The Oil Bodies of Liverworts: Unique and Important Organelles in Land Plants

Abstract Oil bodies of liverworts are intracellular organelles bounded by a single unit membrane containing lipophilic globules suspended in a proteinaceous matrix. They are a prominent and highly distinctive organelle uniquely found in liverworts. Although they have been widely used in taxonomy and chemosystematics, and many of their secondary metabolites are known to be bioactive and are considered as potential sources of medicines, their origin, development and function still remain poorly understood. Recently, biochemical studies have indicated that the isoprenoid biosynthetic pathways in liverworts are similar to those of the seed plants and that oil bodies of Marchantia polymorpha contain a protein complex immunologically related to plastid and cytosolic enzymes of isoprenoid synthesis. Cytoplasmic lipid droplets lacking a bounding membrane have recently been recognized as important dynamic organelles playing active roles in cell physiology. Structural proteins, covering the surface of the lipid droplets and preventing them coalescing during desiccation, have been found in seed plants and also in the moss Physcomitrella patens. However, whether liverwort oil bodies play a dynamic role in cell metabolism, in addition to their role as sites of essential oil accumulation and sequestration, has not been formally tested. In this review, we present current knowledge on the oil bodies of liverworts on their origin and development, their role in taxonomy, chemosystematics and potential pharmaceutical applications leading to their functional significance, and we also identify avenues for future studies on this important but long-overlooked organelle. Keywords: oil bodylipophilic globuleliverwortorganellelipid dropletland plantontogenyhomologyevolutionecologyfunction ACKNOWLEDGMENTS The authors thank Kaarina Pihakaski and Yoshinori Asakawa for kindly providing permissions to reproduce their previously published figures, and Sinikka Piippo, David Glenny and Neil Bell for their valuable comments. Financial support from the Academy of Finland to the first author (no. 114715) and the National Science Foundation of China to the third author (NSFC no. 30825004) is gratefully acknowledged.