Mechanisms Involving Plant Cell Walls in the Immune Response and Its In Situ Non-labeled Imaging Technique

doi:10.11983/CBB25034

Abstract

Abstract: The plant cell wall, which is composed of cellulose, hemicellulose, pectin and lignin, is a dynamically changing network structure, that not only plays the role of a key line of defense in the process of plant resistance to external pressure and adaptation to environmental changes, but also plays the role of an information hub in the process of signal transmission. When the cell wall is damaged, cells sense cell wall changes and initiate early immune responses, such as hormonal changes, alterations in wall composition and modifications, and the production of disease-resistant secondary metabolites. Although the importance of the cell wall in plant immunity is widely recognized, the specific molecular mechanisms by which cell wall damage triggers immune responses remain poorly understood. The application of in situ unlabeled imaging techniques in plant cells is gradually increasing and has become an important tool for studying cell wall structure and function. This paper describes the interaction mechanism between the plant cell wall and the immune response to provide a scientific basis for a deeper understanding of plant life activities and improve crop disease resistance, and describes in situ non-labeled imaging of the cell wall to provide more technological options for advancing the study of the cell wall in the immune response.

Key words: plant cell wall, plant immunity, signal sensing and transmission, in situ non-labeled imaging techniques

Wang Xiao, Xu Changwen, Qian Hongping, Li Sibo, Lin Jinxing, Cui Yaning. Mechanisms Involving Plant Cell Walls in the Immune Response and Its In Situ Non-labeled Imaging Technique[J]. Chinese Bulletin of Botany, 2025, 60(5): 773-785.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.chinbullbotany.com/EN/10.11983/CBB25034

https://www.chinbullbotany.com/EN/Y2025/V60/I5/773

Figures/Tables 3

References 87

[1]	Albersheim P, Anderson AJ (1971). Proteins from plant cell walls inhibit polygalacturonases secreted by plant pathogens. Proc Natl Acad Sci USA 68, 1815-1819. PMID
[2]	Alonso S, Gautam K, Iglesias-Moya J, Martínez C, Jamilena M (2024). Crosstalk between ethylene, jasmonate and ABA in response to salt stress during germination and early plant growth in Cucurbita pepo. Int J Mol Sci 25, 8728.
[3]	Azelee NIW, Mahdi HI, Cheng YS, Nordin N, Illias RM, Rahman RA, Shaarani SM, Bhatt P, Yadav S, Chang SW, Ravindran B, Ashokkumar V (2023). Biomass degradation: challenges and strategies in extraction and fractionation of hemicellulose. Fuel 339, 126982.
[4]	Aziz A, Gauthier A, Bézier A, Poinssot B, Joubert JM, Pugin A, Heyraud A, Baillieul F (2007). Elicitor and resistance-inducing activities of β-1,4 cellodextrins in grapevine, comparison with β-1,3 glucans and α-1,4 oligogalacturonides. J Exp Bot 58, 1463-1472. PMID
[5]	Babosha AV (2008). Inducible lectins and plant resistance to pathogens and abiotic stress. Biochemistry 73, 812-825.
[6]	Berglund J, Mikkelsen D, Flanagan BM, Dhital S, Gaunitz S, Henriksson G, Lindström ME, Yakubov GE, Gidley MJ, Vilaplana F (2020). Wood hemicelluloses exert distinct biomechanical contributions to cellulose fibrillar networks. Nat Commun 11, 4692. DOI PMID
[7]	Bot P, Mun BG, Imran QM, Hussain A, Lee SU, Loake G, Yun BW (2019). Differential expression of AtWAKL10 in response to nitric oxide suggests a putative role in biotic and abiotic stress responses. PeerJ 7, e7383.
[8]	Bronkhorst J, Kots K, De Jong D, Kasteel M, Van Boxmeer T, Joemmanbaks T, Govers F, van der Gucht J, Ketelaar T, Sprakel J (2022). An actin mechanostat ensures hyphal tip sharpness in Phytophthora infestans to achieve host penetration. Sci Adv 8, eabo0875.
[9]	Brutus A, Sicilia F, Macone A, Cervone F, De Lorenzo G (2010). A domain swap approach reveals a role of the plant wall-associated kinase 1 (WAK1) as a receptor of oligogalacturonides. Proc Natl Acad Sci USA 107, 9452-9457. DOI PMID
[10]	Bugbee WM (1993). A pectin lyase inhibitor protein from cell walls of sugar beet. Phytopathology 83, 63-68.
