植物凯氏带起源与功能分化研究进展
Research Progress on the Origin and Functional Differentiation of Plant Casparian Strip
摘要: 凯氏带是维管植物根部皮层细胞壁上特化的木质化带状增厚结构,作为根系质外体运输的关键屏障,在植物选择性吸收矿质元素、响应逆境胁迫以及植物–微生物互作中发挥着重要作用。该结构自被发现以来,长期被认为是维管植物高度保守的结构,但近年来研究逐渐揭示了其在不同植物类群间存在显著的结构、调控与功能分化。本文从进化发育生物学角度系统梳理相关研究进展,论述了凯氏带起源以及凯氏带形成的核心调控模块,阐明了凯氏带在不同植物类型中的发育特征和功能分化,包括内皮层和外皮层屏障的物种特异性、凯氏带响应非生物胁迫可塑性变化、凯氏带介导根系内生菌和根际微生物定殖与富集并参与共生固氮等过程。本文进一步探讨了现有研究的不足以及未来研究方向,以期为作物抗逆育种和养分高效利用提供理论参考。
Abstract: The casparian strip is a specialized lignified band-like thickening structure located in the cell walls of root cortical tissues in vascular plants. As a critical barrier regulating apoplastic transport in roots, it plays essential roles in the selective uptake of mineral nutrients, responses to abiotic stresses, and plant-microbe interactions. Since its discovery, the casparian strip has long been regarded as a highly conserved structure in vascular plants. However, recent studies have gradually revealed significant divergence in its structure, regulation, and function among different plant lineages. From the perspective of evolutionary developmental biology, this review systematically summarizes recent advances in the field, discusses the evolutionary origin of the casparian strip and the core regulatory modules governing its formation, and elucidates the developmental characteristics and functional diversification of casparian strips in different plant groups. These include species-specific differences in endodermal and exodermal barriers, the plastic responses of casparian strips to abiotic stresses, and the roles of casparian strips in mediating root endophyte and rhizosphere microbiota colonization and enrichment, as well as their involvement in symbiotic nitrogen fixation. Furthermore, this review discusses current limitations and future research directions, with the aim of providing theoretical references for crop breeding aimed at improving stress resistance and nutrient use efficiency.
文章引用:陈韵宁, 李唐语, 杨伟, 耿腊, 秦世雯, 黄立钰. 植物凯氏带起源与功能分化研究进展[J]. 植物学研究, 2026, 15(5): 301-310. https://doi.org/10.12677/br.2026.155035

参考文献

[1] Steudle, E. (2000) Water Uptake by Plant Roots: An Integration of Views. Plant and Soil, 226, 45-56.
https://doi.org/10.1023/a:1026439226716
[2] Steudle, E. (2000) Water Uptake by Roots: Effects of Water Deficit. Journal of Experimental Botany, 51, 1531-1542.
https://doi.org/10.1093/jexbot/51.350.1531
[3] 刘鑫, 王沛, 周青平. 植物根系质外体屏障研究进展[J]. 植物学报, 2021, 56(6): 761-773.
[4] Caspary, R. (1865) Remarks on the Protective Sheath and the Formation of Stems and Roots. Yearbooks for Scientific Botany, 4, 101‑124.
https://wp.unil.ch/geldnerlab/files/2013/06/Caspary1865.pdf
[5] Geldner, N. (2013) The Endodermis. Annual Review of Plant Biology, 64, 531-558.
https://doi.org/10.1146/annurev-arplant-050312-120050
[6] Naseer, S., Lee, Y., Lapierre, C., Franke, R., Nawrath, C. and Geldner, N. (2012) Casparian Strip Diffusion Barrier in Arabidopsis Is Made of a Lignin Polymer without Suberin. Proceedings of the National Academy of Sciences, 109, 10101-10106.
https://doi.org/10.1073/pnas.1205726109
[7] 嵇康轩. NaCl和外源ABA对百合生理生化及凯氏带发育的影响[D]: [硕士学位论文]. 苏州: 苏州大学, 2023.
