A Review of the Interaction Mechanism and Law between Vegetation and Rock Geochemical Background in Karst Areas
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摘要:
岩溶地貌主要是由碳酸盐溶解形成的特殊景观，岩溶区植被的生长发育受到基岩的制约，并演化出各种机制来适应岩溶区的独特环境。本文综述了岩溶地区植物对岩溶环境适应机制及植物生长对碳酸盐岩风化的驱动作用。通过总结发现：①植物主要通过分泌碳酸酐酶等有机物促进矿物分解、生物钻孔作用改善岩石表面的持水性能、根劈作用加速破碎岩石的崩解等化学生物和物理作用，促进了碳酸盐岩的风化溶解，形成独特的岩溶地球化学背景。②岩溶区植物通过调整自身结构和生理功能来适应干旱、高钙和营养元素缺乏等逆境。植物的抗旱性主要通过生理生化过程、形态结构和水分的利用方式来适应干旱或缺水环境，不同的植物进化出不同水分利用方式，提高水分利用效率，减少蒸腾；植物的高钙适应性是通过生理结构和生理过程来实现的，在高钙环境下的优势植物可通过形成钙化根、草酸钙含晶细胞和叶片调节等方式保持植株钙含量处于相对稳定的状态，并且植物还可以通过调节体内钙库和控制钙的吸收转运来控制细胞内钙离子浓度；根系分泌的有机酸和菌根能帮助植被在土壤中获取营养元素，以应对土壤的营养元素缺乏。③岩溶植被在正向演替过程中，土壤保水保肥能力增强、稳定性增加，物种的生存几率增加，物种多样性也随着增加。植被演化出的适应机制影响了植物的分布和生长，推进植物群落的演替过程和促进植物多样性的形成。但是多样性与群落生态系统的稳定性间的内在关系，以及与非岩溶区的对比特征仍需要开展深入的研究和探索。
Abstract:Karst landforms are mainly special landscapes formed by carbonate dissolution.They are characterized by calcium abundance, lack of soil resources, and insufficient water resources. The growth and development of vegetation in the karst area is restricted by the bedrock. It is very important to understand the synergistic interaction between vegetation metabolism and geochemistry of carbonate rocks in karst areas to maintain the stability of structure and function of the karst ecosystem.
The mechanism and law of interaction between vegetation and rock geochemical background in karst areas from two aspects is expounded in this paper: vegetation community promotes weathering of carbonate rocks and geochemical background restricts vegetation.
Through summarizing: (1) Plants promote the weathering and dissolution of carbonate rocks through physicochemical and biological actions, such as secreting carbonic anhydrase organic matter, improving the water retention performance of rock surface through boring by organisms and accelerating the disintegration of broken rocks through root splitting, thus forming a unique karst geochemical background of drought, high calcium, shallow soil layer and lack of nutrients in the soil layer.
(2) With the long-term interaction between plants and the karst environment, plants adapt to environmental stress by adjusting their own structures and physiological functions, and even their unique plant succession rules. The plants that survived eventually evolved into unique karst plants that were drought-resistant, adaptable to high-calcium environments, and able to cope with nutrient deficiencies.
Due to the solubility of carbonate rocks, the hydrologic system forms a two-layer spatial structure of surface and underground, which makes it difficult to utilize groundwater resources. As a result, the available water resources of local plants are limited, and are prone to drought stress. The drought resistance of plants adapts to the drought or water shortage environment mainly through physiological and biochemical processes, morphological structure and water use. In the morphological structure of the plant, through the stomatal regulation and the xeric structural characteristics of the leaves, the transpiration water loss of the plant is minimized. Some karst plants can cope with drought stress through physiological and biochemical processes, which can reduce the damage caused by drought stress by increasing the activities of antioxidant enzymes and accumulating osmoregulatory substances through phytochemicals. Karst plants improve water use efficiency and reduce transpiration through different water use methods in the dry season. For example, some plants absorb deep soil, deep bedrock water or groundwater water through developed deep roots, and some plants even use fog water.
The adaptability of plants to high calcium is realized through physiological structure and process. In a high calcium environment, karst plants can limit the excess calcium transfer upward by forming calcified roots and keeping the calcium content in plants in a relatively stable state through the regulation of calcium oxalate crystal cells and leaves. Plants can also control the intracellular calcium ion concentration by regulating the calcium pool in vivo and controlling the absorption and transport of calcium.
Organic acids and mycorrhiza secreted by roots can help vegetation obtain nutrients in the soil to cope with nutrient deficiency in the soil. The organic acid content secreted by the roots of karst plants is usually higher than that of non-karst plants, and the increase of organic acid content can help plants to absorb trace elements. Arbuscular mycorrhizal fungi (AM) and ectomycorrhizal fungi (ECM) help plants adapt to nutrient deficiency by absorbing mineral nutrients from the soil.
(3) The vegetation succession in the karst area is similar to the general stage of vegetation succession in normal landforms, but the vegetation succession in the karst area has the particularity of self-generation. The vegetation succession from the middle to the top of the mountain takes a longer time. Vegetation succession changed physical and chemical properties and soil quality. In the process of succession, soil bulk density gradually decreases, and porosity gradually increases over time. At the same time, soil nutrients accumulate with the positive succession of vegetation. During the positive succession process of karst vegetation, the soil water and fertilizer retention capacity is enhanced, the stability is increased, and the survival probability of species and the species diversity are also increased. The special rock geochemical background in the karst area leads to the obvious changes in the spatial distribution of soil resources. High soil heterogeneity promotes the formation of plant community species diversity. For example, the tropical karst area in Xishuangbanna, Yunnan Province covers an area of 3600km2, accounting for 19% of the land area. The forest survey results in the karst area show that there are 153 families, 640 genera and 1394 species of vascular plants, accounting for 77.7% of the total floristic families, 56.1% of the genera and 37.9% of the species, respectively.
The karst plant community is the result of the long interaction between the plant and the environment, the continuous adaptation to the environment and the growth and reproduction. The karstification of plants, the adaptation mechanism of plants under the typical karst soil environment such as drought, high calcium stress and lack of nutrient elements, as well as the succession characteristics of karst vegetation shown in the adaptation process and the biodiversity of vegetation are a whole interaction. The site growth of vegetation promotes the weathering of carbonate rocks and the formation of soil, creating conditions for their own growth. Meanwhile, soil area and soil thickness are positively correlated with plant diversity. The succession of vegetation changes the physical and chemical properties of soil and improves the quality of soil, and the karstification of carbonate rocks promotes the growth of vegetation. The adaptive mechanism of vegetation affects the distribution and growth of plants and promotes the succession process of plant communities and the formation of plant diversity. Due to the spatial heterogeneity of the karst environment and the diversity of plant habitats, plant diversity in karst areas is manifested as few genera, few species and endemic species. However, the internal relationship between plant diversity and the stability of the community ecosystem, as well as the comparative characteristics with non-karst areas still need to be further studied and explored.
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Key words:
- carbonate rock /
- vegetation /
- plant adaptation strategy /
- biodiversity /
- succession
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