Nikolov Lachezar A, Tomlinson P B, Manickam Sugumaran, Endress Peter K, Kramer Elena M, Davis Charles C
Department of Organismic and Evolutionary Biology, Harvard University Herbaria, Cambridge, MA 02138, USA.
The Kampong, National Tropical Botanical Garden, 4013 Douglas Road, Miami, FL 33133, USA.
Ann Bot. 2014 Aug;114(2):233-42. doi: 10.1093/aob/mcu114. Epub 2014 Jun 18.
Species in the holoparasitic plant family Rafflesiaceae exhibit one of the most highly modified vegetative bodies in flowering plants. Apart from the flower shoot and associated bracts, the parasite is a mycelium-like endophyte living inside their grapevine hosts. This study provides a comprehensive treatment of the endophytic vegetative body for all three genera of Rafflesiaceae (Rafflesia, Rhizanthes and Sapria), and reports on the cytology and development of the endophyte, including its structural connection to the host, shedding light on the poorly understood nature of this symbiosis.
Serial sectioning and staining with non-specific dyes, periodic-Schiff's reagent and aniline blue were employed in order to characterize the structure of the endophyte across a phylogenetically diverse sampling.
A previously identified difference in the nuclear size between Rafflesiaceae endophytes and their hosts was used to investigate the morphology and development of the endophytic body. The endophytes generally comprise uniseriate filaments oriented radially within the host root. The emergence of the parasite from the host during floral development is arrested in some cases by an apparent host response, but otherwise vegetative growth does not appear to elicit suppression by the host.
Rafflesiaceae produce greatly reduced and modified vegetative bodies even when compared with the other holoparasitic angiosperms once grouped with Rafflesiaceae, which possess some vegetative differentiation. Based on previous studies of seeds together with these findings, it is concluded that the endophyte probably develops directly from a proembryo, and not from an embryo proper. Similarly, the flowering shoot arises directly from the undifferentiated endophyte. These filaments produce a protocorm in which a shoot apex originates endogenously by formation of a secondary morphological surface. This degree of modification to the vegetative body is exceptional within angiosperms and warrants additional investigation. Furthermore, the study highlights a mechanical isolation mechanism by which the host may defend itself from the parasite.
全寄生植物科大花草科的物种展现出开花植物中最为高度特化的营养体之一。除了花茎及相关苞片外,该寄生植物是一种生活在葡萄科寄主植物体内的类菌丝内生菌。本研究全面阐述了大花草科三个属(大花草属、寄生花属和蛇菰属)的内生营养体,并报告了内生菌的细胞学及发育情况,包括其与寄主的结构连接,从而揭示了这种共生关系中鲜为人知的本质。
采用连续切片并用非特异性染料、过碘酸希夫试剂和苯胺蓝染色,以便在系统发育多样的样本中表征内生菌的结构。
利用先前确定的大花草科内生菌与其寄主之间核大小的差异,来研究内生体的形态和发育。内生菌通常由在寄主根内呈放射状排列的单列丝状体组成。在某些情况下,花期发育期间寄主将寄生虫从体内排出的过程会因寄主的明显反应而受阻,但除此之外,营养生长似乎并未受到寄主的抑制。
即使与曾经归为大花草科的其他具有一定营养分化的全寄生被子植物相比,大花草科产生的营养体也大幅简化和特化。基于先前对种子的研究以及这些发现,得出的结论是内生菌可能直接由原胚发育而来,而非由成熟胚发育而来。同样,花茎直接源自未分化的内生菌。这些丝状体产生一个原球茎,其中茎尖通过形成次生形态表面而内生起源。营养体这种程度的特化在被子植物中是异常的,值得进一步研究。此外,该研究突出了一种寄主可能用以抵御寄生虫的机械隔离机制。