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DOE OSTI2023

Classical integrability of root-$T\overline{T}$ flows

The root-$T\overline{T}$ flow was recently introduced as a universal and classically marginal deformation of any two-dimensional translation-invariant field theory. The flow commutes with the (irrelevant) $T\overline{T}$ flow, and it can be integrated explicitly for a large class of actions, leading to nonanalytic Lagrangians reminiscent of the four-dimensional modified-Maxwell theory (ModMax). It is not a priori obvious whether the root-$T\overline{T}$ flow preserves integrability, as is the case for the $T\overline{T}$ flow. In this paper we demonstrate that this is the case for a large class of classical models by explicitly constructing a deformed Lax connection. We discuss the principal chiral model and the nonlinear sigma models on symmetric and semisymmetric spaces, without or with the Wess-Zumino term. We also construct Lax connections for the two-parameter families of theories deformed by both root-$T\overline{T}$ and $T\overline{T}$ for all of these models.

72 PHYSICS OF ELEMENTARY PARTICLES AND FIELDS↗

DOE OSTI2020

Serendipita bescii promotes winter wheat growth and modulates the host root transcriptome under phosphorus and nitrogen starvation

Summary Serendipita vermifera ssp. bescii , hereafter referred to as S. bescii , is a root‐associated fungus that promotes plant growth in both its native switchgrass host and a variety of monocots and dicots. Winter wheat ( Triticum aestivum L.), a dual‐purpose crop, used for both forage and grain production, significantly contributes to the agricultural economies of the Southern Great Plains, USA. In this study, we investigated the influence of S. bescii on growth and transcriptome regulation of nitrogen (N) and phosphorus (P) metabolism in winter wheat. Serendipita bescii significantly improved lateral root growth and forage biomass under a limited N or P regime. Further, S. bescii activated sets of host genes regulating N and P starvation responses. These genes include, root‐specific auxin transport, strigolactone and gibberellin biosynthesis, degradation of phospholipids and biosynthesis of glycerolipid, downregulation of ammonium transport and nitrate assimilation, restriction of protein degradation by autophagy and subsequent N remobilization. All these genes are hypothesized to regulate acquisition, assimilation and remobilization of N and P. Based on transcriptional level gene regulation and physiological responses to N or P limitation, we suggest S. bescii plays a critical role in modulating stress imposed by limitation of these two critical nutrients in winter wheat.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2017

Introduction to a Virtual Issue on root traits

Plant traits – ‘morphological, anatomical, physiological, biochem-ical and phenological characteristics of plants and their organs’(Kattge et al., 2011) – are increasingly being harnessed byempiricists and modelers as a framework to understand patternsin the structure and function of specie s across the globe. Trait-basedecology, which emphasizes functional traits over the taxonomicalrelationships among organisms (Laliberte, 2017), promises toimprove generality, synthesis, and predictive ability across ecolog-ical scales (Shipley et al., 2016). Indeed, plant trait studies areincreasingly prominent in the literature: a simple Web of Sciencesearch on the term indicates a surge in publications from 2576during the three-year period from 1999 to 2001 to 13 234 in thethree-year period between 2014 and 2016. However, the mostcommon plant traits described in the literature relate to above -ground organs and their function, including leaf morphology,photosynthetic parameters, and above ground growth rate. Roottraits, particularly those of fine roots associated with criticalbelowground plant functions, are much less studied – they are, afterall, harder to measure and less likely to have a role in ecosystemmodels as they are encoded today. Although the TRY database ofplant traits (Kattge et al., 2011) has been a highly valuable resourcefor plant and ecosystem ecologists, < 1% of the data entries describefine-root functional traits. This glaring gap in our knowledge of thebelowground half of ecosystem function has led to a chorus of pleasin recent years for a stronger emphasis on the measurement andunderstanding of root traits (e.g . Bardgett et al., 2014).

