From Fuel to Fortress: The Defensive Side of Plant Sugars

From Fuel to Fortress: The Defensive Side of Plant Sugars | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 29 January 2026 V1 Latest version Share on From Fuel to Fortress: The Defensive Side of Plant Sugars Authors : Anna Wlazło , Anna Barczak-Brzyżek , and Marcin Filipecki 0000-0003-4107-2484 [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.176966487.75802140/v1 211 views 96 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract The carbohydrate status of plants encompasses both energy resources (sugars and starch) and structural components of the cell wall, shaped by photosynthesis, environmental stress, and growth demands. Carbohydrates are spatially and temporally regulated through subcellular compartmentation, source–sink transitions, and shifts in sugar composition, all of which critically influence plant health and performance.Beyond their metabolic functions, soluble sugars such as sucrose, glucose, and fructose act as key signaling molecules and that integrate carbohydrate metabolism, transport, and energy status with plant defense responses. Dynamic regulation of sugar transporters and sucrose-cleaving enzymes enables rapid adjustment of carbohydrate allocation, providing both energetic resources and regulatory signals during biotic stress. In addition, the plant cell wall contributes to immunity not only as a physical barrier but also as a dynamic source of immune-eliciting oligosaccharides, thereby linking carbohydrate status to pattern-triggered immunity.This review synthesizes recent advances in understanding how carbohydrates coordinate metabolic, signaling, and structural defenses during plant interactions with pathogens and herbivores. Particular emphasis is placed on sugar-mediated priming of immune responses and the emerging potential of sugars and oligosaccharides as sustainable tools for crop protection and enhanced plant resilience. From Fuel to Fortress: The Defensive Side of Plant Sugars Author information Anna Wlazło 1 , Anna Barczak-Brzyżek 1, * , Marcin Filipecki 1, * 1 Department of Plant Genetics, Breeding and Biotechnology, Institute of Biology, Warsaw University of Life Sciences - SGGW, Poland * Corresponding authors: ABB or MF ORCID and e-mails: AW 0000-0001-5417-4119; [email protected] ABB 0000-0003-2541-427X; [email protected] MF 0000-0003-4107-2484; [email protected] The carbohydrate status of plants encompasses both energy resources (sugars and starch) and structural components of the cell wall, shaped by photosynthesis, environmental stress, and growth demands. Carbohydrates are spatially and temporally regulated through subcellular compartmentation, source–sink transitions, and shifts in sugar composition, all of which critically influence plant health and performance. Beyond their metabolic functions, soluble sugars such as sucrose, glucose, and fructose act as key signaling molecules and that integrate carbohydrate metabolism, transport, and energy status with plant defense responses. Dynamic regulation of sugar transporters and sucrose-cleaving enzymes enables rapid adjustment of carbohydrate allocation, providing both energetic resources and regulatory signals during biotic stress. In addition, the plant cell wall contributes to immunity not only as a physical barrier but also as a dynamic source of immune-eliciting oligosaccharides, thereby linking carbohydrate status to pattern-triggered immunity. This review synthesizes recent advances in understanding how carbohydrates coordinate metabolic, signaling, and structural defenses during plant interactions with pathogens and herbivores. Particular emphasis is placed on sugar-mediated priming of immune responses and the emerging potential of sugars and oligosaccharides as sustainable tools for crop protection and enhanced plant resilience. Keywords: carbohydrate metabolism, carbohydrate transport, signal transduction, cell wall metabolism, host-pathogen interactions, herbivory, plant defense responses Author contributions: AW and ABB wrote the original manuscript and prepared the figures; MF came up with the idea, revised the manuscript. Acknowledgements The focus of our research on sugars developed in parallel with studies on plant interactions with spider mite herbivores, conducted within the framework of projects no. 2019/33/B/NZ9/01305 and 2024/53/N/NZ9/01802, funded by the Polish National Science Centre. Introduction In natural environments, plants compete for essential resources such as light, water, and mineral nutrients to sustain photosynthesis and growth, while simultaneously facing continuous pressure from heterotrophic organisms, including pathogens and herbivores. Although plants have evolved a diverse array of defense strategies, their attackers have likewise developed sophisticated mechanisms to circumvent these defenses and exploit the carbon-rich products of photosynthesis. Over time, accumulating experimental evidence has revealed that sugars—traditionally viewed as metabolic energy sources or passive targets for invaders—are active regulators of plant defense, modulating immune responses at both physiological and molecular levels (Horsfall and Dimond 1957). Upon biotic attack, sugar concentrations are dynamically regulated by sucrose-cleaving enzymes and redistributed through specialized transporters. These spatial and temporal changes in carbohydrate allocation act as signaling cues that coordinate immune responses, including transcriptional reprogramming of defense-related genes and modulation of phytohormone pathways. In addition, sugars support defense-associated developmental and anatomical adaptations and provide substrates for the biosynthesis of deterrent compounds, contributing to both physical and chemical resistance mechanisms. While the role of sugars in plant–pathogen interactions has been extensively investigated, their involvement in defense against herbivores remains comparatively underexplored. In this review, we reposition sugars not as static metabolic resources but as central regulators of plant immunity, integrating metabolism, transport, signaling, and structural defenses that contribute to resistance against both pathogens and herbivores. Sugar dynamics at the plant–heterotroph interface: energy, competition, and signaling In higher plants, sucrose is the principal long-distance transport carbohydrate, owing to its non-reducing nature, metabolic stability, and high energy content. Beyond its role as a carbon source, sucrose functions as a signaling molecule that regulates local and systemic responses to environmental stress, including acclimation and defense processes (Koch 1996; Lemoine 2000; Wind et al. 2010). Sucrose transport through the phloem involves either apoplastic or symplastic loading and unloading, enabling the distribution of photoassimilates from source tissues, such as mature leaves, to sink tissues including roots, stems, storage organs, and reproductive structures (Rennie and Turgeon 2009). Under biotic stress, particularly during pathogen or herbivore attack, carbohydrate allocation undergoes extensive reprogramming and represents a key component of the growth–defense trade-off (Huot et al. 2014; Züst and Agrawal 2017). These stresses typically suppress photosynthesis-associated gene expression, reduce photosynthetic efficiency, and increase respiratory demand. As a consequence, affected tissues often shift from source to sink status, accompanied by profound metabolic reorganization (Bilgin et al. 2010; Kanwar and Jha 2019). Sugar levels fluctuate dynamically during defense responses, reflecting intense competition between host plants and their attackers for access to carbohydrates. In many plant–pathogen interactions, the apoplast serves as a nutrient-rich niche that pathogens exploit during colonization (Naseem et al. 2017). To restrict pathogen access to apoplastic sugars, plants tightly regulate carbohydrate transport, metabolism, and signaling, thereby limiting nutrient availability and suppressing pathogen growth. Pathogens, in turn, deploy diverse virulence strategies, including effector proteins that interfere with host sugar transport and immune signaling. Consequently, activation of plant defense responses is closely associated with large-scale and dynamic changes in host metabolism (Jones and Dangl 2006). Sugars thus emerge as key targets of manipulation on both sides of the plant–heterotroph interaction (Bezrutczyk et al. 2018; Ponzio et al. 2017; Ali et al. 2024). First, as readily accessible energy sources, sugars support heterotrophic growth and facilitate pathogen or herbivore development (Naseem et al. 2017). Second, sugars act as signaling molecules: elevated sugar levels can induce defense-related gene expression, modulate phytohormone signaling networks, and stimulate the production of secondary metabolites that enhance resistance (Jeandet et al. 2022; Yamada and Mine 2024). Third, sugars provide the metabolic energy required for highly demanding defense processes, including cell wall remodeling, callose deposition, and trichome formation, enabling plants to mount effective physical barriers against invaders (Tauzin and Giardina 2014; Wei et al. 2025). The diversity of pathogen and herbivore lifestyles further shapes carbohydrate-mediated defense strategies, particularly through interactions with stress hormones (Liu et al. 2022). Biotrophic pathogens predominantly activate salicylic acid (SA)–dependent signaling, whereas necrotrophic and hemibiotrophic pathogens preferentially induce jasmonic acid (JA) and ethylene (ET) pathways, which often antagonize SA responses (Glazebrook 2005; Thaler et al. 2004). A similar dichotomy is observed in plant–herbivore interactions: sap-sucking insects mainly trigger SA-associated defenses, whereas chewing herbivores and mites induce broader responses dominated by JA and frequently involving ET, SA, and abscisic acid (ABA) signaling (Kant et al. 2004; Moran and Thompson 2001; Vos et al. 2013; Glas et al. 2014; Kiełkiewicz et al. 2019; Ali et al. 2024). Despite substantial progress in characterizing hormone-mediated defense pathways, their integration with sugar signaling remains incompletely understood. One notable example is the ABA–glucosyl ester, the major inactive storage form of ABA, which may represent an important regulatory node linking carbohydrate metabolism and hormone signaling (Lee et al. 2006). More broadly, glucosylation of stress-related phytohormones can modulate their activity, stability, and transport, suggesting that sugars play an underappreciated role in fine-tuning hormonal defense networks (Ostrowski and Jakubowska 2014). Molecular mechanisms of sucrose allocation: the role of SWEET, SUT, and STP transporters in plant–pathogen and herbivore interactions Sucrose, a disaccharide composed of glucose and fructose, is the primary product of photosynthesis and the major transport carbohydrate in most plant species. Its allocation relies on a coordinated network of sugar transporters, including sugars will eventually be exported transporters (SWEETs) , sucrose transporters/sucrose carriers (SUTs/SUCs) , and sugar transport proteins (STPs), which collectively regulate sucrose and hexose movement across cellular membranes and tissues. In source tissues, sucrose is transported symplastically through plasmodesmata to phloem parenchyma cells, where clade III SWEETs mediate sucrose efflux into the apoplast (Chen et al. 2012; Geiger 2020). Sucrose is subsequently actively loaded into the sieve element–companion cell complex by proton-coupled SUT/SUC symporters (Riesmeier et al. 1994; Barker et al. 2000). In sink tissues, sucrose unloading occurs via SWEETs or SUTs and is followed either by direct uptake through SUTs or by extracellular hydrolysis into glucose and fructose, which are imported by clade I/II SWEETs or STPs (Sauer et al. 1990; Büttner et al. 2000; Li et al. 2022). Sugars can be transiently stored in the vacuole, with sucrose transported by tonoplast sugar transporters (TSTs) and hexoses by tonoplast monosaccharide transporters (TMTs), early responsive to dehydration (ERD) uniporters, or clade IV SWEETs. Vacuolar sugar release is mediated by proton-coupled transporters such as VGT1 for hexoses and SUT4 for sucrose (Wormit et al. 2006; Schulz et al. 2011; Eom et al. 2015). This dynamic regulation of sugar partitioning is essential for balancing growth, storage, and defense demands. Compelling evidence demonstrates that sugar transporters are major determinants of plant susceptibility or resistance to pathogens, as many microbes actively manipulate host sugar transport to redirect carbon fluxes for their own benefit (Chen et al. 2010; Bezrutczyk et al. 2018; Naseem et al. 2017; Pommerrenig et al. 2020; Liu et al. 2022; Khanna et al. 2023; Lei et al. 2025) (Figure 1). A landmark example is provided by bacterial blight of rice, in which Xanthomonas species