[11]	Carpita NC, Gibeaut DM (1993). Structural models of primary cell walls in flowering plants: consistency of molecular structure with the physical properties of the walls during growth. Plant J 3, 1-30. DOI PMID
[12]	Cayrol B, Delteil A, Gobbato E, Kroj T, Morel JB (2016). Three wall-associated kinases required for rice basal immunity form protein complexes in the plasma membrane. Plant Signal Behav 11, e1149676.
[13]	Cervone F, Ausubel FM, De Lorenzo G (2015). Enhancing immunity by engineering DAMPs. Oncotarget 6, 28523-28524. DOI PMID
[14]	Chen J, Xu F, Qiang XN, Liu HB, Wang L, Jiang LL, Li CY, Wang BQ, Luan S, Wu DS, Zhou F, Yu F (2024). Regulated cleavage and translocation of FERONIA control immunity in Arabidopsis roots. Nat Plants 10, 1761-1774. DOI PMID
[15]	Claverie J, Balacey S, Lemaître-Guillier C, Brulé D, Chiltz A, Granet L, Noirot E, Daire X, Darblade B, Héloir MC, Poinssot B (2018). The cell wall-derived xyloglucan is a new DAMP triggering plant immunity in Vitis vinifera and Arabidopsis thaliana. Front Plant Sci 9, 1725. DOI PMID
[16]	Cosgrove DJ (2001). Wall structure and wall loosening. A look backwards and forwards. Plant Physiol 125, 131-134. PMID
[17]	Cosgrove DJ (2024). Structure and growth of plant cell walls. Nat Rev Mol Cell Biol 25, 340-358.
[18]	Cui XM, Ji DC, Chen T, Tian SP (2016). Advances in the studies on molecular mechanism of receptor-like protein kinase FER regulating host plant-pathogen interaction. Chin Bull Bot 56, 339-346. (in Chinese)
	崔晓敏, 季东超, 陈彤, 田世平 (2016). 类受体激酶FER调节植物与病原菌相互作用的分子机制. 植物学报 56, 339-346.
[19]	Cui YN, Qian HP, Zhao YX, Li XJ (2020). Intracellular trafficking in pattern recognition receptor-triggered plant immunity. Chin Bull Bot 55, 329-339. (in Chinese)
	崔亚宁, 钱虹萍, 赵艳霞, 李晓娟 (2020). 模式识别受体的胞内转运及其在植物免疫中的作用. 植物学报 55, 329-339. DOI
[20]	Das S, Bhati V, Dewangan BP, Gangal A, Mishra GP, Dikshit HK, Pawar PAM (2024). Combining Fourier-transform infrared spectroscopy and multivariate analysis for chemotyping of cell wall composition in mungbean (Vigna radiata (L.) Wizcek). Plant Methods 20, 135.
[21]	de Azevedo Souza C, Li SD, Lin AZ, Boutrot F, Grossmann G, Zipfel C, Somerville SC (2017). Cellulose-derived oligomers act as damage-associated molecular patterns and trigger defense-like responses. Plant Physiol 173, 2383-2398. DOI PMID
[22]	De Lorenzo G, Ferrari S, Giovannoni M, Mattei B, Cervone F (2019). Cell wall traits that influence plant development, immunity, and bioconversion. Plant J 97, 134-147. DOI
[23]	de Souza WR, Mitchell RAC, Cesarino I (2023). Editorial: the plant cell wall: advances and current perspectives. Front Plant Sci 14, 1235749.
[24]	Delteil A, Gobbato E, Cayrol B, Estevan J, Michel-Romiti C, Dievart A, Kroj T, Morel JB (2016). Several wall-associated kinases participate positively and negatively in basal defense against rice blast fungus. BMC Plant Biol 16, 17. DOI PMID
[25]	Diener AC, Ausubel FM (2005). RESISTANCE TO FUSARIUM OXYSPORUM 1, a dominant Arabidopsis disease- resistance gene, is not race specific. Genetics 171, 305-321.
