植物生态学报 ›› 2024, Vol. 48 ›› Issue (3): 269-286.DOI: 10.17521/cjpe.2023.0107 cstr: 32100.14.cjpe.2023.0107
• 综述 • 下一篇
收稿日期:
2023-04-19
接受日期:
2023-08-03
出版日期:
2024-03-20
发布日期:
2024-04-24
通讯作者:
*(liuyh@bjfu.edu.cn)
基金资助:
ZHANG Yu-Jian, LIU Yan-Hong*()
Received:
2023-04-19
Accepted:
2023-08-03
Online:
2024-03-20
Published:
2024-04-24
Contact:
*(liuyh@bjfu.edu.cn)
Supported by:
摘要:
随着全球逐渐暖干化, 林火不仅驱动着森林生态系统结构和功能的变化, 同时也影响树木的生理和生长。林火导致的热损伤引发树木一系列复杂生理响应。揭示火后树木生理的响应机制, 对于进一步理解树木碳水关系和火后恢复生长限制, 以及提高火后树木死亡预测准确性具有重要指导意义。该文从林火对树木的作用途径和方式着手, 基于不同形式林火(树冠火、地表火、地下火)对树木各部分(树冠、树干、根系)造成的损伤, 综述了林火对树木生理的直接影响和间接影响, 以及火后树木生理与非生物和生物因素的互作关系。热损伤诱导的形成层、韧皮部坏死和木质部水力失衡是火后树木生理的主要响应机制, 二者导致的两个生理功能限制——“碳饥饿”和水力失效——严重影响树木的碳水关系, 也决定了火后树木是恢复生长还是延迟死亡。火后树木生理机制还与干旱、昆虫攻击和微生物入侵等其他因素密切相关。该文强调了对林火强度的定量分析和对植物组织死亡阈值的准确判断的迫切性, 同时提出了探究火后树木生理与功能性状和其他因素的互作关系的必要性。精确评估树木生理机制间关系对于深入理解林火如何影响树木功能完整性极为关键, 有助于完善林火风险评估和树木死亡模型预测。在未来气候暖干化驱动的高频、高强度林火发生背景下, 对树木生理响应的深刻认识对于更好地研究火后生态系统动态及其与气候因子的相互关系同样具有重要意义。
张雨鉴, 刘艳红. 林火干扰下的树木生理及主要影响因素. 植物生态学报, 2024, 48(3): 269-286. DOI: 10.17521/cjpe.2023.0107
ZHANG Yu-Jian, LIU Yan-Hong. Tree physiology and major influencing factors under forest fires. Chinese Journal of Plant Ecology, 2024, 48(3): 269-286. DOI: 10.17521/cjpe.2023.0107
图1 林火进程。不同形式的林火(树冠火、地表火和地下火)通过热传递(传导、对流和辐射)影响树木(A), 造成不同组织或器官的损伤和坏死(B)。根据烧伤程度, 树冠火的影响结果分为树冠烧伤、树冠烧死和树冠烧毁(C)。D为森林可燃物热解过程, 燃烧是将挥发性碳氢可燃物和氧气转化为各种产物(如CO2、CO、H2O及焦油、烟灰等)的氧化放热反应, 燃烧方向从左到右。
Fig. 1 Forest fire process. Different classes of forest fires (e.g., crown, surface, and ground fires) affect trees by heat transfer (e.g., conduction, convection, and radiation) (A), causing injury and necrosis of various tissues or organs (B). According to the degree of scorching, the effects of crown fire can be divided into crown scorch, crown kill, and crown consumption (C). Schematic diagram (D) of pyrolysis for forest fuel elements. Combustion is an exothermic oxidation reaction that converts volatile hydrocarbon fuels and oxygen into various products (CO2, CO, H2O, tar, soot, etc.), with a combustion direction from left to right.
图2 不同研究中松属(Pinus)树种死亡率预测与树冠烧伤体积的关系。Prescribed fire、Side wildfire和Bridger-Knoll wildfire分别表示不同的火事件。
Fig. 2 Probability of mortality as a function of crown volume scorched for Pinus species in different studies. Prescribed fire, Side wildfire and Bridger-Knoll wildfire indicate different fire events, respectively.
图3 树冠、树干和根系对火后损伤的一系列潜在生理响应过程。标有“火”符号的方框表示热传递对组织和器官造成的直接损伤(直接影响), 未标记的方框表示火后的一系列生理机制(间接影响)。红色方框表示与形成层和韧皮部坏死相关的过程; 蓝色方框表示与木质部损伤相关的过程; 绿色方框表示火后的生物(昆虫和微生物)攻击可能会加剧热损伤对生理机制的影响。标有“钥匙”符号的方框表示连接树木内在碳水关系的两个关键生理过程, 分别为“碳饥饿”和水力效率降低, 严重的水力效率降低会导致水力失效。两个生理过程与火后环境密切相关, 能够反映树木的生长限制, 决定树木火后是恢复生长还是延迟死亡。
Fig. 3 Conceptual diagram of cascading potential physiological responses of postfire injuries in tree crowns, trunks, and roots. Boxes marked with the “fire” symbol indicate direct damage (direct effect) caused by heat transfer to tissues and organs, while unmarked boxes indicate a series of physiological mechanisms (indirect effect) after a fire. Red-filled boxes indicate processes related to cambium and phloem necrosis. Blue-filled boxes indicate processes related to damage in the xylem. Green-filled boxes indicate that biotic attacks (insects and microorganisms) after a fire may exacerbate the impact of thermal damage on physiological mechanisms. Boxes marked with the “key” symbol indicate two crucial physiological processes involving the carbon-water relationship within a tree, namely, “carbon starvation” and reduced hydraulic efficiency. Severe reduction in hydraulic efficiency leads to hydraulic failure. The two physiological processes are closely related to environmental conditions after a fire, which can demonstrate the growth limitations of trees and determine whether they can restore their growth or delay their death after a fire.
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