deliver transcription activator-like effectors (TALEs) that bind directly to SWEET gene promoters, inducing SWEET expression and elevating apoplastic sugar availability to promote infection (Chen et al. 2010; Römer et al. 2010; Gupta et al. 2021). The classical resistance locus xa13 was later shown to encode OsSWEET11, whose altered promoter prevents TALE binding and confers resistance (Chu et al. 2006; Yang et al. 2006). Similar promoter-based resistance mechanisms have been identified in cotton, where mutations in GhSWEET10 disrupt recognition by Xanthomonas citri TALEs and restrict pathogen access to sugars (Cox et al. 2017). Altered sugar partitioning also influences disease development in other systems. During Plasmodiophora brassicae infection of Arabidopsis roots, gall formation correlates with induction of SWEET11 and SWEET12, while gall development is impaired in sweet11/sweet12 double mutants (Walerowski et al. 2018). Interestingly, enhanced sugar accumulation in these mutants also promotes SA–dependent defense priming, suggesting that modified sugar allocation can simultaneously restrict pathogen nutrition and activate immunity (Gebauer et al. 2017). Importantly, increased SWEET expression does not invariably result in enhanced susceptibility. During Pythium irregulare infection of Arabidopsis roots, SWEET2 sequesters cytosolic sugars into the vacuole, thereby limiting apoplastic sugar availability. Consequently, sweet2 loss-of-function mutants exhibit increased susceptibility (Breia et al. 2021). More generally, plants counteract pathogen-induced sugar efflux by inducing STP transporters that reabsorb hexoses from the apoplast into the cytosol. This mechanism has been documented for STP4 during Erysiphe cichoracearum infection and STP13 during Botrytis cinerea infection (Fotopoulos et al. 2003; Lemonnier et al. 2014). Notably, STP13 physically associates with the flagellin receptor FLS2 and its co-receptor BAK1, and phosphorylation by BAK1 enhances monosaccharide uptake, contributing to the so-called pathogen starvation strategy (Yamada et al. 2016; Bezrutczyk et al. 2018; Chen et al. 2023). Figure 1. Dynamic regulation of sugar transport and metabolism during pathogen attack. (A) Pathogen-favored processes include the induction of sugar efflux transporters, such as clade III sugars will eventually be exported transporters (SWEETs), which are activated by pathogens such as Xanthomonas via transcription activator–like effectors (TALEs) and mediate sucrose release into the apoplast. In parallel, cell wall invertases (CW-Invs) hydrolyze sucrose into glucose and fructose, increasing the availability of accessible carbon sources for invading pathogens. (B) Plant counter-defense mechanisms involve the induction of sugar transporters, including sugar transport proteins (STPs), which promote monosaccharide uptake into the cytosol, and clade IV SWEETs, which facilitate vacuolar import of sugars, thereby limiting apoplastic nutrient availability. Blue circles represent glucose, dark blue triangles represent fructose, and combined symbols represent sucrose. Abbreviations: C, cytosol; N, nucleus; V, vacuole; AS, apoplastic space. Sugar transporter manipulation is also prominent in plant–herbivore interactions (Figure 2). Phloem-feeding insects frequently modulate source–sink relations to enhance nutrient acquisition. For example, infestation of rice by the brown planthopper ( Nilaparvata lugens ) induces expression of OsSUT1 , OsSUT2 , OsSWEET13 , and OsSWEET14 , increasing sugar concentrations in phloem sap and facilitating insect feeding (Xiao et al. 2024; Yu et al. 2024). In Arabidopsis , early infestation by the green peach aphid ( Myzus persicae ) induces several AtSUC genes, supplying energy to damaged tissues; however, plants subsequently downregulate AtSUC expression to limit sugar loss once aphids establish feeding sites (Dubey et al. 2013). Consistently, aphid performance is reduced on suc2 mutants. Chewing herbivores and mechanical damage also influence sugar transport. In Arabidopsis , AtSUC3 expression is strongly induced by wounding (Meyer et al. 2004), while in rice, OsSUT4 expression increases following infestation by rice leaffolder larvae ( Cnaphalocrocis medinalis ) (Chang et al. 2019). This response depends on JA and ABA signaling and is modulated by tissue sink–source status, underscoring the integration of sugar transport with hormonal defense networks. In contrast to antagonistic interactions, sugars also function as exchange currencies in symbiotic relationships. During arbuscular mycorrhizal associations, coordinated regulation of SWEETs, monosaccharide transporters, and cell wall invertases (CW-Invs) enhances sink strength in colonized roots and facilitates carbon transfer to the fungal partner (Helber et al. 2011; Doidy et al. 2012). In tomato, suppression of SlSUT2 increases carbohydrate retention in the peri-arbuscular space and promotes colonization by Funneliformis mosseae (Bitterlich et al. 2014), illustrating how sugar transporters shape outcomes across the mutualism–parasitism continuum. Sugar-cleaving enzymes in plants: from metabolism to immunity Sucrose cleavage represents a pivotal metabolic step linking carbohydrate allocation with plant development and defense. In plants, sucrose is primarily cleaved by invertases (Invs) and sucrose synthase (SUS) . Invs irreversibly hydrolyze sucrose into glucose and fructose, whereas SUS catalyzes the reversible conversion of sucrose and UDP into fructose and UDP-glucose (Koch 2004). Invertases are classified into acid invertases, including CW-Invs and vacuolar invertases (V-Invs), which operate optimally at acidic pH, and alkaline/neutral invertases (A/N-Invs) localized to the cytosol, plastids, and mitochondria (Ruan 2014). SUS occurs predominantly as a cytosolic phosphorylated enzyme but also associates with the plasma membrane in its non-phosphorylated form, particularly in tissues undergoing active cell wall biosynthesis (Brill et al. 2011). Functionally, invertases dominate sucrose breakdown in mature tissues to support respiration, whereas SUS is closely associated with sink strength in rapidly growing or hypoxic tissues such as developing seeds and fruits (Xu et al. 2012; Chibbar et al. 2016). Among sucrolytic enzymes, CW-Invs have emerged as central regulators of plant defense. By altering the apoplastic sucrose-to-hexose ratio, CW-Invs activate sugar signaling pathways that intersect with immune responses (Proels and Hückelhoven 2014). Numerous studies have documented transcriptional induction of CW-Inv genes during pathogen attack (Roitsch et al. 1995; Zhang et al. 1997). Additional post-translational regulation occurs through proteinaceous Inv inhibitors, whose pH-dependent binding fine-tunes enzyme activity in response to apoplastic conditions (Hothorn et al. 2010; Bonfig et al. 2010). Enhanced CW-Inv activity has been observed across a broad spectrum of plant–pathogen interactions, including those involving bacteria, fungi, viruses, oomycetes, and nematodes (Proels and Hückelhoven 2014; Tauzin and Giardina 2014). This induction often correlates with activation of classical defense markers such as pathogenesis–related (PR) genes and WRKY transcription factors. For example, overexpression of CW-Invs in tobacco and rice leads to constitutive PR gene expression and increased resistance to Potato virus Y and Magnaporthe oryzae , respectively (Herbers et al. 1996; Sun et al. 2014). Conversely, silencing CW-Invs compromises PR gene induction and increases susceptibility, as shown for Phytophthora nicotianae infection in tobacco (Essmann et al. 2008). CW-Inv activity is frequently coordinated with monosaccharide transport during defense responses (Figure 1). Concurrent induction of CW-Invs and STP transporters has been reported in several systems, including wheat– Blumeria graminis , grapevine– Erysiphe necator , and Arabidopsis – Erysiphe cichoracearum interactions (Fotopoulos et al. 2003; Sutton et al. 2007; Hayes et al. 2010). This coordination ensures efficient hydrolysis of apoplastic sucrose and rapid uptake of hexoses, thereby optimizing carbohydrate availability for defense while limiting pathogen access. Notably, Xanthomonas campestris pv. vesicatoria has evolved effectors that directly target CW-Invs to suppress host immunity in pepper, underscoring the strategic importance of sucrose cleavage in defense regulation (Sonnewald et al. 2012). Beyond pathogen defense, CW-Invs also play context-dependent roles in plant–herbivore interactions (Figure 2). In hybrid poplar, caterpillar feeding and jasmonate treatment induce CW-Inv activity in developing leaves, enhancing carbon import from source tissues to fuel defense metabolite production (Arnold and Schultz 2002; Arnold et al. 2004). Limiting carbohydrate transport reduces defense compound accumulation, revealing a carbon-driven growth–defense trade-off. Similar induction of CW-Inv activity has been observed in Arabidopsis following methyl jasmonate treatment (Ferrieri et al. 2012, 2013). Conversely, suppression of CW-Inv activity can enhance resistance by restricting nutrient availability to herbivores. In Nicotiana attenuata , the CW-Inv inhibitor Na CWII is strongly induced after Manduca sexta feeding, reducing apoplastic sucrose hydrolysis and limiting larval performance (Ferrieri et al. 2015). In tomato, M. sexta attack triggers transcriptional repression of sugar metabolism and transport genes, lowering sugar availability at feeding sites and impairing larval growth (Ke et al. 2021). Together, these findings indicate that CW-Invs contribute to herbivore defense through two complementary strategies: activation to support defense metabolism or inhibition to restrict nutritional resources. Figure 2. Dynamic regulation of sugar transport and metabolism during herbivore attack. (A) Herbivore-favored processes include activation of apoplastic sucrose efflux mediated by clade III sugars will eventually be exported transporters (SWEETs) and enhanced phloem loading via sucrose transporters (SUTs). Together, these processes increase sugar availability in the apoplast and phloem sap at the feeding site. Cell wall invertases (CW-Invs) further hydrolyze sucrose into glucose and fructose, facilitating nutrient uptake by the herbivore. (B) Plant defense responses include downregulation of SUT expression to limit sucrose allocation to the phloem and induction of cell wall invertase inhibitors (CWIIs), which suppress CW-Inv activity and restrict sugar release into the apoplast. Blue circles indicate glucose, dark blue triangles indicate fructose, and combined symbols represent sucrose. Abbreviations: C, cytosol; N, nucleus; AS, apoplastic space. Compared with CW-Invs, the role of A/N-Invs in plant defense remains less well understood. Reduced expression of a wheat A/N-Inv is associated with decreased susceptibility to stripe rust ( Puccinia striiformis ), while coordinated increases in CW-Inv and A/N-Inv activities have been reported in several plant–pathogen systems, suggesting integrated regulation of sucrose cleavage across cellular compartments (Storr and Hall 1992; Park et al. 2013; Liu et al. 2015). Invertases also contribute to plant tolerance and compensatory growth following herbivory. By supplying glucose to the oxidative pentose phosphate pathway, invertases support regrowth and metabolic recovery after tissue damage. This role has been demonstrated in Arabidopsis through comparative analyses of invertase mutants and natural accessions differing in compensatory capacity (Siddappaji et al. 2015). Transcriptomic studies of spider mite infestation further implicate invertases and sugar sensors such as AtRGS1 in early defense and metabolic reprogramming responses (Zhurov et al. 2014), although disentangling defensive from exploitative functions remains a challenge in large-scale omics analyses. Crosstalk of sugar sensing and energy signaling in plant–pathogen and plant–herbivore interactions Effective coordination of metabolism, growth, and defense depends on a plant’s ability to accurately sense its carbohydrate and energy status. Because downstream physiological and immune responses rely on these inputs, plants continuously monitor sugar availability through multiple sensing mechanisms operating at the plasma membrane, in the cytosol, and within intracellular organelles. This multilevel surveillance enables tight integration of carbon status with stress-responsive signaling networks. The first intracellular sugar sensor identified in plants was hexokinase 1 (HXK1) , a mitochondrion-associated enzyme that functions not only in glucose phosphorylation but also in glucose signaling (Moore et al. 2003). Additional candidates for intracellular glucose sensing include AtHXK3 and AtHXK-like1, whereas sucrose sensing has been proposed to involve SUS , and fructose perception has been linked to nuclear-localized fructose-1,6-bisphosphatase (FINS1) and fructokinase-like proteins (FLN1/FLN2) (Cho and Yoo 2011; Karve et al. 2008; Ruan 2012). Extracellular sugar perception is mediated by plasma membrane–associated sensors, most notably regulator of G-protein signaling 1 (RGS1) , which responds to glucose, fructose, and sucrose, and AtSUT2 , proposed to function as a sucrose sensor in sieve elements (Barker et al. 2000; Huang et al. 2006). Despite clear roles in sugar signaling, direct involvement of these receptors in plant defense has been demonstrated in relatively few cases. Notably, pretreatment of Arabidopsis leaves with glucose enhances resistance to bacterial infection in an HXK1-dependent manner, indicating that glucose perception is required for sugar-induced immune activation (Jing et al. 2020). Beyond individual sugar sensors, plants integrate carbohydrate availability with cellular energy status through two evolutionarily conserved protein kinases: sucrose non-fermenting related kinase 1 (SnRK1) and target of rapamycin (TOR) . These kinases function as central metabolic switches that antagonistically regulate growth and stress responses (Li and Sheen 2016; Li et al. 2021). SnRK1 is activated under conditions of low energy or nutrient availability and promotes catabolic processes and stress-adaptive responses, while repressing energy-intensive anabolic pathways. In contrast, TOR is activated under nutrient-rich conditions and stimulates growth, protein synthesis, and biosynthetic metabolism. Sugar-derived signals tightly regulate both kinases: trehalose-6-phosphate inhibits SnRK1 activity in response to high sucrose levels, whereas glucose-dependent mitochondrial energy signaling activates TOR (Zhang et al. 2009; Xiong et al. 2013). Accumulating evidence positions SnRK1 as a major integrator of sugar and energy signaling in plant defense. Activation of SnRK1 shifts metabolism from a growth-promoting to a stress-resilient state by inducing autophagy, mobilizing stored resources, suppressing sugar and protein biosynthesis, and activating defense-related gene expression, including PR genes (Hulsmans et al. 2016; Xu et al. 2025). Functional studies support a positive role for SnRK1 in resistance against diverse pathogens. In Arabidopsis , silencing AtSnRK1.1 increases susceptibility to clubroot disease caused by Plasmodiophora brassicae , whereas overexpression enhances resistance (Chen et al. 2021). In pepper, SnRK1 is required for hypersensitive response (HR) induction following infection by Xanthomonas campestris pv. vesicatoria ( Xcv ), while the bacterial effector AvrBsT suppresses HR by targeting host SnRK1 signaling components (Szczesny et al. 2010). Similarly, in rice, OsSnRK1a positively regulates resistance to both fungal and bacterial pathogens, and its activity is negatively regulated by K63-linked ubiquitination mediated by OsUbc13 (Filipe et al. 2018; Liu et al. 2023). SnRK1 also modulates immunity through phosphorylation of transcription factors and metabolic regulators. In barley and maize, SnRK1 phosphorylates specific WRKY transcription factors, targeting them for degradation and thereby restricting expression of genes involved in carbohydrate transport. This mechanism limits nutrient availability in the apoplast and enhances resistance to fungal pathogens such as powdery mildew ( Blumeria graminis ) and head smut ( Sporisorium reilianum ) (Han et al. 2020; Zhang et al. 2024). In rice, SnRK1-mediated phosphorylation of the ATPase XB24 enhances defense against Ustilaginoidea virens , while the fungal effector SCRE1 disrupts this interaction to suppress immunity (Yang et al. 2022a). SnRK1 also contributes to defense responses against herbivores, although its roles appear to be more context-dependent. In Arabidopsis infested by the green peach aphid ( Myzus persicae ), expression of several SnRK1 target genes is downregulated without major changes in SnRK1 transcript levels, suggesting post-transcriptional regulation of SnRK1 activity during infestation (Appel et al. 2014). In contrast, feeding by the corn leaf aphid ( Rhopalosiphum maidis ) induces expression of ZmSnRK1.3 in maize, potentially through WRKY transcription factors activated by herbivory (Akbudak et al. 2025). In Nicotiana attenuata , attack by Manduca sexta larvae leads to downregulation of a SnRK1 β-subunit in source leaves, resulting in reallocation of sugars to sink tissues such as roots. This shift promotes root growth and has been proposed to enhance tolerance rather than resistance to herbivory, illustrating how SnRK1-mediated sugar signaling can support alternative defense strategies (Schwachtje et al. 2006). In contrast to SnRK1 , TOR signaling is generally suppressed during biotic stress, and inhibition of TOR often enhances plant resistance. Arabidopsis mutants with impaired TOR complex function exhibit increased resistance to the oomycete Hyaloperonospora arabidopsidis and the fungus Fusarium graminearum , a phenotype attributed to constitutive defense priming and accelerated immune activation (Meteignier et al. 2017; Aznar et al. 2018). Similarly, in rice, TOR acts as a negative regulator of immunity against a broad range of pathogens, including Xanthomonas oryzae , Rhizoctonia solani , and Cochliobolus miyabeanus (De Vleesschauwer et al. 2017). In tomato, pharmacological inhibition of TOR enhances resistance to the necrotrophic fungus Botrytis cinerea (Marash et al. 2022). Together, these findings support a model in which repression of TOR signaling during infection prioritizes defense over growth and biosynthesis. Collectively, these studies identify sugar and energy sensing as central hubs linking carbohydrate availability with stress-adaptive signaling pathways. Through antagonistic regulation by SnRK1 and TOR, plants dynamically balance growth, metabolism, and defense in response to pathogens and herbivores. This integration provides a mechanistic framework for understanding how carbohydrate status shapes immune competence and stress resilience (Rolland et al. 2006; Morkunas and Ratajczak 2014). Plant cell wall remodeling in plant–pathogen and plant–herbivore interactions: constructing the “sweet fortress” The plant cell wall constitutes the first physical barrier that pathogens and herbivores must overcome to access plant-derived carbon resources (Wan et al. 2021; Pérez-Alonso et al. 2025). In plant–heterotroph interactions, both the biochemical composition and mechanical properties of the cell wall critically shape the outcome of attack. Carbohydrates play a dual role in this context: while heterotrophs seek to exploit soluble sugars as energy sources, access to these carbohydrates is restricted by the complex, polysaccharide-rich architecture of the wall. To breach this barrier, pathogens and herbivores secrete suites of cell wall–degrading enzymes (CWDEs), including cellulases, pectinases, xylanases, and xyloglucanases, which weaken wall integrity and release soluble nutrients (Mary Wanjiru et al. 2002). Plants, in turn, have evolved surveillance systems that detect perturbations in cell wall integrity and initiate defense signaling (Bacete and Hamann 2020). Central to this response is the perception of damage-associated molecular patterns (DAMPs) and herbivore-associated molecular patterns (HAMPs), many of which are carbohydrate-derived oligosaccharides generated during wall degradation. These molecular patterns are recognized by pattern-recognition receptors (PRRs) at the plasma membrane, triggering pattern-triggered immunity (PTI) (Jones and Dangl 2006). Among the best-characterized PRRs involved in carbohydrate perception are LysM-containing receptor kinases such as CERK1 and LYK5, which bind oligosaccharide ligands and initiate downstream signaling cascades (Miya et al. 2007; Cao et al. 2014; Yang et al. 2022b). Ligand binding activates receptor-like cytoplasmic kinases, including BIK1, which in turn stimulate mitogen-activated protein kinase (MAPK) cascades, calcium influx, reactive oxygen species (ROS) production via NADPH oxidases, and transcriptional reprogramming of defense-related genes (Veronese et al. 2006; Sun and Zhang 2020). These early signaling events integrate with phytohormone pathways and metabolic reprogramming to reinforce cell wall defenses and restrict further invasion. From the attacker’s perspective, CWDE activity is essential for host colonization. For example, Magnaporthe oryzae secretes the GH12 endoglucanases MoCel12A and MoCel12B, which degrade mixed-linkage glucans in the rice cell wall during infection (Yang et al. 2021). The resulting cellulose-derived oligosaccharides, including cellodextrins, act as potent elicitors of plant immunity. In grapevine, β-1,4-linked cellodextrins induce cytosolic calcium influx, ROS production, and expression of PR genes, ultimately enhancing resistance to Botrytis cinerea (Aziz et al. 2007). Similarly, xyloglucan-derived oligosaccharides activate MAPKs, callose synthase, and defense gene expression in Arabidopsis , conferring increased resistance to B. cinerea (Souza et al. 2017). Pectin degradation represents another major source of immune-eliciting carbohydrates. Hydrolysis of homogalacturonan generates oligogalacturonides (OGs), which function as canonical DAMPs and activate PTI in multiple plant species (Aziz et al. 2004; Denoux et al. 2008; Galletti et al. 2008). OG signaling enhances resistance to necrotrophic pathogens by promoting ROS accumulation, callose deposition, and hormone-dependent defense gene expression. These examples underscore how carbohydrate fragments released during wall degradation serve not only as by-products of attack but also as critical signals that amplify immune responses. Cell wall composition and remodeling are equally important in plant–herbivore interactions, where mechanical resistance and digestibility directly influence feeding behavior (Pérez-Alonso et al. 2025). Alterations in polysaccharide composition can impede stylet penetration by sap-feeding insects and reduce the efficiency of CWDEs present in herbivore saliva (Divol et al. 2005; Mohase and Taiwe 2015). In Arabidopsis , recessive mutations in CESA3 , a primary cell wall cellulose synthase, confer increased resistance to the green peach aphid ( Myzus persicae ), a phenotype associated with elevated JA and ET levels and induction of defense-related genes (Ellis et al. 2002; Divol et al. 2005). Conversely, aphid infestation induces expression of multiple CESA , pectin biosynthesis, and xyloglucan endotransglycosylase/hydrolase ( XTH ) genes, reflecting dynamic remodeling of the wall during feeding. Beyond cellulose, modifications of hemicellulose strongly influence resistance. In Arabidopsis , disruption of α-xylosidase activity alters xyloglucan structure and enhances resistance to Plectosphaerella cucumerina (Sampedro et al. 2010; Delgado-Cerezo et al. 2012). In barley, overexpression of xylan biosynthetic genes increases resistance to powdery mildew, whereas defects in hemicellulose acetylation, as observed in rwa2 mutants, confer enhanced resistance to B. cinerea (Chowdhury et al. 2017; Manabe et al. 2013). These findings highlight how subtle chemical modifications of wall polysaccharides can substantially alter susceptibility to biotic stress. Pectin content and modification further shape plant defense outcomes. Arabidopsis mutants defective in rhamnogalacturonan I biosynthesis ( gae1 gae6 ) display reduced resistance to Pseudomonas syringae and B. cinerea , accompanied by hypersensitivity to JA, indicating a close link between pectin structure and hormone signaling (Bethke et al. 2016). In contrast, mutants with increased pectin content ( pmr5 , pmr6 ) exhibit enhanced resistance to powdery mildew (Vogel et al. 2002, 2004). The degree of pectin methylesterification also plays a pivotal role: high methylesterification generally increases resistance to pathogens, whereas demethylated pectin is more susceptible to pathogen-derived pectinases. Consistently, disruption of AtPME3 increases pectin methylesterification and enhances resistance to B. cinerea and Pectobacterium carotovorum (Lionetti et al. 2012). In herbivore interactions, however, pectin modifications can have opposing effects. Reduced pectin methylesterase activity in tobacco increases susceptibility to chewing insects such as Manduca sexta , which gain more weight when feeding on plants with softened cell walls (Körner et al. 2009). Similarly, impairment of hemicellulose O-acetylation increases susceptibility to Spodoptera exigua larvae, emphasizing that optimal wall rigidity and composition differ between pathogen and herbivore defense contexts (Sun et al. 2020). Callose, a β-1,3-glucan, represents a highly dynamic cell wall component that plays a central role in early defense responses despite its relatively low abundance. Callose is rapidly deposited at papillae in response to pathogen penetration attempts and contributes to the regulation of