[26]	Dong BX, Liu Y, Huang G, Song AP, Chen SM, Jiang JF, Chen FD, Fang WM (2024). Plant NAC transcription factors in the battle against pathogens. BMC Plant Biol 24, 958. DOI PMID
[27]	Du J, Anderson CT, Xiao CW (2022). Dynamics of pectic homogalacturonan in cellular morphogenesis and adhesion, wall integrity sensing and plant development. Nat Plants 8, 332-340. DOI PMID
[28]	Engelsdorf T, Gigli-Bisceglia N, Veerabagu M, McKenna JF, Vaahtera L, Augstein F, Van der Does D, Zipfel C, Hamann T (2018). The plant cell wall integrity maintenance and immune signaling systems cooperate to control stress responses in Arabidopsis thaliana. Sci Signal 11, eaao3070.
[29]	Ferrari S, Savatin DV, Sicilia F, Gramegna G, Cervone F, De Lorenzo G (2013). Oligogalacturonides: plant damage-associated molecular patterns and regulators of growth and development. Front Plant Sci 4, 49. DOI PMID
[30]	Grantham NJ, Wurman-Rodrich J, Terrett OM, Lyczakowski JJ, Stott K, Iuga D, Simmons TJ, Durand-Tardif M, Brown SP, Dupree R, Busse-Wicher M, Dupree P (2017). An even pattern of xylan substitution is critical for interaction with cellulose in plant cell walls. Nat Plants 3, 859-865. DOI PMID
[31]	Guo YY, Wang SF, Yu KJ, Wang HL, Xu HM, Song CW, Zhao YY, Wen JL, Fu CX, Li Y, Wang SZ, Zhang X, Zhang Y, Cao Y, Shao FJ, Wang XH, Deng X, Chen T, Zhao Q, Li L, Wang GD, Grünhofer P, Schreiber L, Li Y, Song GY, Dixon RA, Lin JX (2023). Manipulating microRNA miR408 enhances both biomass yield and saccharification efficiency in poplar. Nat Commun 14, 4285.
[32]	He Q, Zabotina OA, Yu CX (2020). Principal component analysis facilitated fast and noninvasive Raman spectroscopic imaging of plant cell wall pectin distribution and interaction with enzymatic hydrolysis. J Raman Spectrosc 51, 2458-2467.
[33]	Jin KX, Liu XG, Jiang ZH, Tian GL, Yang SM, Shang LL, Ma JF (2019). Delignification kinetics and selectivity in poplar cell wall with acidified sodium chlorite. Ind Crops Prod 136, 87-92.
[34]	Kang X, Kirui A, Dickwella Widanage MC, Mentink-Vigier F, Cosgrove DJ, Wang T (2019). Lignin-polysaccharide interactions in plant secondary cell walls revealed by solid-state NMR. Nat Commun 10, 347. DOI PMID
[35]	Ke JJ, Zhu WT, Yuan Y, Du XY, Xu A, Zhang D, Cao S, Chen W, Lin Y, Xie JT, Cheng JS, Fu YP, Jiang DH, Yu X, Li B (2023). Duality of immune recognition by tomato and virulence activity of the Ralstonia solanacearum exo-polygalacturonase PehC. Plant Cell 35, 2552-2569.
[36]	Keegstra K (2010). Plant cell walls. Plant Physiol 154, 483-486. DOI PMID
[37]	Keegstra K, Talmadge KW, Bauer WD, Albersheim P (1973). The structure of plant cell walls: III. A model of the walls of suspension-cultured sycamore cells based on the interconnections of the macromolecular components. Plant Physiol 51, 188-197. DOI PMID
[38]	Lagaert S, Beliën T, Volckaert G (2009). Plant cell walls: protecting the barrier from degradation by microbial enzymes. Semin Cell Dev Biol 20, 1064-1073. DOI PMID
[39]	Larkan NJ, Ma LS, Haddadi P, Buchwaldt M, Parkin IAP, Djavaheri M, Borhan MH (2020). The Brassica napus wall-associated kinase-like (WAKL) gene Rlm9 provides race-specific blackleg resistance. Plant J 104, 892-900.
[40]	Lee HK, Santiago J (2023). Structural insights of cell wall integrity signaling during development and immunity. Curr Opin Plant Biol 76, 102455.
