Woodward Andrew W, Bartel Bonnie
Department of Biochemistry and Cell Biology, Rice University, 6100 Main Street, Houston, TX 77005, USA.
Ann Bot. 2005 Apr;95(5):707-35. doi: 10.1093/aob/mci083. Epub 2005 Mar 4.
The phytohormone auxin is critical for plant growth and orchestrates many developmental processes.
This review considers the complex array of mechanisms plants use to control auxin levels, the movement of auxin through the plant, the emerging view of auxin-signalling mechanisms, and several interactions between auxin and other phytohormones. Though many natural and synthetic compounds exhibit auxin-like activity in bioassays, indole-3-acetic acid (IAA) is recognized as the key auxin in most plants. IAA is synthesized both from tryptophan (Trp) using Trp-dependent pathways and from an indolic Trp precursor via Trp-independent pathways; none of these pathways is fully elucidated. Plants can also obtain IAA by beta-oxidation of indole-3-butyric acid (IBA), a second endogenous auxin, or by hydrolysing IAA conjugates, in which IAA is linked to amino acids, sugars or peptides. To permanently inactivate IAA, plants can employ conjugation and direct oxidation. Consistent with its definition as a hormone, IAA can be transported the length of the plant from the shoot to the root; this transport is necessary for normal development, and more localized transport is needed for tropic responses. Auxin signalling is mediated, at least in large part, by an SCFTIR1 E3 ubiquitin ligase complex that accelerates Aux/IAA repressor degradation in response to IAA, thereby altering gene expression. Two classes of auxin-induced genes encode negatively acting products (the Aux/IAA transcriptional repressors and GH3 family of IAA conjugating enzymes), suggesting that timely termination of the auxin signal is crucial. Auxin interaction with other hormone signals adds further challenges to understanding auxin response.
Nearly six decades after the structural elucidation of IAA, many aspects of auxin metabolism, transport and signalling are well established; however, more than a few fundamental questions and innumerable details remain unresolved.
植物激素生长素对植物生长至关重要,并调控着许多发育过程。
本综述探讨了植物用于控制生长素水平的一系列复杂机制、生长素在植物体内的运输、生长素信号传导机制的新观点,以及生长素与其他植物激素之间的几种相互作用。尽管许多天然和合成化合物在生物测定中表现出生长素样活性,但吲哚 - 3 - 乙酸(IAA)被认为是大多数植物中的关键生长素。IAA可通过依赖色氨酸(Trp)的途径由色氨酸合成,也可通过不依赖Trp的途径由吲哚类Trp前体合成;这些途径均未完全阐明。植物还可通过第二种内源性生长素吲哚 - 3 - 丁酸(IBA)的β - 氧化或通过水解IAA缀合物(其中IAA与氨基酸、糖或肽相连)来获得IAA。为使IAA永久失活,植物可采用缀合和直接氧化的方式。与其作为激素的定义一致,IAA可从植物地上部分运输到根部;这种运输对正常发育是必需的,而向性反应则需要更局部的运输。生长素信号传导至少在很大程度上由SCFTIR1 E3泛素连接酶复合物介导,该复合物响应IAA加速Aux/IAA阻遏物的降解,从而改变基因表达。两类生长素诱导基因编码负向作用产物(Aux/IAA转录阻遏物和IAA缀合酶的GH3家族),这表明及时终止生长素信号至关重要。生长素与其他激素信号的相互作用给理解生长素反应带来了更多挑战。
在IAA结构阐明近六十年后,生长素代谢、运输和信号传导的许多方面已得到充分确立;然而,仍有一些基本问题和无数细节尚未解决。