花粉致敏相关组分及花粉壁物质组成研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2026-0468

摘要/Abstract

摘要：

花粉导致的过敏发病率逐年上升，对人类健康的影响日益显著。花粉粒是一个复杂的生物复合体系，其所含的免疫活性成分是引发、加剧过敏反应的关键因素。近年来，对花粉中的蛋白过敏原及非蛋白免疫调节剂的鉴定与功能研究取得了显著进展。此外，花粉壁作为花粉与外界接触的首要界面，其多层结构及丰富的物质组分在致敏过程中也发挥了重要作用，现已成为花粉致敏机制研究的重要关注点。染色标记、质谱组学及生物信息学等技术的发展，极大促进了对花粉壁结构与化学组成的解析。本文旨在揭示花粉粒作为致敏相关活性物质载体的结构复杂性，系统分析蛋白过敏原及非蛋白组分的免疫调节机制。同时，综述花粉壁的结构特征及其组分分析技术，以期为系统研究花粉致敏的根源及开发干预措施提供新思路。

关键词: 花粉致敏, 花粉壁, 致敏蛋白, 免疫调节剂

Abstract:

The increasing prevalence of pollen-induced allergies has become a significant public health concern, and its impact on human health is becoming increasingly pronounced. The pollen grain is a complex biological system that harbors immunologically active components, which are key factors in initiating and exacerbating allergic reactions. In recent years, systematic progress has been made in the identification and functional characterization of both pollen-derived allergenic proteins and non-protein immunomodulators in pollen. Furthermore, the pollen wall serving as the primary interface between pollen and the external environment, plays an important role in the sensitization process due to its multilayered structure and abundant material components, which has gradually attracted increasing attention. Advances in techniques including staining and labeling, mass spectrometry-based omics, and bioinformatics have greatly facilitated the elucidation of pollen wall architecture and chemical composition. This review aims to elucidate the structural complexity of the pollen grain as a carrier of sensitization-related active substances and to systematically analyze the immunomodulatory mechanisms of protein allergens and non-protein components. Meanwhile, it summarizes the structural features of the pollen wall and the analytical techniques used for its constituents analysis, with the goal of providing new insights for systematic investigation of the root causes of pollen sensitization and the development of intervention strategies.

Key words: pollen sensitization, pollen wall, allergenic proteins, immunomodulators

王雪晨, 姚丽娟, 王欣颖, 李晓娟. 花粉致敏相关组分及花粉壁物质组成研究进展[J]. 生物技术通报, doi: 10.13560/j.cnki.biotech.bull.1985.2026-0468.

WANG Xue-chen, YAO Li-juan, WANG Xin-ying, LI Xiao-juan. Research Progress on Pollen Sensitization-related Components and Material Composition of the Pollen Wall[J]. Biotechnology Bulletin, doi: 10.13560/j.cnki.biotech.bull.1985.2026-0468.

图/表 4

图1 花粉粒多层次结构示意图左侧展示花粉粒完整球形外观及表面纹饰，右侧为楔形立体剖面，清晰呈现花粉内部雄配子体及外部花粉壁（花粉涂层、外壁、内壁）的分层结构

Fig. 1 Schematic diagram of the multilayered structure of a pollen grainThe left panel shows the intact spherical morphology and surface ornamentation of the pollen grain, while the right panel presents a wedge-shaped three-dimensional section that clearly displays the internal male gametophyte and the layered structure of the pollen wall, including the pollen coat, exine, and intine