[41]	Li DD, Li XM, Wang ZC, Wang HC, Gao JZ, Liu XT, Zhang Z (2024a). Transcription factors RhbZIP17 and RhWRKY30 enhance resistance to Botrytis cinerea by increasing lignin content in rose petals. J Exp Bot 75, 1633-1646.
[42]	Li YJ, Shen WW, Zhang X, Cui YN, Zhao YY, Guo YY, Li XJ, Wang SZ, Song GY, Wang P, Ma JF, Lin JX (2024b). Single-cell characterization of major components of plant cell walls in situ by Raman spectroscopy. Sci China Life Sci 67, 1772-1774.
[43]	Lin WW, Tang WX, Pan X, Huang AB, Gao XQ, Anderson CT, Yang ZB (2022). Arabidopsis pavement cell morphogenesis requires FERONIA binding to pectin for activation of ROP GTPase signaling. Curr Biol 32, 497-507.
[44]	Liu MCJ, Yeh FLJ, Yvon R, Simpson K, Jordan S, Chambers J, Wu HM, Cheung AY (2024). Extracellular pectin-RALF phase separation mediates FERONIA global signaling function. Cell 187, 312-330.
[45]	Malinovsky FG, Fangel JU, Willats WGT (2014). The role of the cell wall in plant immunity. Front Plant Sci 5, 178. DOI PMID
[46]	Man Y, Zhao YY, Ye R, Lin JX, Jing YP (2018). In vivo cytological and chemical analysis of Casparian strips using stimulated Raman scattering microscopy. J Plant Physiol 220, 136-144.
[47]	Molina A, Jordá L, Torres MÁ, Martín-Dacal M, Berlanga DJ, Fernández-Calvo P, Gómez-Rubio E, Martín-Santamaría S (2024). Plant cell wall-mediated disease resistance: current understanding and future perspectives. Mol Plant 17, 699-724. DOI PMID
[48]	Munzert KS, Engelsdorf T (2025). Plant cell wall structure and dynamics in plant-pathogen interactions and pathogen defence. J Exp Bot 76, 228-242.
[49]	Peaucelle A, Braybrook SA, Le Guillou L, Bron E, Kuhlemeier C, Höfte H (2011). Pectin-induced changes in cell wall mechanics underlie organ initiation in Arabidopsis. Curr Biol 21, 1720-1726. DOI PMID
[50]	Philippe G, Sørensen I, Jiao C, Sun XP, Fei ZJ, Domozych DS, Rose JK (2020). Cutin and suberin: assembly and origins of specialized lipidic cell wall scaffolds. Curr Opin Plant Biol 55, 11-20. DOI PMID
[51]	Rawat S, Ali S, Nayankantha NNC, Chandrashekar N, Mittra B, Grover A (2017). Isolation and expression analysis of defensin gene and its promoter from Brassica juncea. J Plant Dis Prot 124, 591-600.
[52]	Reymond P, Farmer EE (1998). Jasmonate and salicylate as global signals for defense gene expression. Curr Opin Plant Biol 1, 404-411. DOI PMID
[53]	Rongpipi S, Ye D, Gomez ED, Gomez EW (2019). Progress and opportunities in the characterization of cellulose—an important regulator of cell wall growth and mechanics. Front Plant Sci 9, 1894.
[54]	Shine MB, Yang JW, El-Habbak M, Nagyabhyru P, Fu DQ, Navarre D, Ghabrial S, Kachroo P, Kachroo A (2016). Cooperative functioning between phenylalanine ammonia lyase and isochorismate synthase activities contributes to salicylic acid biosynthesis in soybean. New Phytol 212, 627-636. DOI PMID
[55]	Silverman P, Seskar M, Kanter D, Schweizer P, Metraux JP, Raskin I (1995). Salicylic acid in rice (biosynthesis, conjugation, and possible role). Plant Physiol 108, 633-639. DOI PMID
[56]	Simmons TJ, Mortimer JC, Bernardinelli OD, Pöppler AC, Brown SP, deAzevedo ER, Dupree R, Dupree P (2016). Folding of xylan onto cellulose fibrils in plant cell walls revealed by solid-state NMR. Nat Commun 7, 13902. DOI PMID
[57]	Song CW, Guo YY, Shen WW, Yao XM, Xu HM, Zhao YY, Li RL, Lin JX (2023). PagUNE12 encodes a basic helix- loop-helix transcription factor that regulates the development of secondary vascular tissue in poplar. Plant Physiol 192, 1046-1062.