[8] 刘梓清, 杨继刚, 吴子涵, 等. 植物根系限制重(类)金属吸收/转运的因素及其机制[J]. 农业现代化研究, 2021, 42(2): 284-293.
[9] 杨朝东, 张霞, 刘国锋, 等. 植物根中质外体屏障结构和生理功能研究进展[J]. 植物研究, 2013, 33(1): 114-119.
[10] 王平, 周青平, 王沛. 植物内皮层的分化及其屏障功能研究进展[J]. 西北植物学报, 2019, 39(4): 752-762.
[11] Manzano, C., Morimoto, K.W., Shaar-Moshe, L., Mason, G.A., Cantó-Pastor, A., Gouran, M., et al. (2024) Regulation and Function of a Polarly Localized Lignin Barrier in the Exodermis. Nature Plants, 11, 118-130.
https://doi.org/10.1038/s41477-024-01864-z
[12] Su, Y., Feng, T., Liu, C., Huang, H., Wang, Y., Fu, X., et al. (2023) The Evolutionary Innovation of Root Suberin Lamellae Contributed to the Rise of Seed Plants. Nature Plants, 9, 1968-1977.
https://doi.org/10.1038/s41477-023-01555-1
[13] 徐黎, 胡海琳, 王晨璨, 等. 植物根内皮层的发育与功能研究进展[J]. 科学通报, 2023, 68(8): 901-914.
[14] Uddin, N., Li, X., Ullah, M.W., Sethupathy, S., Ma, K., Zahoor, et al. (2024) Lignin Developmental Patterns and Casparian Strip as Apoplastic Barriers: A Review. International Journal of Biological Macromolecules, 260, Article 129595.
https://doi.org/10.1016/j.ijbiomac.2024.129595
[15] 张倩倩, 林晓艳, 顾钰峰, 等. 木贼类营养器官凯氏带类型的新观察[J]. 西北植物学报, 2021, 41(7): 1158-1164.
[16] 胡海琳. PagCIF1调控84K杨凯氏带发育的功能研究[D]: [硕士学位论文]. 北京: 北京林业大学, 2023.
[17] Karlova, R., Boer, D., Hayes, S. and Testerink, C. (2021) Root Plasticity under Abiotic Stress. Plant Physiology, 187, 1057-1070.
https://doi.org/10.1093/plphys/kiab392
[18] Roppolo, D., Boeckmann, B., Pfister, A., Boutet, E., Rubio, M.C., Dénervaud-Tendon, V., et al. (2014) Functional and Evolutionary Analysis of the CASPARIAN STRIP MEMBRANE DOMAIN PROTEIN Family. Plant Physiology, 165, 1709-1722.
https://doi.org/10.1104/pp.114.239137
[19] Xue, B., Liang, Z., Liu, Y., Li, D., Cao, P. and Liu, C. (2024) Comparative Analysis of Casparian Strip Membrane Domain Protein Family in Oryza sativa (L.) and Arabidopsis thaliana (L.). International Journal of Molecular Sciences, 25, Article 9858.
https://doi.org/10.3390/ijms25189858
[20] Xue, B., Liang, Z., Li, D., Liu, Y. and Liu, C. (2024) Genome-Wide Identification and Expression Analysis of CASPL Gene Family in Zea mays (L.). Frontiers in Plant Science, 15, Article ID: 1477383.
https://doi.org/10.3389/fpls.2024.1477383
[21] Wang, X., Zhang, Y., Wang, L., Pan, Z., He, S., Gao, Q., et al. (2020) Casparian Strip Membrane Domain Proteins in Gossypium arboreum: Genome-Wide Identification and Negative Regulation of Lateral Root Growth. BMC Genomics, 21, Article No. 340.
