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    绿色裸藻类系统发育研究进展

    RESEARCH ADVANCES ON THE PHYLOGENY OF GREEN EUGLENOIDS

    • 摘要: 绿色裸藻类(Green euglenoids)是兼具动物与植物双重性状的特殊类群, 也是研究真核生物次级内共生起源的经典模式生物, 兼具重要基础研究价值与应用价值。该类群常发现于富营养化水体中, 其对水中高浓度的氮、磷养分具有很强的耐受性, 还是高效的重金属超积累微生物, 对镉、锌、汞、铬、铅等有毒重金属具有一定的吸收能力。同时因其高蛋白、高营养特性被广泛应用于食品、饲料及肥料等领域。形态多样性与系统发育关系的矛盾是该类群分类学的核心科学问题。本文梳理了基于形态特征构建绿色裸藻类传统分类体系的历程, 综述了分子系统学技术对各属系统发育关系的重构, 包括叶绿体基因组、线粒体基因组、染色体水平全基因组等不同层次组学研究对解决分类争议、完善分类体系、明确绿色裸藻类质体起源的核心贡献, 重点厘清其分类体系的动态变化。相较于国际研究进展, 我国绿色裸藻类系统分类研究起步较晚, 虽已报道部分新种及中国新记录种, 但样本覆盖度与分子数据积累仍存在不足。未来研究应强化形态与分子证据的整合, 扩大样本采集范围, 推动我国绿色裸藻类多样性与演化关系的深入探索。

       

      Abstract: Green euglenoids are a freshwater group with photosynthetic autotrophic ability in the phylum Euglenophyta, with over 1000species reported globally. They possess both ecological indicator value (as typical indicator organisms for eutrophic water bodies) and economic value (with high-protein and high-nutrition characteristics, and are widely used in food, feed, and fertilizers). Before the 20th century, their classification mainly relied on classic morphological indicators. However, the morphological similarity of single-celled algae and the unique euglenoid movement made species identification difficult, leading to frequent duplicate reports of the same species and considerable confusion in the classification system. Since the 21st century, phylogenetic studies based on molecular markers have promoted the reconstruction of the classification system. Molecular evidence clearly shows that green euglenoids form a monophyletic group. Due to the polyphyly issue, the genus Euglena has been split into new taxonomic units, whereas the genera Phacus and Lepocinclis have been established as the family Phacaceae. However, the introduction of molecular data has also ledto a disconnection between morphology and phylogenetic results. For important groups such as Trachelomonas and Strombomonas, although molecular studies support their independence, data coverage remains extremely low (molecular data exist for only 25species of Trachelomonas and 10species). The lack of morphological transitional species further hinders the analysis of phylogenetic relationships. The morphological definition of Lepocinclis has become blurred due to the transfer of Euglena species, and the iconic characteristic of cell flattening in Phacus has been broken due to the addition of non-flattened species. Recent breakthroughs in omics technologies have filled long-standing data gaps across multiple key evolutionary lineages of this clade. Accumulated evidence from existing studies has confirmed that green euglenoid chloroplasts trace their origin to a single secondary endosymbiotic event involving green algae, resolved the evolutionary trajectories of organelle genomes within this clade, and provided whole-genome level validation for the shopping bag hypothesis, which posits multiple independent plastid acquisitions by the ancestor of green euglenoids. Furthermore, the first high-quality, chromosome-level de novo genome assembly of euglenoids has recently been completed, breaking the long-standing technical bottleneck that has historically hampered progress in this field. Collectively, these advances establish an unprecedented molecular foundation for the taxonomic delimitation and evolutionary history reconstruction of green euglenoids. Research on green euglenoids in China began with the first report by Wang Jiaji in 1925. In 1999, the Flora Algarum Sinicarum Aquae Dulcis systematically recorded 9 genera and 318species. In recent years, one new species of Trachelomonas and 9 newly recorded species of Euglena in China have been reported by combining morphological and molecular methods, providing basic data for biogeographical research. Future research should integrate morphological and molecular biological methods, establish a classification standard that takes into account both morphological distinguishability and genetic consistency, expand the sampling scope to improve the representativeness of phylogenetic analysis, and leverage the advantages of China's freshwater ecosystem to support further studies. The unique mesokaryotic structure, three-membrane chloroplasts, and non-simplified genome characteristics of green euglenoids serve as a key model for the study of eukaryotic evolution.

       

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