表1 花粉致敏相关的主要蛋白家族

Table 1 Major protein families related to pollen sensitization

分类 Category	蛋白家族 Protein family	花粉过敏原 Pollen allergen	生物学功能 Biological function
病程相关蛋白 Pathogenesis-related proteins （PR-proteins）	PR-2（内切葡聚糖酶）	Ole e 9	水解病原真菌细胞壁中的β-1,3-葡聚糖，释放免疫信号，调控胞间连丝，降解胼胝质促进花粉释放^［22］
	PR-3（几丁质酶I，II，IV-VII）	Cry j chitinase	降解真菌细胞壁的主要成分几丁质，抑制真菌生长，释放激发子激活免疫^［23］
	PR-5（类甜蛋白）	Cup a 3、Cup s 3、Jun a 3、Jun o 3、Jun v 3	增强某些可抑制丝氨酸肽链内切酶的蛋白质的活性，抵御真菌侵染，激活植物防卫反应信号通路^［24］
	PR-8（几丁质酶III）	Ziz m 1	具有水解细菌细胞壁的溶菌酶活性，参与植物防御反应^［25］
	PR-10（Bet v 1样蛋白）	Aln g 1、Bet v 1、Cor a 1	具RNase活性，抗多种生物与非生物胁迫，参与发育和激素信号调控^［26］
	PR-12（防御素）	Art v 1、Amb a 4	通过形成孔道抗真菌、病毒，参与植物防御信号网络，调控植物发育^［25］
	PR-14（脂质转移蛋白）	Ole e 7、Pla a 3、Art v 3	抑制病原菌，参与角质层形成和防御反应信号传导^［27］
细胞结构与生长调控蛋白 Cell structure and growth regulatory proteins	扩张蛋白	Cyn d 1、Lol p 1、Phl p 1、Zea m 1、Ory s1、Dac g1	调节细胞壁松弛，参与植物发育及胁迫响应^［28］
	肌动蛋白结合蛋白	Cyn d 12、Bet v 2、Ole e 2、Phl p 12、Zea m 12	调节肌动蛋白动态，调控细胞形态、运动和胞质流动，参与植物发育及胁迫响应^［29］
	EF-hand钙结合蛋白	Bet v 3/4、Phl p 7、Ole e 3/8、Aln g 4、Amb a 9/10	参与钙转运与储存、细胞信号传导、花粉管生长、细胞增殖与分化等^［30］
代谢相关酶类 Metabolism-related enzymes	果胶酸裂解酶	Amb a 1、Cry j 1、Cup s 1	催化果胶降解，参与植物发育、胁迫响应及细胞壁重塑^［31］
	核糖核酸酶	Phl p 5、Amb t 5	调控RNA的加工、成熟和降解，参与细胞防御^［32］
	多聚半乳糖醛酸酶	Cyn d 4	参与果胶降解，使细胞壁结构解体，导致果实软化、花粉成熟、病原物防御等^［33］

表1 花粉致敏相关的主要蛋白家族

Table 1 Major protein families related to pollen sensitization

分类 Category	蛋白家族 Protein family	花粉过敏原 Pollen allergen	生物学功能 Biological function
病程相关蛋白 Pathogenesis-related proteins （PR-proteins）	PR-2（内切葡聚糖酶）	Ole e 9	水解病原真菌细胞壁中的β-1,3-葡聚糖，释放免疫信号，调控胞间连丝，降解胼胝质促进花粉释放^［22］
	PR-3（几丁质酶I，II，IV-VII）	Cry j chitinase	降解真菌细胞壁的主要成分几丁质，抑制真菌生长，释放激发子激活免疫^［23］
	PR-5（类甜蛋白）	Cup a 3、Cup s 3、Jun a 3、Jun o 3、Jun v 3	增强某些可抑制丝氨酸肽链内切酶的蛋白质的活性，抵御真菌侵染，激活植物防卫反应信号通路^［24］
	PR-8（几丁质酶III）	Ziz m 1	具有水解细菌细胞壁的溶菌酶活性，参与植物防御反应^［25］
	PR-10（Bet v 1样蛋白）	Aln g 1、Bet v 1、Cor a 1	具RNase活性，抗多种生物与非生物胁迫，参与发育和激素信号调控^［26］
	PR-12（防御素）	Art v 1、Amb a 4	通过形成孔道抗真菌、病毒，参与植物防御信号网络，调控植物发育^［25］
	PR-14（脂质转移蛋白）	Ole e 7、Pla a 3、Art v 3	抑制病原菌，参与角质层形成和防御反应信号传导^［27］
细胞结构与生长调控蛋白 Cell structure and growth regulatory proteins	扩张蛋白	Cyn d 1、Lol p 1、Phl p 1、Zea m 1、Ory s1、Dac g1	调节细胞壁松弛，参与植物发育及胁迫响应^［28］
	肌动蛋白结合蛋白	Cyn d 12、Bet v 2、Ole e 2、Phl p 12、Zea m 12	调节肌动蛋白动态，调控细胞形态、运动和胞质流动，参与植物发育及胁迫响应^［29］
	EF-hand钙结合蛋白	Bet v 3/4、Phl p 7、Ole e 3/8、Aln g 4、Amb a 9/10	参与钙转运与储存、细胞信号传导、花粉管生长、细胞增殖与分化等^［30］
代谢相关酶类 Metabolism-related enzymes	果胶酸裂解酶	Amb a 1、Cry j 1、Cup s 1	催化果胶降解，参与植物发育、胁迫响应及细胞壁重塑^［31］
	核糖核酸酶	Phl p 5、Amb t 5	调控RNA的加工、成熟和降解，参与细胞防御^［32］
	多聚半乳糖醛酸酶	Cyn d 4	参与果胶降解，使细胞壁结构解体，导致果实软化、花粉成熟、病原物防御等^［33］