https://doi.org/10.1186/s12864-020-6723-9
[22] Barbosa, I.C.R., Rojas-Murcia, N. and Geldner, N. (2019) The Casparian Strip—One Ring to Bring Cell Biology to Lignification? Current Opinion in Biotechnology, 56, 121-129.
https://doi.org/10.1016/j.copbio.2018.10.004
[23] Doll, N.M. (2024) Filling the Gaps: Monitoring Casparian Strip Integrity in Rice. The Plant Cell, 36, 219-220.
https://doi.org/10.1093/plcell/koad274
[24] Fujita, S. (2021) Casparian Strip Integrity Factor (CIF) Family Peptides-Regulator of Plant Extracellular Barriers. Peptides, 143, Article 170599.
https://doi.org/10.1016/j.peptides.2021.170599
[25] Yang, J., Ding, C., Xu, B., Chen, C., Narsai, R., Whelan, J., et al. (2015) A Casparian Strip Domain-Like Gene, CASPL, Negatively Alters Growth and Cold Tolerance. Scientific Reports, 5, Article No. 14299.
https://doi.org/10.1038/srep14299
[26] Kamiya, T., Borghi, M., Wang, P., Danku, J.M.C., Kalmbach, L., Hosmani, P.S., et al. (2015) The MYB36 Transcription Factor Orchestrates Casparian Strip Formation. Proceedings of the National Academy of Sciences, 112, 10533-10538.
https://doi.org/10.1073/pnas.1507691112
[27] 李朋雪. 番茄根系内皮层凯氏带调控机制的研究[D]: [硕士学位论文]. 福州: 福建农林大学, 2018.
[28] 肖伊辉, 冯志航, 赖雨秋, 等. 植物根系内皮层凯氏带形成的分子机制及对矿质养分转运调控综述[J]. 浙江大学学报(农业与生命科学版), 2025, 51(3): 337-349.
[29] Li, P., Yang, M., Chang, J., Wu, J., Zhong, F., Rahman, A., et al. (2018) Spatial Expression and Functional Analysis of Casparian Strip Regulatory Genes in Endodermis Reveals the Conserved Mechanism in Tomato. Frontiers in Plant Science, 9, Article ID: 832.
https://doi.org/10.3389/fpls.2018.00832
[30] 张琳. 转录因子OsMYB36a调控水稻内皮层凯氏带形成的功能研究[D]: [硕士学位论文]. 南宁: 广西大学, 2024.
[31] Wang, Z., Zhang, B., Chen, Z., Wu, M., Chao, D., Wei, Q., et al. (2022) Three OsMYB36 Members Redundantly Regulate Casparian Strip Formation at the Root Endodermis. The Plant Cell, 34, 2948-2968.
https://doi.org/10.1093/plcell/koac140
[32] Shukla, V., Han, J., Cléard, F., Lefebvre-Legendre, L., Gully, K., Flis, P., et al. (2021) Suberin Plasticity to Developmental and Exogenous Cues Is Regulated by a Set of MYB Transcription Factors. Proceedings of the National Academy of Sciences, 118, Article No. 118.
https://doi.org/10.1073/pnas.2101730118
[33] Hématy, K., De Bellis, D., Wang, X., Mähönen, A.P. and Geldner, N. (2022) Analysis of Exocyst Function in Endodermis Reveals Its Widespread Contribution and Specificity of Action. Plant Physiology, 189, 557-566.
https://doi.org/10.1093/plphys/kiac019
[34] 王志刚. OsCASP1在水稻根内皮层凯氏带形成和矿质元素吸收中的作用机理[D]: [博士学位论文]. 南宁: 广西大学, 2022.
[35] Chen, T., Cai, X., Wu, X., Karahara, I., Schreiber, L. and Lin, J. (2011) Casparian Strip Development and Its Potential Function in Salt Tolerance. Plant Signaling & Behavior, 6, 1499-1502.
https://doi.org/10.4161/psb.6.10.17054
[36] 翁群清. 凯氏带蛋白OsCASP4-2在水稻凯氏带形成及非生物胁迫中的功能研究[D]: [硕士学位论文]. 福州: 福建农林大学, 2018.