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2016

Enrichment of Root Endophytic Bacteria from Populus deltoides and Single-Cell-Genomics Analysis

ABSTRACT Bacterial endophytes that colonize Populus trees contribute to nutrient acquisition, prime immunity responses, and directly or indirectly increase both above- and below-ground biomasses. Endophytes are embedded within plant material, so physical separation and isolation are difficult tasks. Application of culture-independent methods, such as metagenome or bacterial transcriptome sequencing, has been limited due to the predominance of DNA from the plant biomass. Here, we describe a modified differential and density gradient centrifugation-based protocol for the separation of endophytic bacteria from Populus roots. This protocol achieved substantial reduction in contaminating plant DNA, allowed enrichment of endophytic bacteria away from the plant material, and enabled single-cell genomics analysis. Four single-cell genomes were selected for whole-genome amplification based on their rarity in the microbiome (potentially uncultured taxa) as well as their inferred abilities to form associations with plants. Bioinformatics analyses, including assembly, contamination removal, and completeness estimation, were performed to obtain single-amplified genomes (SAGs) of organisms from the phyla Armatimonadetes , Verrucomicrobia , and Planctomycetes , which were unrepresented in our previous cultivation efforts. Comparative genomic analysis revealed unique characteristics of each SAG that could facilitate future cultivation efforts for these bacteria. IMPORTANCE Plant roots harbor a diverse collection of microbes that live within host tissues. To gain a comprehensive understanding of microbial adaptations to this endophytic lifestyle from strains that cannot be cultivated, it is necessary to separate bacterial cells from the predominance of plant tissue. This study provides a valuable approach for the separation and isolation of endophytic bacteria from plant root tissue. Isolated live bacteria provide material for microbiome sequencing, single-cell genomics, and analyses of genomes of uncultured bacteria to provide genomics information that will facilitate future cultivation attempts.

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2012

3-Dimensional root cause diagnosis via co-analysis. In: ICAC '12 Proceedings of the 9th international conference on Autonomic computing

With the growth of system size and complexity, reliability has become a major concern for large-scale systems. Upon the occurrence of failure, system administrators typically trace the events in Reliability, Availability, and Serviceability (RAS) logs for root cause diagnosis. However, RAS log only contains limited diagnosis information. Moreover, the manual processing is time-consuming, error-prone, and not scalable. To address the problem, in this paper we present an automated root cause diagnosis mechanism for large-scale HPC systems. Our mechanism examines multiple logs to provide a 3-D fine-grained root cause analysis. Here, 3-D means that our analysis will pinpoint the failure layer, the time, and the location of the event that causes the problem.

Zheng, Ziming↗

DOE OSTI2021

De novo transcriptome in roots of switchgrass ( Panicum virgatum L. ) reveals gene expression dynamic and act network under alkaline salt stress

Background: Soil salinization is a major limiting factor for crop cultivation. Switchgrass is a perennial rhizomatous bunchgrass that is considered an ideal plant for marginal lands, including sites with saline soil. Here we investigated the physiological responses and transcriptome changes in the roots of Alamo (alkaline-tolerant genotype) and AM314/MS-155 (alkaline-sensitive genotype) under alkaline salt stress. Results: Alkaline salt stress significantly affected the membrane, osmotic adjustment and antioxidant systems in switchgrass roots, and the ASTTI values between Alamo and AM-314/MS-155 were divergent at different time points. A total of 108,319 unigenes were obtained after reassembly, including 73,636 unigenes in AM-314/MS-155 and 65,492 unigenes in Alamo. A total of 10,219 DEGs were identified, and the number of upregulated genes in Alamo was much greater than that in AM-314/MS-155 in both the early and late stages of alkaline salt stress. The DEGs in AM-314/MS-155 were mainly concentrated in the early stage, while Alamo showed greater advantages in the late stage. These DEGs were mainly enriched in plant-pathogen interactions, ubiquitin-mediated proteolysis and glycolysis/gluconeogenesis pathways. We characterized 1480 TF genes into 64 TF families, and the most abundant TF family was the C2H2 family, followed by the bZIP and bHLH families. A total of 1718 PKs were predicted, including CaMK, CDPK, MAPK and RLK. WGCNA revealed that the DEGs in the blue, brown, dark magenta and light steel blue 1 modules were associated with the physiological changes in roots of switchgrass under alkaline salt stress. The consistency between the qRT-PCR and RNA-Seq results confirmed the reliability of the RNA-seq sequencing data. A molecular regulatory network of the switchgrass response to alkaline salt stress was preliminarily constructed on the basis of transcriptional regulation and functional genes. Conclusions: Alkaline salt tolerance of switchgrass may be achieved by the regulation of ion homeostasis, transport proteins, detoxification, heat shock proteins, dehydration and sugar metabolism. These findings provide a comprehensive analysis of gene expression dynamic and act network induced by alkaline salt stress in two switchgrass genotypes and contribute to the understanding of the alkaline salt tolerance mechanism of switchgrass and the improvement of switchgrass germplasm.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2026