symplastic connectivity by restricting plasmodesmatal transport (Germán et al. 2023; Li et al. 2021). In plant–insect interactions, phloem-feeding insects such as aphids trigger calcium-dependent callose deposition, reducing phloem sap loss and limiting nutrient acquisition. Overexpression of the cotton callose synthase GhCalS5 enhances resistance to cotton aphid ( Aphis gossypii ), illustrating the defensive potential of callose-mediated wall reinforcement (Kuśnierczyk et al. 2008; Mbiza et al. 2022). In addition to modifying existing wall structures, plants deploy specialized epidermal outgrowths such as trichomes as part of their defensive arsenal (Wang et al. 2021). Trichome development is increasingly recognized as being regulated by carbohydrate availability, linking sugar status with structural defense formation (Wei et al. 2025). Changes in sugar levels in mature leaves can influence both local and systemic trichome density and morphology, integrating metabolic status with ET signaling pathways to deter herbivores. Glandular trichomes provide an additional layer of carbohydrate-based defense by producing specialized metabolites, including O-acylsugars. Studies in Nicotiana attenuata demonstrate that O-acylsugar abundance correlates with resistance to both herbivores and pathogens. Removal of O-acylsugars from leaf surfaces increases M. sexta larval growth and susceptibility to fungal pathogens, whereas supplementation reduces insect performance and inhibits fungal spore germination (Luu et al. 2017). These findings illustrate how carbohydrate-derived compounds function at the interface between structural and chemical defense. Figure 3. Cell wall integrity disruption and induction of plant immune signaling by cell wall–degrading enzymes (CWDEs). CWDEs secreted by pathogens and herbivores degrade cell wall polysaccharides—cellulose into cellobiose and cellodextrins (CBs and CDs), pectin into oligogalacturonides (OGs), and hemicelluloses into xyloglucan oligosaccharides (XGOs). These degradation products function as damage-associated molecular patterns (DAMPs) and, in the case of herbivore-derived or microbe-associated molecules, as herbivore-associated (HAMPs) or pathogen-associated molecular patterns (PAMPs). Perception of these carbohydrate-derived signals by pattern-recognition receptors (PRRs) at the plasma membrane triggers immune signaling cascades, including cytosolic calcium influx and activation of the NADPH oxidase RBOHD. These early signaling events lead to a rapid oxidative burst, transcriptional activation of defense-related genes, stimulation of phytohormone biosynthesis, and enhanced immune capacity. Abbreviations: CW, cell wall; CM, cell membrane; C, cytosol. Figure created with BioRender.com and Canva.com. “Sweet immunity” and “sweet priming”: carbohydrate-mediated enhancement of plant resistance Accumulating evidence highlights sugars as pivotal regulators of plant defense responses. This concept, often referred to as “sweet immunity,” reflects the emerging view that sugars function not only as metabolic substrates but also as signaling molecules that actively modulate plant immune networks. Closely related is the concept of “sweet priming,” in which sugar-derived signals or carbohydrates of diverse origins prepare plants for faster and more robust defense responses upon subsequent attack. Sugar-mediated priming encompasses the action of soluble sugars as well as carbohydrate-derived DAMPs and HAMPs. These signals induce a physiological “pre-alert” state in plants, commonly termed priming, which enhances responsiveness without constitutive activation of defense pathways. Depending on the plant species, the nature of the attacker, and the concentration and combination of elicitors, priming can activate distinct defense programs, including systemic acquired resistance (SAR) or, in interactions with beneficial microbes, induced systemic resistance (ISR) (Walters and Heil 2007; Leibman-Markus et al. 2023; Wang et al. 2023; del Carmen Orozco-Mosqueda et al. 2023). A defining feature of sugar-mediated priming is its potential to uncouple the growth–defense trade-off. In contrast to constitutive defense activation, which often imposes substantial fitness costs, priming allows plants to maintain growth while retaining enhanced immune competence. In some cases, sugar priming even confers growth benefits, highlighting its promise for agricultural applications (Trouvelot et al. 2014; Meresa et al. 2024). This property positions carbohydrates as attractive candidates for sustainable crop protection strategies under field conditions. Experimental evidence for “sweet priming” has been documented across diverse plant species and pathosystems. Pretreatment of rice and Arabidopsis with exogenous sucrose, glucose, or fructose enhances resistance to the fungal pathogen Magnaporthe oryzae and the bacterial pathogen Pseudomonas syringae pv. tomato DC3000, respectively (Gómez-Ariza et al. 2007; Qian et al. 2015). Similarly, priming of rocket ( Eruca sativa ) with levan oligosaccharides increases resistance to Botrytis cinerea (Versluys and Van den Ende 2022), while trehalose pretreatment of wheat reduces infection by powdery mildew ( Blumeria graminis ) by up to 95% (Reignault et al. 2001). Together, these studies demonstrate that both simple sugars and oligosaccharides can effectively prime immune responses against diverse pathogens. At the mechanistic level, the primed state is characterized by accumulation of dormant defense components that enable rapid signal amplification upon challenge. These include elevated levels of PRRs, enhanced pools of MAPKs, increased abundance of key regulatory proteins such as nonexpressor of pathogenesis-related genes 1 (NPR1) and WRKY transcription factors, and chromatin modifications such as DNA and histone methylation or demethylation. Collectively, these changes reflect the involvement of SA, JA, and ET signaling pathways in sugar-mediated priming (Ahn et al. 2007; Beckers et al. 2009; Jaskiewicz et al. 2011; Kawasaki 2019). The plant cell wall represents a particularly important reservoir of priming-active carbohydrates. Oligosaccharides generated from cellulose, hemicellulose and pectin degradation, including cellodextrins and oligogalacturonides, are among the best-characterized priming agents and have been shown to elicit immune responses in a wide range of plant species (Aziz et al. 2007; Galletti et al. 2008; Ferrari et al. 2013; Rasul et al. 2012; Souza et al. 2017; Locci et al. 2019; Howlader et al. 2020; Silva-Sanzana et al. 2022; Molina et al. 2024a; Molina et al. 2024b; Degli Esposti et al. 2025). These molecules bridge structural defense and immune signaling, reinforcing the concept of the cell wall as a dynamic signaling platform rather than a static barrier. Beyond cell wall–derived oligosaccharides, a wide range of glycomolecules—including mono-, oligo-, and polysaccharides, as well as glycolipids and glycoproteins produced by bacteria, fungi, algae, and higher plants—exhibit biostimulant activity. While defense priming and biostimulation are closely related, biostimulation typically refers to improvements in agronomic traits such as growth or nutrient uptake rather than direct disease resistance (Boulogne et al. 2024). Nevertheless, certain glycomolecules operate at the interface of these processes. A prominent example is chitosan, a fungal-derived polysaccharide that induces defense gene expression in tomato via the octadecanoid pathway and enhances resistance to diverse pathogens across multiple species (Doares et al. 1995; Ghaouth 1994; Trotel-Aziz et al. 2006). From an applied perspective, the exogenous application of sugars, oligosaccharides, and polysaccharides represents an eco-friendly alternative to conventional pesticides. Several carbohydrate-based formulations are already commercially available, including products such as FytoSave® and FytoSol®, which are based on OGs or their combinations with chitosan oligomers (van Aubel et al. 2018; Guarnizo et al. 2020; Degli Esposti et al. 2025). Despite their promise, these approaches face practical challenges, including limited penetration through the plant cuticle and interactions with phyllosphere microorganisms that may modulate efficacy. Advances in formulation technologies, such as nanocarriers, surfactants, and encapsulation systems, offer potential solutions to these limitations by improving stability, delivery, and bioavailability of carbohydrate-based treatments under field conditions (Trouvelot et al. 2014; Sun et al. 2025). Importantly, when considering agricultural applications, indirect effects of exogenously applied or secreted carbohydrates on beneficial aboveground and soil microbial communities should also be taken into account, as these microbes may further influence plant performance through their own carbohydrate-derived signals. Figure 4 . Application of “sweet immunity” knowledge in crop improvement. This scheme illustrates how fundamental research can be translated into the development of resilient crops. By understanding the molecular mechanisms behind “sweet immunity” and “sweet priming”, specific genes of interest can be targeted for genome editing. Alternatively, this knowledge can be used to design effective biopesticide formulations. These strategies can be implement to develop plants with enhanced resistance to biotic stresses or to create efficient biopesticide alternatives, which can be used under field conditions to stimulate plant immune responses. The figure was created via Canva.com Conclusions and future perspectives Sugars are ubiquitous and evolutionarily ancient metabolites that connect primary metabolism with growth, development, and stress adaptation across all domains of life. In plants, carbohydrates function not only as metabolic substrates and structural components but also as central signaling molecules that integrate environmental cues with physiological and immune responses. This review highlights how dynamic regulation of carbohydrate production, transport, cleavage, sensing, and structural incorporation underpins plant defense against both pathogens and herbivores. Through coordinated control of sugar transporters, sucrose-cleaving enzymes, and sugar–energy signaling pathways, plants continuously redistribute carbon resources to balance growth and defense demands. Sugar transport systems such as SWEETs, SUTs, and STPs determine the spatial availability of carbohydrates at host–heterotroph interfaces, while Invs and SUS modulate local sugar composition to fuel defense metabolism or restrict nutrient access to attackers. At the same time, sugar sensing and energy signaling, mediated largely by the antagonistic activities of SnRK1 and TOR, integrate carbohydrate status with immune activation, allowing plants to prioritize survival under biotic stress. Beyond metabolic regulation, carbohydrates play essential structural and signaling roles in plant defense. Remodeling of the cell wall not only reinforces physical barriers but also generates carbohydrate-derived molecular patterns that activate PTI. These processes operate in both plant–pathogen and plant–herbivore interactions and highlight the cell wall as a dynamic signaling platform that links carbohydrate metabolism to immune perception. Together with specialized carbohydrate-derived defenses, such as callose deposition, trichome formation, and O-acylsugar production, these mechanisms form a multilayered “sweet fortress” that limits invasion and damage. The concepts of “sweet immunity” and “sweet priming” further emphasize the capacity of sugars to enhance resistance while minimizing fitness costs. Sugar-mediated priming enables plants to mount faster and stronger defense responses without constitutive activation of costly immune pathways and, in some cases, even promotes growth. This property is particularly attractive for agricultural applications, as it offers opportunities to uncouple the growth–defense trade-off that has traditionally constrained crop improvement strategies. Despite substantial progress, key questions remain unresolved. Future research should aim to identify the molecular nodes that integrate sugar signaling with phytohormone networks and to determine how different attacker lifestyles and feeding strategies shape carbohydrate fluxes at the cellular and tissue levels. Improved spatial and temporal resolution of sugar dynamics, enabled by advanced imaging, metabolomics, and cell-type–specific omics approaches, will be critical for disentangling defensive and exploitative roles of carbohydrate-associated responses. In parallel, genome editing and synthetic biology approaches offer powerful tools to precisely manipulate sugar transport, metabolism, and signaling pathways to enhance resistance without compromising yield. Finally, translating fundamental insights into practical applications remains a major challenge and opportunity. The development of carbohydrate-based biostimulants and biopesticides, improved formulation technologies, and targeted manipulation of sugar-related genes holds considerable promise for sustainable crop protection. By repositioning sugars from passive energy sources to active regulators of plant immunity, this review provides a conceptual framework for harnessing carbohydrate-mediated defenses to improve plant resilience in the face of increasing biotic stress and environmental change. References Ahn, I.