Nogués Isabel, Brilli Federico, Loreto Francesco
Consiglio Nazionale delle Ricerche, Istituto di Biologia Agroambientale e Forestale, 00016 Monterotondo Scalo, Rome, Italy.
Plant Physiol. 2006 Jun;141(2):721-30. doi: 10.1104/pp.105.073213. Epub 2006 Feb 3.
Dimethylallyl diphosphate (DMADP) and geranyl diphosphate (GDP) are the last precursors of isoprene and monoterpenes emitted by leaves, respectively. DMADP and GDP pools were measured in leaves of plants emitting isoprene (Populus alba), monoterpenes (Quercus ilex and Mentha piperita), or nonemitting isoprenoids (Prunus persica). Detectable pools were found in all plant species, but P. persica showed the lowest pool size, which indicates a limitation of the whole pathway leading to isoprenoid biosynthesis in nonemitting species. The pools of DMADP and GDP of nonemitting, isoprene-emitting, and monoterpene-emitting species were partially labeled (generally 40%-60% of total carbon-incorporated (13)C) within the same time by which volatile isoprenoids are fully labeled (15 min). This indicates the coexistence of two pools for both precursors, the rapidly labeled pool presumably occurring in chloroplasts and thereby synthesized by the methylerythritol phosphate pathway and the nonlabeled pool presumably located in the cytosol and synthesized by the mevalonic pathway. In M. piperita storing monoterpenes in specialized leaf structures, the GDP pool remained totally unlabeled, indicating either that monoterpenes are totally formed by the mevalonic pathway or that labeling occurs slowly in comparison to the large pool of stored monoterpenes in this plant. The pools of DMADP and GDP increased during the season (from May to July) but decreased when the leaf was darkened or exposed to very high temperature. In the dark, the pool of DMADP of the isoprene-emitting species decreased faster than the pool of GDP. However, after 6 h of darkness, both pools were depleted to about 10% of the pool size in illuminated leaves. This indicates that both the chloroplastic and the cytosolic pools of precursors are depleted in the dark. When comparing measurements over the season and at different temperatures, an inverse correlation was observed between isoprene emission by P. alba and the DMADP pool size and between monoterpene emission by Q. ilex and the GDP pool size. This suggests that the pool size does not limit the emission of isoprenoids. Rather, it indicates that the flux of volatile isoprenoids effectively controls the size of their pools of precursors.
二甲基烯丙基二磷酸(DMADP)和香叶基二磷酸(GDP)分别是叶片排放异戊二烯和单萜的最后前体物质。对排放异戊二烯的植物(银白杨)、排放单萜的植物(冬青栎和胡椒薄荷)或不排放类异戊二烯的植物(桃)的叶片中的DMADP和GDP库进行了测量。在所有植物物种中均检测到了可测库,但桃树的库大小最低,这表明在不排放类异戊二烯的物种中,通向类异戊二烯生物合成的整个途径存在限制。在挥发性类异戊二烯被完全标记(15分钟)的相同时间内,不排放、排放异戊二烯和排放单萜的物种的DMADP和GDP库被部分标记(通常占总碳掺入量(13C)的40%-60%)。这表明两种前体物质都存在两个库,快速标记的库可能存在于叶绿体中,因此是通过甲基赤藓糖醇磷酸途径合成的,而未标记的库可能位于细胞质中,是通过甲羟戊酸途径合成的。在将单萜储存在特殊叶片结构中的胡椒薄荷中,GDP库完全未被标记,这表明单萜要么完全由甲羟戊酸途径形成,要么与该植物中大量储存的单萜相比,标记发生得很慢。DMADP和GDP库在季节中(从5月到7月)增加,但当叶片遮光或暴露于非常高的温度时会减少。在黑暗中,排放异戊二烯的物种的DMADP库比GDP库减少得更快。然而,在黑暗6小时后,两个库都减少到光照叶片中库大小的约10%。这表明前体物质的叶绿体库和细胞质库在黑暗中都会耗尽。在比较整个季节和不同温度下的测量结果时,观察到银白杨的异戊二烯排放与DMADP库大小之间以及冬青栎的单萜排放与GDP库大小之间呈负相关。这表明库大小并不限制类异戊二烯的排放。相反,这表明挥发性类异戊二烯的通量有效地控制了它们前体库的大小。