[58]	Sultana R, Imam Z, Kumar RR, Banu VS, Nahakpam S, Bharti R, Bharadwaj C, Singh AK, Pasala RK, Singh DR, Siddiqui MW (2025). Signaling and defence mechanism of jasmonic and salicylic acid response in pulse crops: role of WRKY transcription factors in stress response. J Plant Growth Regul 44, 5-21.
[59]	Sun Y, Wang Y, Zhang XX, Chen ZD, Xia YQ, Wang L, Sun YJ, Zhang MM, Xiao Y, Han ZF, Wang YC, Chai JJ (2022). Plant receptor-like protein activation by a microbial glycoside hydrolase. Nature 610, 335-342.
[60]	Tang WX, Lin WW, Zhou X, Guo JZ, Dang X, Li BQ, Lin DS, Yang ZB (2022). Mechano-transduction via the pectin-FERONIA complex activates ROP6 GTPase signaling in Arabidopsis pavement cell morphogenesis. Curr Biol 32, 508-517.
[61]	Wan JX, He M, Hou QQ, Zou LJ, Yang YH, Wei Y, Chen XW (2021). Cell wall associated immunity in plants. Stress Biol 1, 3. DOI PMID
[62]	Wang WB, Fei Y, Wang YJ, Song BB, Li L, Zhang WJ, Cheng HY, Zhang XJ, Chen S, Zhou JM (2023). SHOU4/4L link cell wall cellulose synthesis to pattern-triggered immunity. New Phytol 238, 1620-1635.
[63]	Wang Y, Xu YP, Sun YJ, Wang HB, Qi JM, Wan BW, Ye WW, Lin YC, Shao YY, Dong SM, Tyler BM, Wang YC (2018). Leucine-rich repeat receptor-like gene screen reveals that Nicotiana RXEG1 regulates glycoside hydrolase 12 MAMP detection. Nat Commun 9, 594. DOI PMID
[64]	Wasternack C, Strnad M (2016). Jasmonate signaling in plant stress responses and development-active and inactive compounds. New Biotechnol 33, 604-613. DOI PMID
[65]	Wildermuth MC, Dewdney J, Wu G, Ausubel FM (2001). Isochorismate synthase is required to synthesize salicylic acid for plant defence. Nature 414, 562-565.
[66]	Wojtasik W, Kulma A, Dymińska L, Hanuza J, Czemplik M, Szopa J (2016). Evaluation of the significance of cell wall polymers in flax infected with a pathogenic strain of Fusarium oxysporum. BMC Plant Biol 16, 75. DOI PMID
[67]	Wormit A, Usadel B (2018). The multifaceted role of pectin methylesterase inhibitors (PMEIs). Int J Mol Sci 19, 2878.
[68]	Wu HC, Bulgakov VP, Jinn TL (2018). Pectin methylesterases: cell wall remodeling proteins are required for plant response to heat stress. Front Plant Sci 9, 1612.
[69]	Xia YQ, Sun GZ, Xiao JH, He XY, Jiang HB, Zhang ZC, Zhang Q, Li KN, Zhang SC, Shi XC, Wang ZY, Liu L, Zhao Y, Yang YH, Duan KX, Ye WW, Wang YM, Dong SM, Wang Y, Ma ZC, Wang YC (2024). AlphaFold-guided redesign of a plant pectin methylesterase inhibitor for broad-spectrum disease resistance. Mol Plant 17, 1344-1368.
[70]	Xiao Y, Sun GZ, Yu QS, Gao T, Zhu QS, Wang R, Huang SJ, Han ZF, Cervone F, Yin H, Qi TC, Wang YC, Chai JJ (2024). A plant mechanism of hijacking pathogen virulence factors to trigger innate immunity. Science 383, 732-739. DOI PMID
[71]	Yalpani N, Leon J, Lawton MA, Raskin I (1993). Pathway of salicylic acid biosynthesis in healthy and virus-inoculated tobacco. Plant Physiol 103, 315-321. DOI PMID
[72]	Yang C, Liu R, Pang JH, Ren B, Zhou HB, Wang G, Wang ET, Liu J (2021a). Poaceae-specific cell wall-derived oligosaccharides activate plant immunity via OsCERK1 during Magnaporthe oryzae infection in rice. Nat Commun 12, 2178.