-P., Lee, S.-W., & Suh, S.-C., 2007. Rhizobacteria-Induced Priming in Arabidopsis Is Dependent on Ethylene, Jasmonic Acid, and NPR1. The American Phytopathological Society. https://doi.org/10.1094/MPMI-20-7-0759 Akbudak, M. A., Yildiz, K., Cetin, D., Filiz, E., Yukselbaba, U., & Srivastava, V., 2025. Characterization of ZmSnRK1 genes and their response to aphid feeding, drought and cold stress. Genetic Resources and Crop Evolution, 72(1), 735–749. https://doi.org/10.1007/s10722-024-02006-2 Ali, J., Tonğa, A., Islam, T., Mir, S., Mukarram, M., Konôpková, A. S., & Chen, R., 2024. Defense strategies and associated phytohormonal regulation in Brassica plants in response to chewing and sap-sucking insects. Frontiers in Plant Science, 15, 1376917. https://doi.org/10.3389/fpls.2024.1376917 Appel, H. M., Fescemyer, H., Ehlting, J., Weston, D., Rehrig, E., Joshi, T., Xu, D., Bohlmann, J., & Schultz, J., 2014. Transcriptional responses of Arabidopsis thaliana to chewing and sucking insect herbivores. Frontiers in Plant Science, 5, 565. https://doi.org/10.3389/fpls.2014.00565 Arnold, T., Appel, H., Patel, V., Stocum, E., Kavalier, A., & Schultz, J., 2004. Carbohydrate translocation determines the phenolic content of Populus foliage: A test of the sink–source model of plant defense. New Phytologist, 164(1), 157–164. https://doi.org/10.1111/j.1469-8137.2004.01157.x Arnold, T. M., & Schultz, J. C., 2002. Induced sink strength as a prerequisite for induced tannin biosynthesis in developing leaves of Populus. Oecologia, 130(4), 585–593. https://doi.org/10.1007/s00442-001-0839-7 Aziz, A., Gauthier, A., Bézier, A., Poinssot, B., Joubert, J.-M., Pugin, A., Heyraud, A., & Baillieul, F., 2007. Elicitor and resistance-inducing activities of beta-1,4 cellodextrins in grapevine, comparison with beta-1,3 glucans and alpha-1,4 oligogalacturonides. Journal of Experimental Botany, 58(6), 1463–1472. https://doi.org/10.1093/jxb/erm008 Aziz, A., Heyraud, A., & Lambert, B., 2004. Oligogalacturonide signal transduction, induction of defense-related responses and protection of grapevine against Botrytis cinerea. Planta, 218(5), 767–774. https://doi.org/10.1007/s00425-003-1153-x Aznar, N. R., Consolo, V. F., Salerno, G. L., & Martínez-Noël, G. M. A., 2018. TOR signaling downregulation increases resistance to the cereal killer Fusarium graminearum. Plant Signaling & Behavior, 13(2), e1414120. https://doi.org/10.1080/15592324.2017.1414120 Bacete, L., & Hamann, T., 2020. The Role of Mechanoperception in Plant Cell Wall Integrity Maintenance. Plants (Basel, Switzerland), 9(5), 574. https://doi.org/10.3390/plants9050574 Barker, L., Kühn, C., Weise, A., Schulz, A., Gebhardt, C., Hirner, B., Hellmann, H., Schulze, W., Ward, J. M., & Frommer, W. B., 2000. SUT2, a Putative Sucrose Sensor in Sieve Elements. The Plant Cell, 12(7), 1153–1164. https://doi.org/10.1105/tpc.12.7.1153 Beckers, G. J. M., Jaskiewicz, M., Liu, Y., Underwood, W. R., He, S. Y., Zhang, S., & Conrath, U., 2009. Mitogen-Activated Protein Kinases 3 and 6 Are Required for Full Priming of Stress Responses in Arabidopsis thaliana. The Plant Cell, 21(3), 944–953. https://doi.org/10.1105/tpc.108.062158 Bethke, G., Thao, A., Xiong, G., Li, B., Soltis, N. E., Hatsugai, N., Hillmer, R. A., Katagiri, F., Kliebenstein, D. J., Pauly, M., & Glazebrook, J., 2016. Pectin Biosynthesis Is Critical for Cell Wall Integrity and Immunity in Arabidopsis thaliana. The Plant Cell, 28(2), 537–556. https://doi.org/10.1105/tpc.15.00404 Bezrutczyk, M., Yang, J., Eom, J.-S., Prior, M., Sosso, D., Hartwig, T., Szurek, B., Oliva, R., Vera-Cruz, C., White, F. F., Yang, B., & Frommer, W. B., 2018. Sugar flux and signaling in plant-microbe interactions. The Plant Journal: For Cell and Molecular Biology, 93(4), 675–685. https://doi.org/10.1111/tpj.13775 Bilgin, D. D., Zavala, J. A., Zhu, J., Clough, S. J., Ort, D. R., & DeLucia, E. H., 2010. Biotic stress globally downregulates photosynthesis genes. Plant Cell Environ, 33. https://doi.org/10.1111/j.1365-3040.2010.02167.x Bitterlich, M., Krügel, U., Boldt-Burisch, K., Franken, P., & Kühn, C., 2014. The sucrose transporter SlSUT2 from tomato interacts with brassinosteroid functioning and affects arbuscular mycorrhiza formation. 78(5), 877–889. https://doi.org/10.1111/tpj.12515 Bonfig, K. B., Gabler, A., Simon, U. K., Luschin-Ebengreuth, N., Hatz, M., Berger, S., Muhammad, N., Zeier, J., Sinha, A. K., & Roitsch, T., 2010. Post-Translational Derepression of Invertase Activity in Source Leaves via Down-Regulation of Invertase Inhibitor Expression Is Part of the Plant Defense Response. Molecular Plant, 3(6), 1037–1048. https://doi.org/10.1093/mp/ssq053 Boulogne, I., Mirande-Ney, C., Bernard, S., Bardor, M., Mollet, J.-C., Lerouge, P., & Driouich, A., 2024. Glycomolecules: From “sweet immunity” to “sweet biostimulation”? Physiologia Plantarum, 176(6), e14640. https://doi.org/10.1111/ppl.14640 Breia, R., Conde, A., Badim, H., Fortes, A. M., Gerós, H., & Granell, A., 2021. Plant SWEETs: From sugar transport to plant–pathogen interaction and more unexpected physiological roles. Plant Physiology, 186(2), 836–852. https://doi.org/10.1093/plphys/kiab127 Brill, E., van Thournout, M., White, R. G., Llewellyn, D., Campbell, P. M., Engelen, S., Ruan, Y.-L., Arioli, T., & Furbank, R. T., 2011. A Novel Isoform of Sucrose Synthase Is Targeted to the Cell Wall during Secondary Cell Wall Synthesis in Cotton Fiber. Plant Physiology, 157(1), 40–54. https://doi.org/10.1104/pp.111.178574 Büttner, M., Truernit, E., Baier, K., Scholz-Starke, J., Sontheim, M., Lauterbach, C., Huss, V. a. R., & Sauer, N., 2000. AtSTP3, a green leaf‐specific, low affinity monosaccharide‐H+ symporter of Arabidopsis thaliana. Plant, Cell & Environment, 23(2), 175–184. Cao, Y., Liang, Y., Tanaka, K., Nguyen, C. T., Jedrzejczak, R. P., Joachimiak, A., & Stacey, G. 2014. The kinase LYK5 is a major chitin receptor in Arabidopsis and forms a chitin-induced complex with related kinase CERK1. elife, 3, e03766. https://doi.org/10.7554/eLife.03766 Chang, Y.-A., Dai, N.-C., Chen, H.-J., Tseng, C.-H., Huang, S.-T., & Wang, S.-J., 2019. Regulation of rice sucrose transporter 4 gene expression in response to insect herbivore chewing. Journal of Plant Interactions, 14(1), 525–532. https://doi.org/10.1080/17429145.2019.1662099 Chen, J., Sun, M., Xiao, G., Shi, R., Zhao, C., Zhang, Q., Yang, S., & Xuan, Y., 2023. Starving the enemy: How plant and microbe compete for sugar on the border. Frontiers in Plant Science, 14. https://doi.org/10.3389/fpls.2023.1230254 Chen, L.-Q., Hou, B.-H., Lalonde, S., Takanaga, H., Hartung, M. L., Qu, X.-Q., Guo, W.-J., Kim, J.-G., Underwood, W., Chaudhuri, B., Chermak, D., Antony, G., White, F. F., Somerville, S. C., Mudgett, M. B., & Frommer, W. B., 2010. Sugar transporters for intercellular exchange and nutrition of pathogens. Nature, 468(7323), 527–532. https://doi.org/10.1038/nature09606 Chen, L.-Q., Qu, X.-Q., Hou, B.-H., Sosso, D., Osorio, S., Fernie, A. R., & Frommer, W. B., 2012. Sucrose Efflux Mediated by SWEET Proteins as a Key Step for Phloem Transport. Science, 335(6065), 207–211. https://doi.org/10.1126/science.1213351 Chen, W., Li, Y., Yan, R., Ren, L., Liu, F., Zeng, L., Sun, S., Yang, H., Chen, K., Xu, L., Liu, L., Fang, X., & Liu, S., 2021. SnRK1.1-mediated resistance of Arabidopsis thaliana to clubroot disease is inhibited by the novel Plasmodiophora brassicae effector PBZF1. Molecular Plant Pathology, 22(9), 1057–1069. https://doi.org/10.1111/mpp.13095 Chibbar, R. N., Jaiswal, S., Gangola, M., & Båga, M., 2016. Carbohydrate Metabolism. Reference Module in Food Science. https://doi.org/10.1016/B978-0-08-100596-5.00089-5 Cho, Y.-H., & Yoo, S.-D., 2011. Signaling Role of Fructose Mediated by FINS1/FBP in Arabidopsis thaliana. PLOS Genetics, 7(1), e1001263. https://doi.org/10.1371/journal.pgen.1001263 Chowdhury, J., Lück, S., Rajaraman, J., Douchkov, D., Shirley, N. J., Schwerdt, J. G., Schweizer, P., Fincher, G. B., Burton, R. A., & Little, A., 2017. Altered Expression of Genes Implicated in Xylan Biosynthesis Affects Penetration Resistance against Powdery Mildew. Frontiers in Plant Science, 8, 445. https://doi.org/10.3389/fpls.2017.00445 Chu, Z., Yuan, M., Yao, J., Ge, X., Yuan, B., Xu, C., Li, X., Fu, B., Li, Z., Bennetzen, J. L., Zhang, Q., & Wang, S., 2006. Promoter mutations of an essential gene for pollen development result in disease resistance in rice. Genes & Development, 20(10), 1250–1255. https://doi.org/10.1101/gad.1416306 Cox, K. L., Meng, F., Wilkins, K. E., Li, F., Wang, P., Booher, N. J., Carpenter, S. C. D., Chen, L.-Q., Zheng, H., Gao, X., Zheng, Y., Fei, Z., Yu, J. Z., Isakeit, T., Wheeler, T., Frommer, W. B., He, P., Bogdanove, A. J., & Shan, L., 2017. TAL effector driven induction of a SWEET gene confers susceptibility to bacterial blight of cotton. Nature Communications, 8(1), 15588. https://doi.org/10.1038/ncomms15588 De Vleesschauwer, D., Filipe, O., Hoffman, G., Seifi, H. S., Haeck, A., Canlas, P., Van Bockhaven, J., De Waele, E., Demeestere, K., Ronald, P., & Hofte, M., 2017. Target of rapamycin signaling orchestrates growth–defense trade‐offs in plants. New Phytologist, 217(1), 305–319. https://doi.org/10.1111/nph.14785 Degli Esposti, C., Guerrisi, L., Peruzzi, G., Giulietti, S., & Pontiggia, D., 2025. Cell wall bricks of defence: The case study of oligogalacturonides. Frontiers in Plant Science, 16, 1552926. https://doi.org/10.3389/fpls.2025.1552926 del Carmen Orozco-Mosqueda, M., Fadiji, A.E., Babalola, O.O. and Santoyo, G., 2023. Bacterial elicitors of the plant immune system: An overview and the way forward. Plant Stress, 7, 100138. https://doi.org/10.1016/j.stress.2023.100138 Delgado-Cerezo, M., Sánchez-Rodríguez, C., Escudero, V., Miedes, E., Fernández, P. V., Jordá, L., Hernández-Blanco, C., Sánchez-Vallet, A., Bednarek, P., Schulze-Lefert, P., Somerville, S., Estevez, J. M., Persson, S., & Molina, A., 2012. Arabidopsis heterotrimeric G-protein regulates cell wall defense and resistance to necrotrophic fungi. Molecular Plant, 5(1), 98–114. https://doi.org/10.1093/mp/ssr082 Denoux, C., Galletti, R., Mammarella, N., Gopalan, S., Werck, D., De Lorenzo, G., Ferrari, S., Ausubel, F. M., & Dewdney, J., 2008. Activation of defense response pathways by OGs and Flg22 elicitors in Arabidopsis seedlings. Molecular Plant, 1(3), 423–445. https://doi.org/10.1093/mp/ssn019 Divol, F., Vilaine, F., Thibivilliers, S., Amselem, J., Palauqui, J.-C., Kusiak, C., & Dinant, S., 2005. Systemic response to aphid infestation by Myzus persicae in the phloem of Apium graveolens. Plant Molecular Biology, 57(4), 517–540. https://doi.org/10.1007/s11103-005-0338-z Doares, S. H., Syrovets, T., Weiler, E. W., & Ryan, C. A., 1995. Oligogalacturonides and chitosan activate plant defensive genes through the octadecanoid pathway. Proceedings of the National Academy of Sciences of the United States of America, 92(10), 4095–4098. https://doi.org/10.1073/pnas.92.10.4095 Doidy, J., Grace, E., Kühn, C., Simon-Plas, F., Casieri, L., & Wipf, D., 2012. Sugar transporters in plants and in their interactions with fungi. Trends in Plant Science, 17(7), 413–422. https://doi.org/10.1016/j.tplants.2012.03.009 Dubey, N. K., Idris, A., Verma, A. K., Chandrashekar, K., & Pandey, K. D., 2013. Expression Pattern of Sucrose Transporters in Arabidopsis thaliana During Aphid (Myzus persicae) Infestation. American Journal of Plant Sciences, 4(12C), 47-51. https://doi.org/10.4236/ajps.2013.412A3006 Ellis, C., Karafyllidis, I., Wasternack, C., & Turner, J. G., 2002. The Arabidopsis mutant cev1 links cell wall signaling to jasmonate and ethylene responses. The Plant Cell, 14(7), 1557–1566. https://doi.org/10.1105/tpc.002022 Eom, J.-S., Chen, L.-Q., Sosso, D., Julius, B. T., Lin, I., Qu, X.