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林业科学 ›› 2022, Vol. 58 ›› Issue (2): 196-205.doi: 10.11707/j.1001-7488.20220219

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基于具缘纹孔膜特征的木质部栓塞机制研究进展

李姗1,2,3,4,李玉军5,张亚6,王杰1,2,3,万贤崇7,姜笑梅1,2,3,孙熙佑4,朱新京4,殷亚方1,2,3,*   

  1. 1. 中国林业科学研究院木材工业研究所 北京 100091
    2. 中国林业科学研究院木材标本馆 北京 100091
    3. 国家林业和草原局木材科学与技术重点实验室 北京 100091
    4. 陕西科技大学环境科学与工程学院 西安 710021
    5. 西北工业大学机电学院 西安 710072
    6. 安徽师范大学生命科学学院 芜湖 241000
    7. 中国林业科学研究院林业新技术研究所 北京 100091
  • 收稿日期:2021-02-18 出版日期:2022-02-25 发布日期:2022-04-26
  • 通讯作者: 殷亚方
  • 基金资助:
    国家自然科学基金青年项目(32001291);北京市自然科学基金青年项目(61840048);“十三五”国家重点研发计划课题(2017YFD0600202)

Research Progress in Mechanism of Xylem Embolism Based on Characteristics of Bordered Pit Membrane

Shan Li1,2,3,4,Yujun Li5,Ya Zhang6,Jie Wang1,2,3,Xianchong Wan7,Xiaomei Jiang1,2,3,Xiyou Sun4,Xinjing Zhu4,Yafang Yin1,2,3,*   

  1. 1. Research Institute of Wood Industry, CAF Beijing 100091
    2. Wood Collections (WOODPEDIA), CAF Beijing 100091
    3. Key Laboratory of Wood Science and Technology, National Forestry and Grassland Administration Beijing 100091
    4. School of Environmental Science and Engineering, Shaanxi University of Science and Technology Xi'an 710021
    5. School of Mechanical Engineering, Northwestern Polytechnical University Xi'an 710072
    6. School of Life Sciences, Anhui Normal University Wuhu 241000
    7. Research Institute of Forestry New Technology, CAF Beijing 100091
  • Received:2021-02-18 Online:2022-02-25 Published:2022-04-26
  • Contact: Yafang Yin

摘要:

木质部栓塞指外界气泡进入木质部充水管道导致的植物水分传导功能受阻。木质部输水管道间具缘纹孔膜是木质部栓塞形成、扩散和疲劳产生的关键部位,近年来,具缘纹孔膜特征在木质部栓塞机制研究中日益受到关注。为更全面和清晰理解木质部栓塞形成、扩散和疲劳过程中具缘纹孔膜的作用,本研究在整理分析国内外木质部栓塞相关研究的基础上,首先探讨被子植物、裸子植物木质部栓塞的形成机制和扩散规律,阐释栓塞形成和扩散的"气种假说",明确具缘纹孔膜的核心作用; 然后分别针对具缘纹孔膜的构造、化学、物理和微力学性质与微液流行为等主要特征以及不同特征间的相互关系,归纳具缘纹孔膜在木质部栓塞形成、扩散和疲劳等不同阶段的重要作用,概述三维结构模型构建、化学成分原位检测、力学行为模拟和单个纹孔膜微液流量化分析等具缘纹孔膜特征关键表征方法的发展现状和问题; 最后提出木质部栓塞应优先开展的3方面研究: 1)不同水分条件下具缘纹孔膜化学成分、物理和力学性质的变化; 2)气泡在水分与具缘纹孔膜三维孔隙结构间水-固耦合界面的形成与扩散方式; 3)不同水分条件下具缘纹孔膜结构变化及其对微液流效率的影响。同时,亟待突破具缘纹孔膜三维空间结构的原位精准表征与生物模型构建及优化的技术瓶颈。通过进一步研究具缘纹孔膜构造、化学、物理、力学性质和微液流行为,揭示具缘纹孔膜特征对木质部栓塞的影响规律这一关键科学问题,将有助于深入理解木质部栓塞机制,为探究木质部抗旱机制、选育优良耐旱种质资源提供科学依据。

关键词: 木质部栓塞, 具缘纹孔膜, 构造, 性质, 微液流行为

Abstract:

Xylem embolism refers to the interrupt of plant water transport function caused by external air bubbles entering the xylem water-filled conduits. Bordered pit membrane is the crucial location where xylem embolism, spread and fatigue occur. In recent years, the characteristics of bordered pit membrane have received increasing attention in the study of xylem embolism mechanisms. In order to have more comprehensive and clear understanding of the role of bordered pit membrane in the formation, spread and fatigue of xylem embolism, we firstly discussed the formation mechanism and diffusion pattern of xylem embolism in angiosperms and gymnosperms, based on relevant research on xylem embolism at home and abroad. We elucidated the "air seeding hypothesis" in xylem embolism formation and diffusion, and clarified the crucial role of the bordered pit membrane; we also reviewed the main characteristics of the structural, chemical, physical and micromechanical properties of the bordered pit membrane, as well as the interrelationships between the different characteristics. Then we summarized the important role of bordered pit membranes in different stages of xylem embolism, diffusion and fatigue. We also summarized the recent advances and problems of the key characterization method of bordered pit membrane, such as the construction of three-dimensional structure model, in-situ detection of chemical composition, and mechanical behavior of the bordered pit membrane, microflow quantification and analysis of the single pit membrane. Finally, we proposed three research topics which should be given priority in xylem embolism: 1) changes in the chemical composition, physical and mechanical properties of the bordered membrane under different moisture conditions; 2) the formation and diffusion of air bubbles at the water-solid interface between water and the three-dimensional porous structure of the bordered pit membrane; 3) structural changes of the bordered pit membrane under different moisture conditions and their effects on the efficiency of microflow. Meanwhile, it is urgent to break through the technical bottleneck of the accurate in-situ characterization of the three-dimensional structure of the bordered pit membrane, and the construction and optimization of biological models. The further investigation of the structural, chemical, physical, mechanical properties and microflow behavior of the bordered pit membrane to reveal the influences of the characteristics of the bordered pit membrane on xylem embolism would be helpful to understand the mechanisms of xylem embolism, to provide scientific basis for exploring the drought-resistant mechanisms of xylem and breeding excellent drought-tolerant germplasm resources.

Key words: xylem embolism, bordered pit membrane, structure, property, microflow behavior

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