Shaping with water: linking moisture perception to development in plant roots

Water is the most limiting resource for plant growth and development. Heterogeneity in the environmental distribution of water requires plants to direct root growth toward water and to avoid investing resources in areas that lack water. Roots use hydrosignaling pathways—hydrotropism, hydropatterning, and xerobranching—to sense and respond to water availability. While molecular mechanisms of water perception remain unclear, recent studies suggest that organ-level processes using proxies like ethylene help detect spatial water patterns. This review summarizes advances in hydrosignaling and identifies key knowledge gaps to address how plants sense water. Understanding these processes will guide strategies to improve root water capture for sustainable agriculture.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2025

SPRUCE Aboveground Vegetation Coverage in Root Ingrowth Core Plots, Marcell Experimental Forest, Minnesota, August 2022

This dataset contains vegetation survey measurements from root ingrowth core plots (Määttä et al. 2025) inside SPRUCE Experiment plots at the Marcell Experimental Forest in northern Minnesota. Vegetation surveys were conducted in 0.25 meter2 plots containing root ingrowth cores on August 8th and 9th, 2022 (2022-08-08 to 2022-08-09). The warming and elevated carbon dioxide (CO2) treatments in the dataset include the full treatment gradient: +0 degrees Celsius (C) (+0 and +500 parts per million (ppm) elevated CO2), +2.25 degrees C (+0 and +500 ppm), +4.5 degrees C (+0 and +500 ppm elevated CO2), +6.75 degrees C (+0 and +500 ppm) and +9 degrees C (+0 and +500 ppm elevated CO2) for both hummocks and hollows. This dataset includes measurements of the height and absolute coverage (%) for each vascular plant and moss species, as well as organic litter and dead overstory vascular plants, and the distance from the grid center to the nearest tree and the species of the nearest tree. These data were used as species-specific aboveground plant metadata for assessing 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 ↗

DOE OSTI2026

Depth-resolved sagebrush root metabolomics, rhizosphere microbial communities, and geochemistry at the East River Watershed

This data set consists of results from soil nutrient profile, untargeted metabolomics, mass spec imaging, and amplicon sequencing. Data for soil nutrient profile includes common cations (Ca, Mg, Na, and K etc.) extracted from 3 digesting steps – ammonia acetate (for exchangeable cations), nitric acid (for acid dissolved fraction), and hydrofluoric acid/perchloric acid (HF/HClO4) for whole soil digestion. It also includes concentration of organic carbon, inorganic nitrogen (ammonia and nitrate) and phosphorus (Bray-1 P and nitric acid extract), and total nitrogen and phosphorus. Data for untargeted metabolomics includes metabolomic profile for root exudate/tissues and soil extracts from depths at surface soil to saprolite, that were measured using gas chromatography – mass spectrometry (GC-MS), and liquid chromatography – tandem mass spectrometry (LC-MS/MS). Data for mass spec imaging includes spatial distribution of metabolites that were detected and annotated with Fourier transformation ion cyclotron resonance mass spectrometer (FTICR-MS). Data for amplicon sequencing includes the base paired 16S and ITS ribosomal RNA sequences from Miseq Illumina sequencing. All samples were collected from 2 sampling campaign October 2022 and June 2023. Collectively, these datasets enable a mechanistic evaluation of how nutrient acquisition, especially nitrogen and phosphorus, differs between shallow roots operating in soil and deep roots functioning within the fractured bedrock zone. All files are provided as comma-separated values (CSV) fies (.csv) and (GZIP) file (.gz). The compressed .gz FASTQ files can be read directly in R using the dada2 package as part of the amplicon sequence analysis workflow. This work was supported by the Watershed Function Science Focus Area at Lawrence Berkeley National Laboratory funded by the US Department of Energy, Office of Science, Biological and Environmental Research under Contract No. DE-AC02-05CH11231. This research was performed on a project award 60563 (https://dx.doi.org/10.46936/expl.proj.2022.60563/60008727) from the Environmental Molecular Sciences Laboratory, a DOE Office of Science User Facility sponsored by the Biological and Environmental Research program under Contract No. DE-AC05-76RL01830.