-Q., Braun, D. M., & Frommer, W. B., 2015. SWEETs, transporters for intracellular and intercellular sugar translocation. Current Opinion in Plant Biology, 25, 53–62. https://doi.org/10.1016/j.pbi.2015.04.005 Essmann, J., Schmitz-Thom, I., Schön, H., Sonnewald, S., Weis, E., & Scharte, J., 2008. RNA Interference-Mediated Repression of Cell Wall Invertase Impairs Defense in Source Leaves of Tobacco. Plant Physiology, 147(3), 1288–1299. https://doi.org/10.1104/pp.108.121418 Ferrari, S., Savatin, D. V., Sicilia, F., Gramegna, G., Cervone, F., & De Lorenzo, G., 2013. Oligogalacturonides: Plant damage-associated molecular patterns and regulators of growth and development. Frontiers in Plant Science, 4. https://doi.org/10.3389/fpls.2013.00049 Ferrieri, A. P., Agtuca, B., Appel, H. M., Ferrieri, R. A., & Schultz, J. C., 2013. Temporal Changes in Allocation and Partitioning of New Carbon as 11C Elicited by Simulated Herbivory Suggest that Roots Shape Aboveground Responses in Arabidopsis. Plant Physiology, 161(2), 692–704. https://doi.org/10.1104/pp.112.208868 Ferrieri, A. P., Appel, H., Ferrieri, R. A., & Schultz, J. C., 2012. Novel application of 2-[18F]fluoro-2-deoxy-d-glucose to study plant defenses. Nuclear Medicine and Biology, 39(8), 1152–1160. https://doi.org/10.1016/j.nucmedbio.2012.06.005 Ferrieri, A. P., Arce, C. C. M., Machado, R. A. R., Meza-Canales, I. D., Lima, E., Baldwin, I. T., & Erb, M., 2015. A Nicotiana attenuata cell wall invertase inhibitor (NaCWII) reduces growth and increases secondary metabolite biosynthesis in herbivore-attacked plants. New Phytologist, 208(2), 519–530. https://doi.org/10.1111/nph.13475 Filipe, O., De Vleesschauwer, D., Haeck, A., Demeestere, K., & Höfte, M. 2018. The energy sensor OsSnRK1a confers broad-spectrum disease resistance in rice. Scientific Reports, 8(1), 3864. https://doi.org/10.1038/s41598-018-22101-6 Fotopoulos, V., Gilbert, M. J., Pittman, J. K., Marvier, A. C., Buchanan, A. J., Sauer, N., Hall, J. L., & Williams, L. E., 2003. The Monosaccharide Transporter Gene, AtSTP4, and the Cell-Wall Invertase, Atβfruct1, Are Induced in Arabidopsis during Infection with the Fungal Biotroph Erysiphe cichoracearum. Plant Physiology, 132(2), 821–829. https://doi.org/10.1104/pp.103.021428 Galletti, R., Denoux, C., Gambetta, S., Dewdney, J., Ausubel, F. M., De Lorenzo, G., & Ferrari, S., 2008. The AtrbohD-Mediated Oxidative Burst Elicited by Oligogalacturonides in Arabidopsis Is Dispensable for the Activation of Defense Responses Effective against Botrytis cinerea. Plant Physiology, 148(3), 1695–1706. https://doi.org/10.1104/pp.108.127845 Gebauer, P., Korn, M., Engelsdorf, T., Sonnewald, U., Koch, C., & Voll, L. M., 2017. Sugar Accumulation in Leaves of Arabidopsis sweet11/sweet12 Double Mutants Enhances Priming of the Salicylic Acid-Mediated Defense Response. Frontiers in Plant Science, 8. https://doi.org/10.3389/fpls.2017.01378 Geiger, D., 2020. Plant glucose transporter structure and function. Pflügers Archiv - European Journal of Physiology, 472(9), 1111–1128. https://doi.org/10.1007/s00424-020-02449-3 German, L., Yeshvekar, R. and Benitez‐Alfonso, Y., 2023. Callose metabolism and the regulation of cell walls and plasmodesmata during plant mutualistic and pathogenic interactions. Plant, Cell & Environment, 46(2), 391-404. https://doi.org/10.1111/pce.14510 Ghaouth, A. E., 1994. Effect of Chitosan on Cucumber Plants: Suppression of Pythium aphanidermatum and Induction of Defense Reactions. Phytopathology, 84(3), 313. https://doi.org/10.1094/Phyto-84-313 Glas, J. J., Alba, J. M., Simoni, S., Villarroel, C. A., Stoops, M., Schimmel, B. C., Schuurink, R. C., Sabelis, M. W., & Kant, M. R., 2014. Defense suppression benefits herbivores that have a monopoly on their feeding site but can backfire within natural communities. BMC Biology, 12(1), 98. https://doi.org/10.1186/s12915-014-0098-9 Glazebrook, J., 2005. Contrasting mechanisms of defense against biotrophic and necrotrophic pathogens. Annual Review of Phytopathology, 43, 205–227. https://doi.org/10.1146/annurev.phyto.43.040204.135923 Gómez-Ariza, J., Campo, S., Rufat, M., Estopà, M., Messeguer, J., Segundo, B.S. and Coca, M., 2007. Sucrose-mediated priming of plant defense responses and broad-spectrum disease resistance by overexpression of the maize pathogenesis-related PRms protein in rice plants. Molecular plant-microbe interactions, 20(7), 832-842. https://doi.org/10.1094/MPMI-20-7-0832 Guarnizo, N., Oliveros, D., Murillo-Arango, W., & Bermúdez-Cardona, M. B., 2020. Oligosaccharides: Defense Inducers, Their Recognition in Plants, Commercial Uses and Perspectives. Molecules, 25(24), 5972. https://doi.org/10.3390/molecules25245972 Gupta, P. K., Balyan, H. S., & Gautam, T., 2021. SWEET genes and TAL effectors for disease resistance in plants: present status and future prospects. Molecular Plant Pathology, 22(8), 1014-1026. https://doi.org/10.1111/mpp.13075 Han, X., Zhang, L., Zhao, L., Xue, P., Qi, T., Zhang, C., Yuan, H., Zhou, L., Wang, D., Qiu, J., Shen, QH., & Shen, Q. H. 2020. SnRK1 phosphorylates and destabilizes WRKY3 to enhance barley immunity to powdery mildew. Plant Communications, 1(4). https://doi.org/10.1016/j.xplc.2020.100083 Hayes, M. A., Feechan, A., & Dry, I. B., 2010. Involvement of Abscisic Acid in the Coordinated Regulation of a Stress-Inducible Hexose Transporter (VvHT5) and a Cell Wall Invertase in Grapevine in Response to Biotrophic Fungal Infection. Plant Physiology, 153(1), 211–221. https://doi.org/10.1104/pp.110.154765 Helber, N., Wippel, K., Sauer, N., Schaarschmidt, S., Hause, B., & Requena, N., 2011. A Versatile Monosaccharide Transporter That Operates in the Arbuscular Mycorrhizal Fungus Glomus sp Is Crucial for the Symbiotic Relationship with Plants. The Plant Cell, 23(10), 3812–3823. https://doi.org/10.1105/tpc.111.089813 Herbers, K., Meuwly, P., Frommer, W. B., Metraux, J. P., & Sonnewald, U., 1996. Systemic Acquired Resistance Mediated by the Ectopic Expression of Invertase: Possible Hexose Sensing in the Secretory Pathway. The Plant Cell, 8(5), 793–803. https://doi.org/10.1105/tpc.8.5.793 Horsfall, J. G., & Dimond, A. E., 1957. Interactions of Tissue Sugar, Growth Substances, and Disease Susceptibility. Zeitschrift Für Pflanzenkrankheiten (Pflanzenpathologie) Und Pflanzenschutz, 415–421. Hothorn, M., Van den Ende, W., Lammens, W., Rybin, V., & Scheffzek, K., 2010. Structural insights into the pH-controlled targeting of plant cell-wall invertase by a specific inhibitor protein. Proceedings of the National Academy of Sciences, 107(40), 17427–17432. https://doi.org/10.1073/pnas.1004481107 Howlader, P., Bose, S. K., Jia, X., Zhang, C., Wang, W., & Yin, H., 2020. Oligogalacturonides induce resistance in Arabidopsis thaliana by triggering salicylic acid and jasmonic acid pathways against Pst DC3000. International Journal of Biological Macromolecules, 164, 4054–4064. https://doi.org/10.1016/j.ijbiomac.2020.09.026 Huang, J., Taylor, J. P., Chen, J.-G., Uhrig, J. F., Schnell, D. J., Nakagawa, T., Korth, K. L., & Jones, A. M., 2006. The Plastid Protein THYLAKOID FORMATION1 and the Plasma Membrane G-Protein GPA1 Interact in a Novel Sugar-Signaling Mechanism in Arabidopsis. The Plant Cell, 18(5), 1226–1238. https://doi.org/10.1105/tpc.105.037259 Hulsmans, S., Rodriguez, M., De Coninck, B. and Rolland, F., 2016. The SnRK1 energy sensor in plant biotic interactions. Trends in Plant Science, 21(8), 648-661. https://doi.org/10.1016/j.tplants.2016.04.008 Huot, B., Yao, J., Montgomery, B. L., & He, S. Y., 2014. Growth–Defense Tradeoffs in Plants: A Balancing Act to Optimize Fitness. Molecular Plant, 7(8), 1267–1287. https://doi.org/10.1093/mp/ssu049 Jaskiewicz, M., Conrath, U., & Peterhänsel, C., 2011. Chromatin modification acts as a memory for systemic acquired resistance in the plant stress response. EMBO Reports, 12(1), 50–55. https://doi.org/10.1038/embor.2010.186 Jeandet, P., Formela-Luboińska, M., Labudda, M., & Morkunas, I., 2022. The Role of Sugars in Plant Responses to Stress and Their Regulatory Function during Development. International Journal of Molecular Sciences, 23(9), 5161. https://doi.org/10.3390/ijms23095161 Jing, W., Uddin, S., Chakraborty, R., Van Anh, D. T., Macoy, D. M., Park, S. O., Ryu, G. R., Kim, Y. H., Cha, J., Kim, W.-Y., & Kim, M. G., 2020. Molecular characterization of HEXOKINASE1 in plant innate immunity. Applied Biological Chemistry, 63(1), 76. https://doi.org/10.1186/s13765-020-00560-8 Jones, J. D. G., & Dangl, J. L., 2006. The plant immune system. Nature, 444(7117), 323–329. https://doi.org/10.1038/nature05286 Kant, M. R., Ament, K., Sabelis, M. W., Haring, M. A., & Schuurink, R. C., 2004. Differential Timing of Spider Mite-Induced Direct and Indirect Defenses in Tomato Plants. Plant Physiology, 135(1), 483–495. https://doi.org/10.1104/pp.103.038315 Kanwar, P., & Jha, G., 2019. Alterations in plant sugar metabolism: Signatory of pathogen attack. Planta, 249(2), 305–318. https://doi.org/10.1007/s00425-018-3018-3 Karve, A., Rauh, B. L., Xia, X., Kandasamy, M., Meagher, R. B., Sheen, J., & Moore, B. D., 2008. Expression and evolutionary features of the hexokinase gene family in Arabidopsis. Planta, 228(3), 411–425. https://doi.org/10.1007/s00425-008-0746-9 Kawasaki, T., 2019. PRR Cross-Talk Jump Starts Plant Immunity. Cell Host & Microbe, 26(6), 707–709. https://doi.org/10.1016/j.chom.2019.11.007 Ke, L., Wang, Y., Schäfer, M., Städler, T., Zeng, R., Fabian, J., Pulido, H., De Moraes, C. M., Song, Y., & Xu, S., 2021. Transcriptomic Profiling Reveals Shared Signalling Networks Between Flower Development and Herbivory-Induced Responses in Tomato. Frontiers in Plant Science, 12, 722810. https://doi.org/10.3389/fpls.2021.722810 Khanna, K., Ohri, P., & Bhardwaj, R., 2023. Decoding sugar regulation and homeostasis in plants: Cracking functional roles under stresses. Journal of Plant Growth Regulation, 42(8), 4797-4817. https://doi.org/10.1007/s00344-022-10727-w Kiełkiewicz, M., Barczak-Brzyżek, A., Karpińska, B., & Filipecki, M., 2019. Unravelling the Complexity of Plant Defense Induced by a Simultaneous and Sequential Mite and Aphid Infestation. International Journal of Molecular Sciences, 20(4), 806. https://doi.org/10.3390/ijms20040806 Koch, K., 2004. Sucrose metabolism: Regulatory mechanisms and pivotal roles in sugar sensing and plant development. Current Opinion in Plant Biology, 7(3), 235–246. https://doi.org/10.1016/j.pbi.2004.03.014 Koch, K. E., 1996. CARBOHYDRATE-MODULATED GENE EXPRESSION IN PLANTS. Annual Review of Plant Biology, 47(Volume 47, 1996), 509–540. https://doi.org/10.1146/annurev.arplant.47.1.509 Körner, E., von Dahl, C. C., Bonaventure, G., & Baldwin, I. T., 2009. Pectin methylesterase NaPME1 contributes to the emission of methanol during insect herbivory and to the elicitation of defence responses in Nicotiana attenuata. Journal of Experimental Botany, 60(9), 2631–2640. https://doi.org/10.1093/jxb/erp106 Kuśnierczyk, A., Winge, P., Jørstad, T. S., Troczyńska, J., Rossiter, J. T., & Bones, A. M., 2008. Towards global understanding of plant defence against aphids—Timing and dynamics of early Arabidopsis defence responses to cabbage aphid (Brevicoryne brassicae) attack. Plant, Cell & Environment, 31(8), 1097–1115. https://doi.org/10.1111/j.1365-3040.2008.01823.x Lee, K. H., Piao, H. L., Kim, H. Y., Choi, S. M., Jiang, F., Hartung, W., Hwang, I., Kwak, J.M., Lee, IJ., Hwang, I. & Hwang, I. (2006). Activation of glucosidase via stress-induced polymerization rapidly increases active pools of abscisic acid. Cell , 126 (6), 1109-1120. Lei, M., Wang, X., Chen, K., Wei, Q., Zhou, M., Chen, G., Su, S., Tai, Y., Zhuang, K., Li, D., Liu, M., Zhang, S., & Wang, Y., 2025. Sugar transporters: Mediators of carbon flow between plants and microbes. Frontiers in Plant Science, 16. https://doi.org/10.3389/fpls.2025.1536969 Leibman-Markus, M., Schneider, A., Gupta, R., Marash, I., Rav-David, D., Carmeli-Weissberg, M., Elad, Y. and Bar, M., 2023. Immunity priming uncouples the growth–defense trade-off in tomato. Development, 150(21), p.dev201158. https://doi.org/10.1242/dev.201158 Lemoine, R., 2000. Sucrose transporters in plants: Update on function and structure. Biochimica et Biophysica Acta (BBA) - Biomembranes, 1465(1–2), 246–262. https://doi.org/10.1016/S0005-2736(00)00142-5 Lemonnier, P., Gaillard, C., Veillet, F., Verbeke, J., Lemoine, R., Coutos-Thévenot, P., & La Camera, S., 2014. Expression of Arabidopsis sugar transport protein STP13 differentially affects glucose transport activity and basal resistance to Botrytis cinerea. Plant Molecular Biology, 85(4–5), 473–484. https://doi.org/10.1007/s11103-014-0198-5 Li, L., & Sheen, J., 2016. Dynamic and diverse sugar signaling. Current Opinion in Plant Biology, 33, 116–125. https://doi.org/10.1016/j.pbi.2016.06.018 Li, L., Liu, K.H. and Sheen, J., 2021. Dynamic nutrient signaling networks in plants. Annual Review of Cell and Developmental Biology, 37(1), 341-367. https://doi.org/10.1146/annurev-cellbio-010521-015047 Li, Y., Liu, H., Yao, X., Sun, L., & Sui, X., 2022. The Role of Sugar Transporter CsSWEET7a in Apoplasmic Phloem Unloading in Receptacle and Nectary During Cucumber Anthesis. Frontiers in Plant Science, 12. https://doi.org/10.3389/fpls.2021.758526 Lionetti, V., Cervone, F., & Bellincampi, D., 2012. Methyl esterification of pectin plays a role during plant–pathogen interactions and affects plant resistance to diseases. Journal of Plant Physiology, 169(16), 1623–1630. https://doi.org/10.1016/j.jplph.2012.05.006 Liu, J., Han, L., Huai, B., Zheng, P., Chang, Q., Guan, T., Li, D., Huang, L., & Kang, Z., 2015. Down-regulation of a wheat alkaline/neutral invertase correlates with reduced host susceptibility to wheat stripe rust caused by Puccinia striiformis. Journal of Experimental Botany, 66(22), 7325–7338. https://doi.org/10.1093/jxb/erv428 Liu, Y.