[73]	Yang J, Xie MX, Wang XF, Wang GN, Zhang Y, Li ZK, Ma ZY (2021b). Identification of cell wall-associated kinases as important regulators involved in Gossypium hirsutum resistance to Verticillium dahliae. BMC Plant Biol 21, 220.
[74]	Yang K, Qi L, Zhang ZY (2014). Isolation and characterization of a novel wall-associated kinase gene TaWAK5 in wheat (Triticum aestivum). Crop J 2, 255-266. DOI
[75]	Yang P, Praz C, Li BB, Singla J, Robert CAM, Kessel B, Scheuermann D, Lüthi L, Ouzunova M, Erb M, Krattinger SG, Keller B (2019). Fungal resistance mediated by maize wall-associated kinase ZmWAK-RLK1 correlates with reduced benzoxazinoid content. New Phytol 221, 976-987. DOI PMID
[76]	Yao XM, Zhang GF, Zhang G, Sun Q, Liu CM, Chu JF, Jing YP, Niu SH, Fu CX, Lew TTS, Lin JX, Li XJ (2024). PagARGOS promotes low-lignin wood formation in poplar. Plant Biotechnol J 22, 2201-2215. DOI PMID
[77]	Zeng YN, Yarbrough JM, Mittal A, Tucker MP, Vinzant TB, Decker SR, Himmel ME (2016). In situ label-free imaging of hemicellulose in plant cell walls using stimulated Raman scattering microscopy. Biotechnol Biofuels 9, 256.
[78]	Zhai K, Rhodes J, Zipfel C (2024). A peptide-receptor module links cell wall integrity sensing to pattern-triggered immunity. Nat Plants 10, 1799-1811.
[79]	Zhang BC, Gao YH, Zhang LJ, Zhou YH (2021a). The plant cell wall: biosynthesis, construction, and functions. J Integr Plant Biol 63, 251-272.
[80]	Zhang DM, Xu ZP, Cao SX, Chen KL, Li SC, Liu XL, Gao CX, Zhang BC, Zhou YH (2018). An uncanonical CCCH- tandem zinc-finger protein represses secondary wall synthesis and controls mechanical strength in rice. Mol Plant 11, 163-174.
[81]	Zhang LS, Kars I, Essenstam B, Liebrand TWH, Wagemakers L, Elberse J, Tagkalaki P, Tjoitang D, van den Ackerveken G, van Kan JAL (2014). Fungal endopolygalacturonases are recognized as microbe-associated molecular patterns by the Arabidopsis receptor-like protein RESPONSIVENESS TO BOTRYTIS POLYGALACTURONASES1. . Plant Physiol 164, 352-364.
[82]	Zhang N, Pombo MA, Rosli HG, Martin GB (2020). Tomato wall-associated kinase SLWak1 depends on Fls2/Fls3 to promote apoplastic immune responses to Pseudomonas syringae. Plant Physiol 183, 1869-1882. DOI PMID
[83]	Zhang ZQ, Ma WY, Ren ZY, Wang XX, Zhao JJ, Pei XY, Liu YG, He KL, Zhang F, Huo WQ, Li W, Yang DG, Ma XF (2021b). Characterization and expression analysis of wall-associated kinase (WAK) and WAK-like family in cotton. Int J Biol Macromol 187, 867-879.
[84]	Zhao YY, Man Y, Wen JL, Guo YY, Lin JX (2019). Advances in imaging plant cell walls. Trends Plant Sci 24, 867-878. DOI PMID
[85]	Zhou JG, Mu Q, Wang XY, Zhang J, Yu HZ, Huang TZ, He YX, Dai SJ, Meng XZ (2022). Multilayered synergistic regulation of phytoalexin biosynthesis by ethylene, jasmonate, and MAPK signaling pathways in Arabidopsis. Plant Cell 34, 3066-3087.
[86]	Zhu JW, Ren WT, Guo F, Wang HK, Yu Y (2024). Revealing spatial distribution and accessibility of cell wall polymers in bamboo through chemical imaging and mild chemical treatments. Carbohydr Polym 339, 122261.
[87]	Zhu N, Yang YF, Ji MB, Wu D, Chen KS (2019). Label-free visualization of lignin deposition in loquats using complementary stimulated and spontaneous Raman microscopy. Hortic Res 6, 72.

