克隆植物对异质生境的适应对策研究进展

生境异质性是自然生态系统的基本特征，植物生长的必需资源和环境胁迫因子均存在着复杂的时间和空间异质性。克隆植物是指在自然条件下具有克隆特性的植物，即可通过与母株相连的芽、根茎、分蘖或枝条等繁殖体产生无性繁殖的植物，这些繁殖体一旦定居便可成为潜在的独立个体。克隆植物具有独特的生境适应策略(如形态可塑性、克隆整合、克隆分工、觅食行为、风险分摊等),面对异质性的生境条件，它可以通过调整自身的生理和形态结构来适应异质生境。目前，对于克隆植物在异质生境适应行为的研究已有很多报道，然而系统性的归纳和总结尚有欠缺。综述了克隆植物在不同资源异质生境(光照、养分、水分)和不同胁迫生境(盐碱胁迫、风沙胁迫、重金属胁迫)下独特的适应对策。最后，针对克隆植物对异质生境的适应对策，进行了总结并对未来的重点研究方向提出建议：(1)时间异质性尺度上的考量；(2)异质性生境中生物因子的调控作用；(3)克隆植物入侵机制；(4)克隆植物在生态修复中的应用潜力。

Reviewing the adaptation strategies of clonal plants to heterogeneous habitats

Habitat heterogeneity is a basic characteristic of natural ecosystem. The essential resources and environmental factors for plant growth are highly spatiotemporal heterogeneity. Clonal plants are plants with the clonal characteristics under natural condition, which can produce asexual reproduction through buds, rhizomes, tillers or branches connected with the mother plant. Once these propagules settle down, they can become potential independent individuals. They can adapt to the heterogeneous habitats through their unique habitat adaptation strategies (including morphological plasticity, clonal integration, clonal division of labor, foraging behavior, etc.) to obtain resources from heterogeneous habitats or avoid stress to the greatest extent. Based on analysis of the different characteristics of habitat heterogeneity in different classification criteria, this paper holds that habitat heterogeneity can be divided into temporal heterogeneity and spatial heterogeneity, or resource heterogeneity and environmental heterogeneity in terms of habitat function. Meanwhile, the adaptive strategies of the clonal plants in different resource heterogeneous habitats (light, nutrient, and water) or heterogeneous stress conditions (saline alkali stress, sand burial and heavy metal stress) were reviewed. Finally, the adaptation strategies of clonal plants to heterogeneous habitats were summarized. The suggestions for future research directions were put forward. (1) In the future research, we should consider combining the variation characteristics of habitat factors with spatial heterogeneity to explore the response characteristics of clonal plants more comprehensively. It is necessary to analyze the characteristics of clonal plants to habitat heterogeneity in different growth stages from the perspective of field in-situ control and long-term monitoring. (2) The previous studies have focused on the responses of clonal plants to abiotic heterogeneity (such as nutrients). However, soil biological factors (soil animals and soil microorganisms) also play an important role in regulation, and may even reshape the response of clonal plants in nature. (3) The invasion mechanism of clonal plants should also be deeply researched. (4) The application potential of clonal plants in ecosystem restoration should be widely concerned in future research.