-H., Song, Y.-H., & Ruan, Y.-L., 2022. Sugar conundrum in plant-pathogen interactions: Roles of invertase and sugar transporters depend on pathosystems. Journal of Experimental Botany, 73(7), 1910–1925. https://doi.org/10.1093/jxb/erab562 Liu, J., Nie, B., Yu, B., Xu, F., Zhang, Q., Wang, Y., & Xu, W. 2023. Rice ubiquitin‐conjugating enzyme OsUbc13 negatively regulates immunity against pathogens by enhancing the activity of OsSnRK1a. Plant Biotechnology Journal, 21(8), 1590-1610. https://doi.org/10.1111/pbi.14059 Locci, F., Benedetti, M., Pontiggia, D., Citterico, M., Caprari, C., Mattei, B., Cervone, F., & De Lorenzo, G., 2019. An Arabidopsis berberine bridge enzyme-like protein specifically oxidizes cellulose oligomers and plays a role in immunity. The Plant Journal: For Cell and Molecular Biology, 98(3), 540–554. https://doi.org/10.1111/tpj.14237 Luu, V. T., Weinhold, A., Ullah, C., Dressel, S., Schoettner, M., Gase, K., Gaquerel, E., Xu, S., & Baldwin, I. T., 2017. O-Acyl Sugars Protect a Wild Tobacco from Both Native Fungal Pathogens and a Specialist Herbivore. Plant Physiology, 174(1), 370–386. https://doi.org/10.1104/pp.16.01904 Manabe, Y., Verhertbruggen, Y., Gille, S., Harholt, J., Chong, S.-L., Pawar, P. M.-A., Mellerowicz, E. J., Tenkanen, M., Cheng, K., Pauly, M., & Scheller, H. V., 2013. Reduced Wall Acetylation proteins play vital and distinct roles in cell wall O-acetylation in Arabidopsis. Plant Physiology, 163(3), 1107–1117. https://doi.org/10.1104/pp.113.225193 Marash, I., Leibman‐Markus, M., Gupta, R., Avni, A., & Bar, M., 2022. TOR inhibition primes immunity and pathogen resistance in tomato in a salicylic acid‐dependent manner. Molecular Plant Pathology, 23(7), 1035–1047. https://doi.org/10.1111/mpp.13207 Mary Wanjiru, W., Zhensheng, K., & Buchenauer, H., 2002. Importance of Cell Wall Degrading Enzymes Produced by Fusarium graminearum during Infection of Wheat Heads. European Journal of Plant Pathology, 108(8), 803–810. https://doi.org/10.1023/A:1020847216155 Mbiza, N. I. T., Hu, Z., Zhang, H., Zhang, Y., Luo, X., Wang, Y., Wang, Y., Liu, T., Li, J., Wang, X., Zhang, J., & Yu, Y., 2022. GhCalS5 is involved in cotton response to aphid attack through mediating callose formation. Frontiers in Plant Science, 13. https://doi.org/10.3389/fpls.2022.892630 Meresa, B. K., Ayimut, K. M., Weldemichael, M. Y., Geberemedhin, K. H., Kassegn, H. H., Geberemikael, B. A., & Egigu, E. M., 2024. Carbohydrate elicitor-induced plant immunity: Advances and prospects. Heliyon, 10(15). https://doi.org/10.1016/j.heliyon.2024.e34871 Meteignier, L.V., El Oirdi, M., Cohen, M., Barff, T., Matteau, D., Lucier, J.F., Rodrigue, S., Jacques, P.E., Yoshioka, K. and Moffett, P., 2017. Translatome analysis of an NB-LRR immune response identifies important contributors to plant immunity in Arabidopsis. Journal of Experimental Botany, 68(9), pp.2333-2344. https://doi.org/10.1093/jxb/erx078 Meyer, S., Lauterbach, C., Niedermeier, M., Barth, I., Sjolund, R. D., & Sauer, N., 2004. Wounding enhances expression of AtSUC3, a sucrose transporter from Arabidopsis sieve elements and sink tissues. Plant Physiology, 134(2), 684–693. https://doi.org/10.1104/pp.103.033399 Miya, A., Albert, P., Shinya, T., Desaki, Y., Ichimura, K., Shirasu, K., Narusaka, Y., Kawakami, N., Kaku, H., & Shibuya, N. 2007. CERK1, a LysM receptor kinase, is essential for chitin elicitor signaling in Arabidopsis. Proceedings of the National Academy of Sciences, 104(49), 19613-19618. https://doi.org/10.1073/pnas.0705147104 Mohase, L., & Taiwe, B., 2015. Saliva fractions from South African Russian wheat aphid biotypes induce differential defence responses in wheat. South African Journal of Plant and Soil, 32(4), Article 4. Molina, A., Jordá, L., Torres, M. Á., Martín-Dacal, M., Berlanga, D. J., Fernández-Calvo, P., Gómez-Rubio, E., & Martín-Santamaría, S., 2024a. Plant cell wall-mediated disease resistance: Current understanding and future perspectives. Molecular Plant, 17(5), 699–724. https://doi.org/10.1016/j.molp.2024.04.003 Molina, A., Sánchez-Vallet, A., Jordá, L., Carrasco-López, C., Rodríguez-Herva, J. J., & López-Solanilla, E., 2024b. Plant cell walls: Source of carbohydrate-based signals in plant-pathogen interactions. Current Opinion in Plant Biology, 82, 102630. https://doi.org/10.1016/j.pbi.2024.102630 Moore, B., Zhou, L., Rolland, F., Hall, Q., Cheng, W.-H., Liu, Y.-X., Hwang, I., Jones, T., & Sheen, J., 2003. Role of the Arabidopsis Glucose Sensor HXK1 in Nutrient, Light, and Hormonal Signaling. Science, 300(5617). https://doi.org/10.1126/science.1080585 Moran, P. J., & Thompson, G. A., 2001. Molecular Responses to Aphid Feeding in Arabidopsis in Relation to Plant Defense Pathways. Plant Physiology, 125(2), 1074–1085. https://doi.org/10.1104/pp.125.2.1074 Morkunas, I., & Ratajczak, L. 2014. The role of sugar signaling in plant defense responses against fungal pathogens. Acta Physiologiae Plantarum, 36(7), 1607-1619. https://doi.org/10.1007/s11738-014-1559-z Naseem, M., Kunz, M., & Dandekar, T., 2017. Plant-Pathogen Maneuvering over Apoplastic Sugars. Trends in Plant Science, 22(9), 740–743. https://doi.org/10.1016/j.tplants.2017.07.001 Ostrowski, M., & Jakubowska, A. (2014). UDP-glycosyltransferases of plant hormones. Adv. Cell Biol , 4 (1), 43-60. Park, J., Kim, S., Choi, E., Auh, C.-K., Park, J.-B., Kim, D.-G., Chung, Y.-J., Lee, T.-K., & Lee, S., 2013. Altered invertase activities of symptomatic tissues on Beet severe curly top virus (BSCTV) infected Arabidopsis thaliana. Journal of Plant Research, 126(5), 743–752. https://doi.org/10.1007/s10265-013-0562-6 Perez-Alonso, M. M., Talavera-Mateo, L., Ojeda-Martinez, D., Barcenilla-Valcárcel, L., Montesinos, Á., Garcia, A., Frey, C., Boter, M., Martinez, M., Diaz, I., & Santamaria, M. E., 2025. The Role of Pant Cell Walls in Pest Resistance: Current Insights and Future Perspectives. Journal of Experimental Botany, eraf306. https://doi.org/10.1093/jxb/eraf306 Pommerrenig, B., Müdsam, C., Kischka, D., & Neuhaus, H. E., 2020. Treat and trick: Common regulation and manipulation of sugar transporters during sink establishment by the plant and the pathogen. Journal of Experimental Botany, 71(14), 3930–3940. https://doi.org/10.1093/jxb/eraa168 Ponzio, Camille, Papazian, Stefano, Albrectsen, Benedicte R., Dicke, Marcel, & Gols, Rieta (2017). Dual herbivore attack and herbivore density affect metabolic profiles of Brassica nigra leaves. Plant, Cell & Environment, 40(8), 1356-1367. Proels, R. K., & Hückelhoven, R., 2014. Cell‐wall invertases, key enzymes in the modulation of plant metabolism during defence responses. Molecular Plant Pathology, 15(8), 858–864. https://doi.org/10.1111/mpp.12139 Qian, Y., Tan, D.X., Reiter, R.J. and Shi, H., 2015. Comparative metabolomic analysis highlights the involvement of sugars and glycerol in melatonin-mediated innate immunity against bacterial pathogen in Arabidopsis. Scientific Reports, 5(1), 15815. https://doi.org/10.1038/srep15815 Rasul, S., Dubreuil-Maurizi, C., Lamotte, O., Koen, E., Poinssot, B., Alcaraz, G., Wendehenne, D., & Jeandroz, S., 2012. Nitric oxide production mediates oligogalacturonide-triggered immunity and resistance to Botrytis cinerea in Arabidopsis thaliana. Plant, Cell & Environment, 35(8), 1483–1499. https://doi.org/10.1111/j.1365-3040.2012.02505.x Reignault, P., Cogan, A., Muchembled, J., Sahraoui, A. L.-H., Durand, R., & Sancholle, M., 2001. Trehalose induces resistance to powdery mildew in wheat. 149(3), 519–529. https://doi.org/10.1046/j.1469-8137.2001.00035.x Rennie, E. A., & Turgeon, R., 2009. A comprehensive picture of phloem loading strategies. Proceedings of the National Academy of Sciences, 106(33), 14162–14167. https://doi.org/10.1073/pnas.0902279106 Riesmeier, J. W., Willmitzer, L., & Frommer, W. B., 1994. Evidence for an essential role of the sucrose transporter in phloem loading and assimilate partitioning. The EMBO Journal, 13(1), 1–7. Roitsch, T., Bittner, M., & Godt, D. E., 1995. Induction of Apoplastic Invertase of Chenopodium rubrum by D-Glucose and a Glucose Analog and Tissue-Specific Expression Suggest a Role in Sink-Source Regulation. Plant Physiology, 108(1), 285–294. https://doi.org/10.1104/pp.108.1.285 Rolland, F., Baena-Gonzalez, E., & Sheen, J. 2006. Sugar sensing and signaling in plants: conserved and novel mechanisms. Annu. Rev. Plant Biol., 57(1), 675-709. https://doi.org/10.1146/annurev.arplant.57.032905.105441 Römer, P., Recht, S., Strauß, T., Elsaesser, J., Schornack, S., Boch, J., Wang, S., & Lahaye, T., 2010. Promoter elements of rice susceptibility genes are bound and activated by specific TAL effectors from the bacterial blight pathogen, Xanthomonas oryzae pv. Oryzae. New Phytologist, 187(4), 1048–1057. https://doi.org/10.1111/j.1469-8137.2010.03217.x Ruan, Y.-L., 2012. Signaling Role of Sucrose Metabolism in Development. Molecular Plant, 5(4), 763–765. https://doi.org/10.1093/mp/sss046 Ruan, Y.-L., 2014. Sucrose Metabolism: Gateway to Diverse Carbon Use and Sugar Signaling. Annual Review of Plant Biology, 65(Volume 65, 2014), 33–67. https://doi.org/10.1146/annurev-arplant-050213-040251 Sampedro, J., Pardo, B., Gianzo, C., Guitián, E., Revilla, G., & Zarra, I., 2010. Lack of α-Xylosidase Activity in Arabidopsis Alters Xyloglucan Composition and Results in Growth Defects. Plant Physiology, 154(3), 1105–1115. https://doi.org/10.1104/pp.110.163212 Sauer, N., Friedländer, K., & Gräml‐Wicke, U., 1990. Primary structure, genomic organization and heterologous expression of a glucose transporter from Arabidopsis thaliana. The EMBO Journal, 9(10), 3045–3050. Schulz, A., Beyhl, D., Marten, I., Wormit, A., Neuhaus, E., Poschet, G., Büttner, M., Schneider, S., Sauer, N., & Hedrich, R., 2011. Proton‐driven sucrose symport and antiport are provided by the vacuolar transporters SUC4 and TMT1/2. The Plant Journal, 68(1), 129–136. https://doi.org/10.1111/j.1365-313X.2011.04672.x Schwachtje, J., Minchin, P. E., Jahnke, S., van Dongen, J. T., Schittko, U., & Baldwin, I. T. 2006. SNF1-related kinases allow plants to tolerate herbivory by allocating carbon to roots. Proceedings of the National Academy of Sciences, 103(34), 12935-12940. https://doi.org/10.1073/pnas.0602316103 Siddappaji, M. H., Scholes, D. R., Krishnankutty, S. M., Calla, B., Clough, S. J., Zielinski, R. E., & Paige, K. N., 2015. The role of invertases in plant compensatory responses to simulated herbivory. BMC Plant Biology, 15(1), 278. https://doi.org/10.1186/s12870-015-0655-6 Silva-Sanzana, C., Zavala, D., Moraga, F., Herrera-Vásquez, A., & Blanco-Herrera, F., 2022. Oligogalacturonides Enhance Resistance against Aphids through Pattern-Triggered Immunity and Activation of Salicylic Acid Signaling. International Journal of Molecular Sciences, 23(17), Article 17. https://doi.org/10.3390/ijms23179753 Sonnewald, S., Priller, J. P. R., Schuster, J., Glickmann, E., Hajirezaei, M.-R., Siebig, S., Mudgett, M. B., & Sonnewald, U., 2012. Regulation of Cell Wall-Bound Invertase in Pepper Leaves by Xanthomonas campestris pv. Vesicatoria Type Three Effectors. PLOS ONE, 7(12), e51763. https://doi.org/10.1371/journal.pone.0051763 Souza, C. de A., Li, S., Lin, A. Z., Boutrot, F., Grossmann, G., Zipfel, C., & Somerville, S. C., 2017. Cellulose-Derived Oligomers Act as Damage-Associated Molecular Patterns and Trigger Defense-Like Responses. Plant Physiology, 173(4), 2383–2398. https://doi.org/10.1104/pp.16.01680 Storr, T., & Hall, J. L., 1992. The effect of infection by Erysiphe pisi DC on acid and alkaline invertase activities and aspects of starch biochemistry in leaves of Pisum sativum L. New Phytologist, 121(4), 535–543. https://doi.org/10.1111/j.1469-8137.1992.tb01123.x Sun, A., Yu, B., Zhang, Q., Peng, Y., Yang, J., Sun, Y., Qin, P., Jia, T., Smeekens, S., & Teng, S., 2020. MYC2-Activated TRICHOME BIREFRINGENCE-LIKE37 Acetylates Cell Walls and Enhances Herbivore Resistance. Plant Physiology, 184(2), 1083–1096. https://doi.org/10.1104/pp.20.00683 Sun, G., Xiao, Y., Yin, H., Yu, K., Wang, Y., & Wang, Y., 2025. Oligosaccharide elicitors in plant immunity: Molecular mechanisms and disease resistance strategies. Plant Communications. https://doi.org/10.1016/j.xplc.2025.101469 External Link Sun, L., Yang, D., Kong, Y., Chen, Y., Li, X.-Z., Zeng, L.-J., Li, Q., Wang, E.-T., & He, Z.