	基因(编码蛋白)	作用	识别病原体后的变化	参考文献
纤维素	CesA (cellulose synthase)	正向调节纤维素合成	下调, 纤维素合成减少	Wojtasik et al., 2016; Molina et al., 2024
半纤维素	GMT (glucomannan 4β-mannosyltransferase) GGT (galactomannan galactosyltransferase) XXT (xyloglucan xylosyltransferase) XYN (endo-1,4-β-xylanase)	正向调节半纤维素合成	下调, 半纤维素合成减少	Wojtasik et al., 2016; Molina et al., 2024
半纤维素	XYLa (1,4-α-xylosidase) GS (α-galactosidase)	正向调节半纤维素降解	上调, 半纤维素降解增多
果胶	GAE (UDP-glucuronate 4-epimerase) GAUT1/7 (galacturonosyltransferase 1/7) RGXT (rhamnogalacturonan II xylosyltransferase) ARAD (arabinose transferase) PMT (pectin methyltransferase)	正向调节果胶合成	下调, 果胶合成减少	Wojtasik et al., 2016; Molina et al., 2024
果胶	PG (polygalacturonase) PaL (pectin lyase)	正向调节果胶裂解	上调, 果胶降解增多
木质素	PAL (phenylalanine ammonia lyase) HCT (p-hydroxycinnamoyl CoA: quinate/shikimate p-hydroxycinnamoyl transferase) CCoAOMT (caffeoyl-CoA O-methyltransferase) COMT (caffeic acid/5-hydroxyferulic acid 3/5-O-methyltransferase) SAD (synaptic dehydrogenase) GT (glucosyltransferase) CAD1 (cinnamyl alcohol dehydrogenase 1)	正向调节木质素合成	上调, 木质素合成增多	Wojtasik et al., 2016; Li et al., 2024a; Molina et al., 2024
水杨酸合成	ICS1 (isochorismate synthase 1) PAL (phenylalanine ammonia lyase) PAD4 (phytoalexin-deficient 4) EDS1 (enhanced disease susceptibility 1) SAGT1 (salicylic acid glucosyltransferase 1) BSMT1 (benzoic acid and salicylic acid methyltransferase 1)	与水杨酸(SA)合成相关, 参与植物防御反应	ICS1、PAD4、EDS1和SAGT1上调/BSMT1下调, SA合成增多	Silverman et al., 1995; Wildermuth et al., 2001; Dong et al., 2024
茉莉酸合成	VSP1 (vegetative storage protein 1) LOX1 (lipoxygenase 1) MYC2 (myelocytomatosis protein 2) AOC (allene oxide cyclase)	与茉莉酸(JA)合成相关, 参与植物防御反应	VSP1、LOX1和AOC上调/MYC2下调, JA合成增多	Wasternack and Strnad, 2016; Dong et al., 2024
乙烯合成	ACC (1-aminocyclopropane-1-carboxylic acid synthase)	与乙烯(ET)合成相关, 参与植物防御反应	上调, ET合成增多	Alonso et al., 2024
抗病次生代谢产物	PR (pathogenesis-related) PDF (plant defensin)	与抗病次生代谢产物生成有关	上调, 防御素和凝集素合成增多	Babosha, 2008; Rawat et al., 2017

	基因(编码蛋白)	作用	识别病原体后的变化	参考文献
纤维素	CesA (cellulose synthase)	正向调节纤维素合成	下调, 纤维素合成减少	Wojtasik et al., 2016; Molina et al., 2024
半纤维素	GMT (glucomannan 4β-mannosyltransferase) GGT (galactomannan galactosyltransferase) XXT (xyloglucan xylosyltransferase) XYN (endo-1,4-β-xylanase)	正向调节半纤维素合成	下调, 半纤维素合成减少	Wojtasik et al., 2016; Molina et al., 2024
半纤维素	XYLa (1,4-α-xylosidase) GS (α-galactosidase)	正向调节半纤维素降解	上调, 半纤维素降解增多