EARTH SCIENCE > AGRICULTURE > SOILS > CARBON↗

DOE OSTI2022

ROOT for the HL-LHC: data format

This document discusses the state, roadmap, and risks of the foundational components of ROOT with respect to the experiments at the HL-LHC (Run 4 and beyond). As foundational components, the document considers in particular the ROOT input/output (I/O) subsystem. The current HEP I/O is based on the TFile container file format and the TTree binary event data format. The work going into the new RNTuple event data format aims at superseding TTree, to make RNTuple the production ROOT event data I/O that meets the requirements of Run 4 and beyond.

72 PHYSICS OF ELEMENTARY PARTICLES AND FIELDS↗

DOE OSTI2026

L'Arlesienne de ROOT

Over many years, ROOT users have repeatedly stumbled over—and loudly rediscovered—the infamous 1 GB limit on individual I/O operations, a constraint that somehow survived long past the era when anyone thought a gigabyte was “a lot.” As experiments embraced ever-larger objects and collections, this limit became an increasingly unavoidable rite of passage. This contribution recounts the sustained, multi-year quest by ROOT I/O developers to finally retire this relic, navigating a maze of legacy APIs, memory-management assumptions, and integer boundaries that seemed determined to preserve the status quo. We describe how internal interfaces were carefully modernized to introduce fully 64-bit–capable code paths without breaking the mountains of existing user code that would definitely have noticed. With the limit now lifted, ROOT can finally handle multi-gigabyte objects in a single read or write operation, even when splitting them into an RNTuple is not an option (we’re looking at you, large RooWorkspaces and giant histograms), liberating users from yet another “fun” debugging adventure and clearing the way for the massive analyses of the HL-LHC and beyond.

Canal, Philippe G. [Fermilab] (ORCID:0000000277487↗

DOE OSTI2026

L'Arlesienne de ROOT

Canal, Philippe G. [Fermilab] (ORCID:0000000277487↗

DOE OSTI2019

Chemical and Isotopic Evidence for Organic Matter Sulfurization in Redox Gradients Around Mangrove Roots

Coastal environments like mangrove forests are increasingly recognized as potential hotspots for organic carbon burial, giving them a crucial and yet poorly constrained role in the global carbon cycle. Mangrove sediments are frequently anoxic, which facilitates elevated organic matter (OM) burial via several mechanisms, including sulfurization – abiotic reactions between dissolved (poly)sulfide and OM that decrease its lability. Although sulfurization was estimated to account for roughly half of OM preservation in a Bermuda mangrove forest, both its mechanisms and its global significance remain poorly understood. In this study, we investigate S cycling in mangrove forest sediments from Little Ambergris Cay, Turks and Caicos Islands, an environment with predominantly microbial OM inputs and no major source of terrestrial iron. We characterize the S- and C-isotope composition and organic S speciation of sedimentary OM fractions with varying degrees of resistance to acid hydrolysis, along with other inorganic S phases. Near the surface of a 3-mm-diameter, O 2 -releasing root, abundant organic and elemental S with a 34 S-depleted composition indicates microbial sulfur cycling and OM sulfurization. A mixture of pyrite, elemental S, and organic S form a plaque within the outer 50 μm of the root, which also contains strongly 34 S-depleted sulfate in its xylem. OM sulfurization products in the sediments include both the alkyl sulfides and disulfides associated with the root plaque and more oxidized forms, especially sulfonates. Hydrolysis-resistant organic S in the sediments is consistently 3–5‰ more 34S-enriched than coexisting elemental S, matching the reported kinetic isotope fractionation factor for OM sulfurization via reaction with polysulfides. These sediments also contain a substantial pool of solid-phase, hydrolyzable organic S with a seawater sulfate-like isotope composition, largely in the form of sulfate esters, which may represent excretions from abundant gastropods. The coexistance of sulfurized OM and aerobic macrofauna highlights how understanding spatial scales and/or temporal cycles in local redox state is critical for predicting net OM preservation, especially in dynamic, coastal environments. Future attempts to mechanistically predict changes in carbon storage in coastal systems will benefit from incorporating OM sulfurization as both a sink for microbially produced sulfide and a mechanism for enhanced carbon sequestration.

