_ESS_DOE_HR_RS_HB2_QAQC_Survey_Data_20260223.csv. Two multiday deployments were also conducted prior to the root exclusion using long-term automated chambers to continuously monitor greenhouse gas soil efflux. Processed, QAQC’d data for both long-term deployments can be found in the file: 2_DATA_ESS_DOE_HR_RS_HB2_QAQC_Longterm_Data_20260209.csv. Raw and working data files (.json, .81x, & .82z format) from LI-COR equipment are included for reference and can be accessed using SoilFluxPro software. CSV metadata files describe the raw data and modifications made using SoilFluxPro v5 and Matlab R2024b, as well as formatting and units for processed CSVs. Matlab code is included for reading in the processed CSVs, with sample figures comparing treatment and control. This research was performed as part of the project: “Improving models of stand and watershed carbon and water fluxes with more accurate representations of soil-plant-water dynamics in southern pine ecosystems”, which examines in part the effects hydraulic redistribution on soil efflux of carbon dioxide, water vapor and methane, as well as soil moisture and temperature in a southern pine ecosystem with sandy soils and high water table.

CARBON DIOXIDE FLUX↗

An integrated Imaging and Modeling Toolbox for Accelerated Development of Root-focused Crops at Field Scales

This project, led by Yuxin Wu at the LBNL, was proposed to develop an integrated imaging-modeling toolbox allowing for accelerated development of root-focused crops at field scales. Our approach is based on a novel root phenotyping method we termed Tomographic Electrical Rhizosphere Imaging (TERI), which provides advanced phenotyping of key root traits by sensing the electrical impedance response of roots and soil to external electrical excitations, and mapping these responses to root and soil properties. The role of Noble Research Institute in this project is to assist TERI model establishment and validation by collecting various above- and below-ground traits data from container- and field-grown plants (mainly wheat). Over the last three years, Noble Research Institute has conducted 3 phases of experiments as proposed including: (Phase 1) greenhouse and hoop-house experiments in wheat and pecan plants grown under controlled container-grown conditions; (Phase 2) space-planted individual plant field experiment using different wheat varieties; and (Phase 3) small-plot planted field experiment using different wheat varieties, as well as using different small grains species including wheat, rye, triticale, oat and barley for increasing plant variation.

60 APPLIED LIFE SCIENCES↗

Root gravitropism in response to a signal originating outside of the cap

We have developed image analysis software linked to a rotating stage, allowing constraint of any user-selected region of a root at a prescribed angle during root gravitropism. This device allows the cap of a graviresponding root to reach vertical while maintaining a selected region within the elongation zone at a gravistimulated angle. Under these conditions gravitropic curvature of roots of Zea mays L. continues long after the root cap reaches vertical, indicating that a signal from outside of the cap can contribute to the curvature response.

Non-NASA Center↗