果胶	GAE (UDP-glucuronate 4-epimerase) GAUT1/7 (galacturonosyltransferase 1/7) RGXT (rhamnogalacturonan II xylosyltransferase) ARAD (arabinose transferase) PMT (pectin methyltransferase)	正向调节果胶合成	下调, 果胶合成减少	Wojtasik et al., 2016; Molina et al., 2024
果胶	PG (polygalacturonase) PaL (pectin lyase)	正向调节果胶裂解	上调, 果胶降解增多
木质素	PAL (phenylalanine ammonia lyase) HCT (p-hydroxycinnamoyl CoA: quinate/shikimate p-hydroxycinnamoyl transferase) CCoAOMT (caffeoyl-CoA O-methyltransferase) COMT (caffeic acid/5-hydroxyferulic acid 3/5-O-methyltransferase) SAD (synaptic dehydrogenase) GT (glucosyltransferase) CAD1 (cinnamyl alcohol dehydrogenase 1)	正向调节木质素合成	上调, 木质素合成增多	Wojtasik et al., 2016; Li et al., 2024a; Molina et al., 2024
水杨酸合成	ICS1 (isochorismate synthase 1) PAL (phenylalanine ammonia lyase) PAD4 (phytoalexin-deficient 4) EDS1 (enhanced disease susceptibility 1) SAGT1 (salicylic acid glucosyltransferase 1) BSMT1 (benzoic acid and salicylic acid methyltransferase 1)	与水杨酸(SA)合成相关, 参与植物防御反应	ICS1、PAD4、EDS1和SAGT1上调/BSMT1下调, SA合成增多	Silverman et al., 1995; Wildermuth et al., 2001; Dong et al., 2024
茉莉酸合成	VSP1 (vegetative storage protein 1) LOX1 (lipoxygenase 1) MYC2 (myelocytomatosis protein 2) AOC (allene oxide cyclase)	与茉莉酸(JA)合成相关, 参与植物防御反应	VSP1、LOX1和AOC上调/MYC2下调, JA合成增多	Wasternack and Strnad, 2016; Dong et al., 2024
乙烯合成	ACC (1-aminocyclopropane-1-carboxylic acid synthase)	与乙烯(ET)合成相关, 参与植物防御反应	上调, ET合成增多	Alonso et al., 2024
抗病次生代谢产物	PR (pathogenesis-related) PDF (plant defensin)	与抗病次生代谢产物生成有关	上调, 防御素和凝集素合成增多	Babosha, 2008; Rawat et al., 2017

技术	优点	缺点	应用	参考文献
傅里叶红外光谱(Fourier transform infrared, FTIR)	无损, 可获得多维信息, 适用范围广	空间分辨率受限, 水吸收影响成像, 不适合活体成像	研究细胞壁成分特性, 研究发育和抗病过程中的成分变化	Song et al., 2023; Das et al., 2024; Zhu et al., 2024
共聚焦拉曼光谱成像(confocal Raman microscopy, CRM)	高光谱和空间分辨率, 非侵入性, 分辨率可达微米级	成像速度慢, 拉曼散射效应弱, 荧光背景干扰限制成像质量	监测细胞壁成分变化	Jin et al., 2019; He et al., 2020; Guo et al., 2023
受激拉曼光谱成像(stimulated Raman scattering, SRS)	高成像速度, 可进行定量分析, 具有高空间分辨率和高信噪比	对待测物质丰度要求高, 设备成本高	实时监测高度动态过程中的细胞壁成分与结构变化	Zhu et al., 2019; Yao et al., 2024

技术	优点	缺点	应用	参考文献
傅里叶红外光谱(Fourier transform infrared, FTIR)	无损, 可获得多维信息, 适用范围广	空间分辨率受限, 水吸收影响成像, 不适合活体成像	研究细胞壁成分特性, 研究发育和抗病过程中的成分变化	Song et al., 2023; Das et al., 2024; Zhu et al., 2024
共聚焦拉曼光谱成像(confocal Raman microscopy, CRM)	高光谱和空间分辨率, 非侵入性, 分辨率可达微米级	成像速度慢, 拉曼散射效应弱, 荧光背景干扰限制成像质量	监测细胞壁成分变化	Jin et al., 2019; He et al., 2020; Guo et al., 2023
受激拉曼光谱成像(stimulated Raman scattering, SRS)	高成像速度, 可进行定量分析, 具有高空间分辨率和高信噪比	对待测物质丰度要求高, 设备成本高	实时监测高度动态过程中的细胞壁成分与结构变化	Zhu et al., 2019; Yao et al., 2024