37 INORGANIC, ORGANIC, PHYSICAL, AND ANALYTICAL CH↗

DOE OSTI2023

Nitrogen addition alters soil fungal communities, but root fungal communities are resistant to change

Plants are colonized by numerous microorganisms serving important symbiotic functions that are vital to plant growth and success. Understanding and harnessing these interactions will be useful in both managed and natural ecosystems faced with global change, but it is still unclear how variation in environmental conditions and soils influence the trajectory of these interactions. In this study, we examine how nitrogen addition alters plant-fungal interactions within two species of Populus - Populus deltoides and P. trichocarpa. In this experiment, we manipulated plant host, starting soil (native vs. away for each tree species), and nitrogen addition in a fully factorial replicated design. After ~10 weeks of growth, we destructively harvested the plants and characterized plant growth factors and the soil and root endosphere fungal communities using targeted amplicon sequencing of the ITS2 gene region. Overall, we found nitrogen addition altered plant growth factors, e.g., plant height, chlorophyll density, and plant N content. Interestingly, nitrogen addition resulted in a lower fungal alpha diversity in soils but not plant roots. Further, there was an interactive effect of tree species, soil origin, and nitrogen addition on soil fungal community composition. Starting soils collected from Oregon and West Virginia were dominated by the ectomycorrhizal fungi Inocybe (55.8% relative abundance), but interestingly when P. deltoides was grown in its native West Virginia soil, the roots selected for a high abundance of the arbuscular mycorrhizal fungi, Rhizophagus. These results highlight the importance of soil origin and plant species on establishing plant-fungal interactions.

59 BASIC BIOLOGICAL SCIENCES↗

DOE OSTI2023

Effects of different cultivation media on root bacterial community characteristics of greenhouse tomatoes

Tomato, as a typical greenhouse crop, is commonly first planted as seedlings in a variety of substrates before being transplanted into soil. However, there is rare research on the characteristics of the bacterial community in tomato roots under this planting mode. In this study, tomatoes were planted in pots containing three different cultivation media, including soil and two types of substrates in a greenhouse, followed by a transplanting treatment. After collecting tomato root samples, high-throughput sequencing and bioinformatic analysis were used to compare the differences in bacterial diversity and functions between tomato roots before and after transplanting in different cultivation media. In total, 702776 sequences were obtained, and the OTUs were belonging to 109 genera, 58 families, 41 orders, 14 classes, and 12 phyla. Among the three cultivation media, the β -diversity was significant, and there was a slight difference in bacterial species diversity along with a large difference in their abundance at the genus level. Soil and both substrates had 79 bacterial genera in common, these genera accounted for 68.70%, 76.70%, and 71.17% of the total genera found in the soil, substrate 1, and substrate 2, respectively. After being transplanted from the two substrates to the soil, the bacterial community structure and abundance exhibited similarities with those found in the soil. Furthermore, based on microbial function prediction, the microbial communities in the two-substrate environment demonstrated a greater potential for promoting growth, while the microbial communities in the soil exhibited a greater tendency to exert their antibacterial potential. Our findings offer theoretical support for the creation of artificially reconstructed microbial communities in greenhouse cultivation.

Zhang, Xinjian↗

DOE OSTI2024

Continuous cropping of potato changed the metabolic pathway of root exudates to drive rhizosphere microflora