图2 花粉脂质相关介质的免疫调节通路花粉接触呼吸道上皮后先于蛋白过敏原释放PALMs：免疫刺激性类白三烯B4（LTB4-like）脂质募集并激活嗜酸性粒细胞（Eo）及多形核粒细胞（PMN），这些细胞释放颗粒蛋白、细胞因子、活性氧（ROS）等物质以构建促炎微环境；免疫调节性植物前列腺素E1（PPE1）抑制树突状细胞（DC）产生白细胞介素12（IL-12），为2型辅助性T细胞（Th2）极化创造条件。同时，DC表面的CD1分子能结合花粉磷脂，将其作为脂质抗原呈递给磷脂特异性T细胞，这些T细胞表现出Th2偏向的功能特征，通过分泌白细胞介素4（IL-4）辅助浆细胞产生免疫球蛋白E（IgE），从而促进过敏反应。花粉脂质还可上调DC表面CD1分子表达，进一步增强脂质抗原的呈递效率与iNKT细胞活化，最终协同参与Th2型过敏炎症

Fig. 2 Immunomodulatory pathways of pollen-associated lipid mediatorsUpon contacting respiratory epithelium, pollen releases PALMs prior to protein allergens. Immunostimulatory leukotriene B4 (LTB4)-like lipids recruit and activate eosinophil (Eo) and polymorphonuclear neutrophil (PMN), which release granule proteins, cytokines, and reactive oxygen species (ROS) to establish a pro-inflammatory microenvironment. Immunomodulatory phytoprostane E1 (PPE1) suppresses dendritic cell (DC) production of interleukin-12 (IL-12), favoring T helper 2 (Th2) polarization. Meanwhile, CD1 molecules on DCs binds pollen phospholipids and presents them as lipid antigens to phospholipid-specific T cells. These Th2-biased T cells secrete interleukin-4 (IL-4), supporting plasma cell immunoglobulin E (IgE) production and promoting the allergic response. Furthermore, pollen lipids upregulate CD1 molecules on DC, further enhancing lipid antigen presentation and iNKT cell activation, synergistically contributing to Th2 inflammation

图3 傅里叶变换拉曼光谱与共聚焦拉曼光谱原理对比示意图左栏：傅里叶变换拉曼光谱用于宏观分析，激光激发样品后，背向散射信号经干涉仪调制与FT探测器采集得到干涉图，再通过快速傅里叶变换（FFT）转换为拉曼光谱。右栏：共聚焦拉曼光谱用于微区分析，激光经聚焦后照射花粉微区，散射信号经共聚焦针孔滤波、色散光谱仪分光后由电荷耦合器（CCD）检测，实现精细亚结构的化学表征

Fig. 3 Schematic comparison of the principles of Fourier transform Raman spectroscopy and confocal Raman spectroscopyLeft panel: Fourier transform Raman spectroscopy is used for macroscopic analysis. After laser excitation of the sample, the backscattered signal is modulated by an interferometer and collected by an FT detector to generate an interferogram, which is then converted into a Raman spectrum via fast Fourier transform (FFT). Right panel: Confocal Raman spectroscopy is used for microanalysis. The laser beam is focused onto a micro-region of the pollen grain, and the scattered signal is filtered through a confocal pinhole, dispersed by a spectrometer, and detected by a charge-coupled device (CCD), enabling chemical characterization of fine substructures

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