-H., 2014. Sugar homeostasis mediated by cell wall invertase GRAIN INCOMPLETE FILLING 1 (GIF1) plays a role in pre-existing and induced defence in rice. Molecular Plant Pathology, 15(2), 161–173. https://doi.org/10.1111/mpp.12078 Sun, L., & Zhang, J. 2020. Regulatory role of receptor-like cytoplasmic kinases in early immune signaling events in plants. FEMS Microbiology Reviews, 44(6), 845-856. https://doi.org/10.1093/femsre/fuaa035 Sutton, P. N., Gilbert, M. J., Williams, L. E., & Hall, J. L., 2007. Powdery mildew infection of wheat leaves changes host solute transport and invertase activity. Physiologia Plantarum, 129(4), 787–795. https://doi.org/10.1111/j.1399-3054.2007.00863.x Szczesny, R., Büttner, D., Escolar, L., Schulze, S., Seiferth, A., & Bonas, U. 2010. Suppression of the AvrBs1‐specific hypersensitive response by the YopJ effector homolog AvrBsT from Xanthomonas depends on a SNF1‐related kinase. New Phytologist, 187(4), 1058-1074. https://doi.org/10.1111/j.1469-8137.2010.03346.x Tauzin, A. S., & Giardina, T., 2014. Sucrose and invertases, a part of the plant defense response to the biotic stresses. Frontiers in Plant Science, 5, 293. https://doi.org/10.3389/fpls.2014.00293 Thaler, J. S., Owen, B., & Higgins, V. J., 2004. The role of the jasmonate response in plant susceptibility to diverse pathogens with a range of lifestyles. Plant Physiology, 135(1), 530–538. https://doi.org/10.1104/pp.104.041566 Trotel-Aziz, P., Couderchet, M., Vernet, G., & Aziz, A., 2006. Chitosan Stimulates Defense Reactions in Grapevine Leaves and Inhibits Development of Botrytis Cinerea. European Journal of Plant Pathology, 114(4), 405–413. https://doi.org/10.1007/s10658-006-0005-5 Trouvelot, S., Héloir, M.-C., Poinssot, B., Gauthier, A., Paris, F., Guillier, C., Combier, M., Trdá, L., Daire, X., & Adrian, M., 2014. Carbohydrates in plant immunity and plant protection: Roles and potential application as foliar sprays. Frontiers in Plant Science, 5. https://doi.org/10.3389/fpls.2014.00592 van Aubel, G., Serderidis, S., Ivens, J., Clinckemaillie, A., Legrève, A., Hause, B., & Van Cutsem, P., 2018. Oligosaccharides successfully thwart hijacking of the salicylic acid pathway by Phytophthora infestans in potato leaves. Plant Pathology, 67(9), 1901–1911. https://doi.org/10.1111/ppa.12908 Veronese, P., Nakagami, H., Bluhm, B., AbuQamar, S., Chen, X., Salmeron, J., Dietrich, R. A., Hirt, H., & Mengiste, T. 2006. The membrane-anchored BOTRYTIS-INDUCED KINASE1 plays distinct roles in Arabidopsis resistance to necrotrophic and biotrophic pathogens. The Plant Cell, 18(1), 257-273. https://doi.org/10.1105/tpc.105.035576 Versluys, M., & Van den Ende, W., 2022. Sweet Immunity Aspects during Levan Oligosaccharide-Mediated Priming in Rocket against Botrytis cinerea. Biomolecules, 12(3), 370. https://doi.org/10.3390/biom12030370 Vogel, J. P., Raab, T. K., Schiff, C., & Somerville, S. C., 2002. PMR6, a pectate lyase-like gene required for powdery mildew susceptibility in Arabidopsis. The Plant Cell, 14(9), 2095–2106. https://doi.org/10.1105/tpc.003509 Vogel, J. P., Raab, T. K., Somerville, C. R., & Somerville, S. C., 2004. Mutations in PMR5 result in powdery mildew resistance and altered cell wall composition. The Plant Journal: For Cell and Molecular Biology, 40(6), 968–978. https://doi.org/10.1111/j.1365-313X.2004.02264.x Vos, I. A., Verhage, A., Schuurink, R. C., Watt, L. G., Pieterse, C. M. J., & Van Wees, S. C. M., 2013. Onset of herbivore-induced resistance in systemic tissue primed for jasmonate-dependent defenses is activated by abscisic acid. Frontiers in Plant Science, 4. https://doi.org/10.3389/fpls.2013.00539 Walerowski, P., Gündel, A., Yahaya, N., Truman, W., Sobczak, M., Olszak, M., Rolfe, S., Borisjuk, L., & Malinowski, R., 2018. Clubroot Disease Stimulates Early Steps of Phloem Differentiation and Recruits SWEET Sucrose Transporters within Developing Galls. The Plant Cell, 30(12), 3058–3073. https://doi.org/10.1105/tpc.18.00283 Walters, D. and Heil, M., 2007. Costs and trade-offs associated with induced resistance. Physiological and Molecular Plant Pathology, 71(1-3), 3-17. https://doi.org/10.1016/j.pmpp.2007.09.008 Wan, J., He, M., Hou, Q., Zou, L., Yang, Y., Wei, Y., & Chen, X., 2021. Cell wall associated immunity in plants. Stress Biology, 1(1), 3. https://doi.org/10.1007/s44154-021-00003-4 Wang, D., Wei, L., Liu, T., Ma, J., Huang, K., Guo, H., Huang, Y., Zhang, L., Zhao, J., Tsuda, K. and Wang, Y., 2023. Suppression of ETI by PTI priming to balance plant growth and defense through an MPK3/MPK6-WRKYs-PP2Cs module. Molecular plant, 16(5), 903-918. https://doi.org/10.1016/j.molp.2023.04.004 Wang, X., Shen, C., Meng, P., Tan, G., & Lv, L., 2021. Analysis and review of trichomes in plants. BMC Plant Biology, 21(1), 70. https://doi.org/10.1186/s12870-021-02840-x Wei, Y.-T., Bao, Q.-X., Shi, Y.-N., Mu, X.-R., Wang, Y.-B., Jiang, J.-H., Yu, F.-H., & Meng, L.-S., 2025. Trichome development of systemic developing leaves is regulated by a nutrient sensor-relay mechanism within mature leaves. Science Advances, 11(6), eadq5820. https://doi.org/10.1126/sciadv.adq5820 Wind, J., Smeekens, S., & Hanson, J., 2010. Sucrose: Metabolite and signaling molecule. Phytochemistry, 71(14–15), 1610–1614. https://doi.org/10.1016/j.phytochem.2010.07.007 Wormit, A., Trentmann, O., Feifer, I., Lohr, C., Tjaden, J., Meyer, S., Schmidt, U., Martinoia, E., & Neuhaus, H. E., 2006. Molecular Identification and Physiological Characterization of a Novel Monosaccharide Transporter from Arabidopsis Involved in Vacuolar Sugar Transport. The Plant Cell, 18(12), 3476–3490. https://doi.org/10.1105/tpc.106.047290 Xiao, L., Gheysen, G., Yang, M., Xiao, X., Xu, L., Guo, X., Yang, L., Liu, W., He, Y., Peng, D., Peng, H., Ma, K., Long, H., Wang, G., & Xiao, Y., 2024. Brown planthopper infestation on rice reduces plant susceptibility to Meloidogyne graminicola by reducing root sugar allocation. New Phytologist, 242(1), 262–277. https://doi.org/10.1111/nph.19570 Xiong, Y., McCormack, M., Li, L., Hall, Q., Xiang, C., & Sheen, J., 2013. Glucose–TOR signalling reprograms the transcriptome and activates meristems. Nature, 496(7444), 181–186. https://doi.org/10.1038/nature12030 Xu, S.-M., Brill, E., Llewellyn, D. J., Furbank, R. T., & Ruan, Y.-L., 2012. Overexpression of a Potato Sucrose Synthase Gene in Cotton Accelerates Leaf Expansion, Reduces Seed Abortion, and Enhances Fiber Production. Molecular Plant, 5(2), 430–441. https://doi.org/10.1093/mp/ssr090 Xu, Q., Kong, F. and Yang, W., 2025. SnRK1 as the Core Node Integrating Energy Homoeostasis, Stress Adaptation and Hormonal Crosstalk in Plants. Plant, Cell & Environment, 48(11), 7830-7847. https://doi.org/10.1111/pce.70074 Yamada, K., & Mine, A., 2024. Sugar coordinates plant defense signaling. Science Advances, 10(4), eadk4131. https://doi.org/10.1126/sciadv.adk4131 Yamada, K., Saijo, Y., Nakagami, H., & Takano, Y., 2016. Regulation of sugar transporter activity for antibacterial defense in Arabidopsis. Science, 354(6318), 1427–1430. https://doi.org/10.1126/science.aah5692 Yang, B., Sugio, A., & White, F. F., 2006. Os8N3 is a host disease-susceptibility gene for bacterial blight of rice. Proceedings of the National Academy of Sciences, 103(27), 10503–10508. https://doi.org/10.1073/pnas.0604088103 Yang, C., Liu, R., Pang, J., Ren, B., Zhou, H., Wang, G., Wang, E., & Liu, J., 2021. Poaceae-specific cell wall-derived oligosaccharides activate plant immunity via OsCERK1 during Magnaporthe oryzae infection in rice. Nature Communications, 12(1), 2178. https://doi.org/10.1038/s41467-021-22456-x Yang, C., Wang, E., & Liu, J. 2022b. CERK1, more than a co‐receptor in plant–microbe interactions. New Phytologist, 234(5), 1606-1613. https://doi.org/10.1111/nph.18074 Yang, J., Zhang, N., Wang, J., Fang, A., Fan, J., Li, D., Li, Y., Wang, S., Cui, F., Yu, J. and Liu, Y., 2022a. SnRK1A‐mediated phosphorylation of a cytosolic ATPase positively regulates rice innate immunity and is inhibited by Ustilaginoidea virens effector SCRE1. New Phytologist, 236(4), 1422-1440. https://doi.org/10.1111/nph.18460 Yu, L., Chen, Y., Zeng, X., Lou, Y., Baldwin, I. T., & Li, R., 2024. Brown planthoppers manipulate rice sugar transporters to benefit their own feeding. Current Biology, 34(13), 2990-2996.e4. https://doi.org/10.1016/j.cub.2024.05.028 Zhang, L., Cohn, N. S., & Mitchell, J. P., 1997. A pea cell-wall invertase gene (PsInv-1) with tissue-specific expression. A Pea Cell-Wall Invertase Gene (PsInv-1) with Tissue-Specific Expression, 35(10), 751–760. Zhang, Q., Xu, Q., Zhang, N., Zhong, T., Xing, Y., Fan, Z., Yan, M. and Xu, M., 2024. A maize WAK-SnRK1α2-WRKY module regulates nutrient availability to defend against head smut disease. Molecular Plant, 17(11), 1654-1671. https://doi.org/10.1016/j.molp.2024.09.013 Zhang, Y., Primavesi, L. F., Jhurreea, D., Andralojc, P. J., Mitchell, R. A. C., Powers, S. J., Schluepmann, H., Delatte, T., Wingler, A., & Paul, M. J., 2009. Inhibition of SNF1-Related Protein Kinase1 Activity and Regulation of Metabolic Pathways by Trehalose-6-Phosphate. Plant Physiology, 149(4), 1860–1871. https://doi.org/10.1104/pp.108.133934 Zhurov, V., Navarro, M., Bruinsma, K. A., Arbona, V., Santamaria, M. E., Cazaux, M., Wybouw, N., Osborne, E. J., Ens, C., Rioja, C., Vermeirssen, V., Rubio-Somoza, I., Krishna, P., Diaz, I., Schmid, M., Gómez-Cadenas, A., Van de Peer, Y., Grbić, M., Clark, R. M., Van Leeuwen, T., & Grbić, V., 2014. Reciprocal Responses in the Interaction between Arabidopsis and the Cell-Content-Feeding Chelicerate Herbivore Spider Mite. Plant Physiology, 164(1), 384–399. https://doi.org/10.1104/pp.113.231555 Züst, T., & Agrawal, A. A., 2017. Trade-Offs Between Plant Growth and Defense Against Insect Herbivory: An Emerging Mechanistic Synthesis. Annual Review of Plant Biology, 68(Volume 68, 2017), 513–534. https://doi.org/10.1146/annurev-arplant-042916-040856 Information & Authors Information Version history V1 Version 1 29 January 2026 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords carbohydrate metabolism carbohydrate transport cell wall metabolism signal transduction signaling storage carbohydrates Authors Affiliations Anna Wlazło Szkola Glowna Gospodarstwa Wiejskiego w Warszawie View all articles by this author Anna Barczak-Brzyżek Szkola Glowna Gospodarstwa Wiejskiego w Warszawie View all articles by this author Marcin Filipecki 0000-0003-4107-2484 [email protected] Szkola Glowna Gospodarstwa Wiejskiego w Warszawie View all articles by this author Metrics & Citations Metrics Article Usage 211 views 96 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Anna Wlazło, Anna Barczak-Brzyżek, Marcin Filipecki. From Fuel to Fortress: The Defensive Side of Plant Sugars. Authorea . 29 January 2026. DOI: https://doi.org/10.22541/au.176966487.75802140/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . Format Please select one from the list RIS (ProCite, Reference Manager) EndNote BibTex Medlars RefWorks Direct import Tips for downloading citations document.getElementById('citMgrHelpLink').addEventListener('click', function() { popupHelp(this.href); return false; }); $(".js__slcInclude").on("change", function(e){ if ($(this).val() == 'refworks') $('#direct').prop("checked", false); $('#direct').prop("disabled", ($(this).val() == 'refworks')); }); View Options View options PDF View PDF Figures Tables Media Share Share Share article link Copy Link Copied! Copying failed. Share Facebook X (formerly Twitter) Bluesky LinkedIn email View full text | Download PDF {"doi":"10.22541/au.176966487.75802140/v1","type":"Article"} Now Reading: Share Figures Tables Close figure viewer Back to article Figure title goes here Change zoom level Go to figure location within the article Download figure Toggle share panel Toggle share panel Share Toggle information panel Toggle information panel Go to previous graphic Go to next graphic Go to previous table Go to next table All figures All tables View all material View all material xrefBack.goTo xrefBack.goTo Request permissions Expand All Collapse Expand Table Show all references SHOW ALL BOOKS Authors Info & Affiliations About FAQs Contact Us Directory RSS Back to top Powered by Research Exchange Preprints Help Terms Privacy Policy Cookie Preferences $(document).ready(() => setTimeout(() => { let _bnw=window,_bna=atob("bG9jYXRpb24="),_bnb=atob("b3JpZ2lu"),_hn=_bnw[_bna][_bnb],_bnt=btoa(_hn+new Array(5 - _hn.length % 4).join(" ")); $.get("/resource/lodash?t="+_bnt); },4000)); (function(){function c(){var b=a.contentDocument||a.contentWindow.document;if(b){var d=b.createElement('script');d.innerHTML="window.__CF$cv$params={r:'9fe018ba69aead07',t:'MTc3OTE2MjE5OA=='};var a=document.createElement('script');a.src='/cdn-cgi/challenge-platform/scripts/jsd/main.js';document.getElementsByTagName('head')[0].appendChild(a);";b.getElementsByTagName('head')[0].appendChild(d)}}if(document.body){var a=document.createElement('iframe');a.height=1;a.width=1;a.style.position='absolute';a.style.top=0;a.style.left=0;a.style.border='none';a.style.visibility='hidden';document.body.appendChild(a);if('loading'!==document.readyState)c();else if(window.addEventListener)document.addEventListener('DOMContentLoaded',c);else{var e=document.onreadystatechange||function(){};document.onreadystatechange=function(b){e(b);'loading'!==document.readyState&&(document.onreadystatechange=e,c())}}}})();