For potato production, continuous cropping (CC) could lead to autotoxicity buildup and microflora imbalance in the field soil, which may result in failure of crops and reduction in yield. In this study, non-targeted metabolomics (via liquid chromatography with tandem mass spectrometry (LC–MS/MS)) combined with metagenomic profiling (via high-throughput amplicon sequencing) were used to evaluate correlations between metabolomics of potato root exudates and communities of bacteria and fungi around potato plants to illustrate the impacts of CC. Potato plants were grown in soil collected from fields with various CC years (0, 1, 4, and 7 years). Metabolomic analysis showed that the contents and types of potential autotoxins in potato root exudates increased significantly in CC4 and CC7 plants (i.e., grown in soils with 4 and 7 years of CC). The differentially expressed metabolites were mainly produced via alpha-linolenic acid metabolism in plant groups CC0 and CC1 (i.e., no CC or 1 year CC). The metabolomics of the groups CC4 and CC7 became dominated by styrene degradation, biosynthesis of siderophore group non-ribosomal peptides, phenylpropanoid biosynthesis, and biosynthesis of various plant secondary metabolites. Continuous cropping beyond 4 years significantly changed the bacterial and fungal communities in the soil around the potato crops, with significant reduction of beneficial bacteria and accumulation of harmful fungi. Correlations between DEMs and microflora biomarkers were established with strong significances. These results suggested that continuous cropping of potato crops changed their metabolism as reflected in the plant root exudates and drove rhizosphere microflora to directions less favorable to plant growth, and it needs to be well managed to assure potato yield.

Xing, Yanhong↗

DOE OSTI2016

Cell Wall Ultrastructure of Stem Wood, Roots, and Needles of a Conifer Varies in Response to Moisture Availability

The composition, integrity, and architecture of the macromolecular matrix of cell walls, collectively referred to as cell wall ultrastructure, exhibits variation across species and organs and among cell types within organs. Indirect approaches have suggested that modifications to cell wall ultrastructure occur in response to abiotic stress; however, modifications have not been directly observed. Glycome profiling was used to study cell wall ultrastructure by examining variation in composition and extractability of non-cellulosic glycans in cell walls of stem wood, roots, and needles of loblolly pine saplings exposed to high and low soil moisture. Soil moisture influenced physiological processes and the overall composition and extractability of cell wall components differed as a function of soil moisture treatments. The strongest response of cell wall ultrastructure to soil moisture was increased extractability of pectic backbone epitopes in the low soil moisture treatment. The higher abundance of these pectic backbone epitopes in the oxalate extract indicate that the loosening of cell wall pectic components could be associated with the release of pectic signals as a stress response. The increased extractability of pectic backbone epitopes in response to low soil moisture availability was more pronounced in stem wood than in roots or needles. Additional responses to low soil moisture availability were observed in lignin associated carbohydrates released in chlorite extracts of stem wood, including an increased abundance of pectic arabinogalactan epitopes. Overall, these results indicate that cell walls of loblolly pine organs undergo changes in their ultrastructural composition and extractability as a response to soil moisture availability and that cell walls of the stem wood are more responsive to low soil moisture availability compared to cell walls of roots and needles. In conclusion, to our knowledge, this is the first direct evidence, delineated by glycomic analyses, that abiotic stress affects cell wall ultrastructure. This study is also unique in that glycome profiling of pine needles has never before been reported.

54 ENVIRONMENTAL SCIENCES↗

DOE OSTI2022

Spatial mapping of plant N-glycosylation cellular heterogeneity inside soybean root nodules provided insights into legume-rhizobia symbiosis

Although ubiquitously present, information on the function of complex N-glycan posttranslational modification in plants is very limited and it is often neglected. In this work, we adopted an enzyme-assisted matrix-assisted laser desorption/ionization mass spectrometry imaging strategy to visualize the distribution and identity of N-glycans in soybean root nodules at a cellular resolution. We additionally performed proteomics analysis to probe the potential correlation to proteome changes during symbiotic rhizobia-legume interactions. Our ion images reveal that intense N-glycosylation occurs in the sclerenchyma layer, and inside the infected cells within the infection zone, while morphological structures such as the cortex, uninfected cells, and cells that form the attachment with the root are less N-glycosylated. Notably, we observed different N-glycan profiles between soybean root nodules infected with wild type rhizobia and those infected with mutant rhizobia incapable of efficiently fixing atmospheric nitrogen. The majority of complex N-glycan structures, particularly those with characteristic Lewis-a epitopes, are overexpressed in the mutant nodules. Our proteomic results revealed that these glycans likely originated from proteins that maintain the redox balance crucial for proper nitrogen fixation, but also from enzymes involved in N-glycan and phenylpropanoid biosynthesis. These findings indicate possible involvement of Lewis-a glycans in these critical pathways during legume-rhizobia symbiosis.

59 BASIC BIOLOGICAL SCIENCES↗