Abstract Bryophytes are close extant relatives of the ancestral land plant. As such, they have retained many innovations that had enabled the adaptation of early land plants to the terrestrial environment. One of such important innovations is the elaboration of an enormously diverse array of secondary metabolites. This article reviews current knowledge of terpenoid secondary metabolites, which constitute the largest family of plant metabolites, in bryophytes from three perspectives: chemical diversity, biosynthesis, and biological functions. The diversity of terpenoids in bryophytes, particularly in liverworts, is enormously rich. More than 1600 terpenoids have been reported from this plant group. While many terpenoids are observed in both bryophytes and seed plants, some are unique to bryophytes. It is just the beginning for us to understand the molecular and biochemical basis underlying terpenoid biosynthesis in bryophytes. Compared to seed plants, which have only one type of terpene synthase genes, so-called typical plant terpene synthase genes, bryophytes employ not only typical plant terpene synthase genes but also another class of terpene synthase genes called microbial-terpene synthase-like (MTPSL) genes for terpenoid biosynthesis. Biochemical studies suggest the MTPSLs are largely responsible for the terpenoid diversity in bryophytes, particularly sesquiterpenes and monoterpenes. Existing literature indicates that terpenoids made by bryophytes have important functions in diverse biological and ecological processes, particularly as defenses against biotic and abiotic stresses. The continued development of genomic resources and molecular tool kit for bryophytes will accelerate characterization of terpenoids biosynthesis and their biological functions in this important lineage of plants.

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苔藓植物中萜类次级代谢物：化学多样性、生物合成和生物功能

摘要 苔藓植物是现存陆地植物祖先的近亲。因此，他们保留了许多使早期陆生植物适应陆地环境的创新。此类重要创新之一是精心设计了种类繁多的次生代谢物。本文从化学多样性、生物合成和生物学功能三个角度回顾了苔藓植物中萜类次生代谢物的最新知识，萜类次生代谢物构成了苔藓植物中最大的植物代谢物家族。苔藓植物中萜类化合物的多样性非常丰富，尤其是苔类植物。据报道，该植物群中含有 1600 多种萜类化合物。虽然在苔藓植物和种子植物中都观察到了许多萜类化合物，但有些是苔藓植物独有的。这只是我们了解苔藓植物萜类生物合成的分子和生化基础的开始。与种子植物只有一种萜烯合酶基因，即所谓的典型植物萜烯合酶基因相比，苔藓植物不仅利用了典型的植物萜烯合酶基因，还利用了另一类萜烯合酶基因，称为微生物萜类合酶基因（MTPSL）。 ) 萜类生物合成基因。生化研究表明，MTPSL 是苔藓植物中萜类多样性的主要原因，尤其是倍半萜和单萜。现有文献表明，苔藓植物产生的萜类化合物在多种生物和生态过程中具有重要功能，特别是作为抵御生物和非生物胁迫的防御。