[37] 刘鑫. 根系质外体屏障在老芒麦适应干旱胁迫中的作用[D]: [硕士学位论文]. 成都: 西南民族大学, 2022.
[38] Moore, C.A., Bowen, H.C., Scrase‐Field, S., Knight, M.R. and White, P.J. (2002) The Deposition of Suberin Lamellae Determines the Magnitude of Cytosolic Ca2+ Elevations in Root Endodermal Cells Subjected to Cooling. The Plant Journal, 30, 457-465.
https://doi.org/10.1046/j.1365-313x.2002.01306.x
[39] Beisson, F., Li, Y., Bonaventure, G., Pollard, M. and Ohlrogge, J.B. (2007) The Acyltransferase GPAT5 Is Required for the Synthesis of Suberin in Seed Coat and Root of Arabidopsis. The Plant Cell, 19, 351-368.
https://doi.org/10.1105/tpc.106.048033
[40] Yadav, V., Molina, I., Ranathunge, K., Castillo, I.Q., Rothstein, S.J. and Reed, J.W. (2014) ABCG Transporters Are Required for Suberin and Pollen Wall Extracellular Barriers in Arabidopsis. The Plant Cell, 26, 3569-3588.
https://doi.org/10.1105/tpc.114.129049
[41] Ranathunge, K. and Schreiber, L. (2011) Water and Solute Permeabilities of Arabidopsis Roots in Relation to the Amount and Composition of Aliphatic Suberin. Journal of Experimental Botany, 62, 1961-1974.
https://doi.org/10.1093/jxb/erq389
[42] Barberon, M., Vermeer, J.E.M., De Bellis, D., Wang, P., Naseer, S., Andersen, T.G., et al. (2016) Adaptation of Root Function by Nutrient-Induced Plasticity of Endodermal Differentiation. Cell, 164, 447-459.
https://doi.org/10.1016/j.cell.2015.12.021
[43] 孟利芳. 凯氏带和木栓层在宁夏枸杞根系Na+吸收中的作用研究[D]: [硕士学位论文]. 银川: 宁夏大学, 2024.
[44] 朱进, 彭玉全, 沈若妍, 等. 淹水胁迫对小白菜幼苗根系生长、生理及凯氏带的影响[J]. 植物生理学报, 2019, 55(8): 1089-1097.
[45] Krishnamurthy, P., Ranathunge, K., Franke, R., Prakash, H.S., Schreiber, L. and Mathew, M.K. (2009) The Role of Root Apoplastic Transport Barriers in Salt Tolerance of Rice (Oryza sativa L.). Planta, 230, 119-134.
https://doi.org/10.1007/s00425-009-0930-6
[46] Shen, D., Micic, N., Venado, R.E., Bjarnholt, N., Crocoll, C., Persson, D.P., et al. (2025) Apoplastic Barriers Are Essential for Nodule Formation and Nitrogen Fixation in Lotus japonicus. Science, 387, 1281-1286.
https://doi.org/10.1126/science.ado8680
[47] Ranathunge, K., Thomas, R.H., Fang, X., Peterson, C.A., Gijzen, M. and Bernards, M.A. (2008) Soybean Root Suberin and Partial Resistance to Root Rot Caused by Phytophthora sojae. Phytopathology®, 98, 1179-1189.
https://doi.org/10.1094/phyto-98-11-1179
[48] Mahiwal, S., Andersen, T.G. and Shen, D. (2025) Establishment and Functions of the Casparian Strip. Molecular Plant, 18, 1249-1252.
https://doi.org/10.1016/j.molp.2025.07.001
[49] Ramakrishna, P. and Somssich, M. (2022) Exocyst Function and Specificity during Casparian Strip Formation—Insights via a Gene-Edited Endodermis. Plant Physiology, 189, 435-437.
https://doi.org/10.1093/plphys/kiac101