Department of Biological Sciences, Lehman College, City University of New York, Bronx, New York 10468, USA.
Plant Physiol. 2010 May;153(1):66-79. doi: 10.1104/pp.110.153916. Epub 2010 Mar 24.
Metabolic engineering of plant carotenoids in food crops has been a recent focus for improving human health. Pathway manipulation is predicated on comprehensive knowledge of this biosynthetic pathway, which has been extensively studied. However, there existed the possibility of an additional biosynthetic step thought to be dispensable because it could be compensated for by light. This step, mediated by a putative Z-ISO, was predicted to occur in the sequence of redox reactions that are coupled to an electron transport chain and convert the colorless 15-cis-phytoene to the red-colored all-trans-lycopene. The enigma of carotenogenesis in the absence of light (e.g. in endosperm, a target for improving nutritional content) argued for Z-ISO as a pathway requirement. Therefore, understanding of plant carotenoid biosynthesis was obviously incomplete. To prove the existence of Z-ISO, maize (Zea mays) and Arabidopsis (Arabidopsis thaliana) mutants were isolated and the gene identified. Functional testing of the gene product in Escherichia coli showed isomerization of the 15-cis double bond in 9,15,9'-tri-cis-zeta-carotene, proving that Z-ISO encoded the missing step. Z-ISO was found to be important for both light-exposed and "dark" tissues. Comparative genomics illuminated the origin of Z-ISO found throughout higher and lower plants, algae, diatoms, and cyanobacteria. Z-ISO evolved from an ancestor related to the NnrU (for nitrite and nitric oxide reductase U) gene required for bacterial denitrification, a pathway that produces nitrogen oxides as alternate electron acceptors for anaerobic growth. Therefore, plant carotenogenesis evolved by recruitment of genes from noncarotenogenic bacteria.
植物类胡萝卜素的代谢工程一直是近年来改善人类健康的重点。途径操作的前提是对该生物合成途径有全面的了解,该途径已得到广泛研究。然而,由于有一种额外的生物合成步骤被认为是可有可无的,因为它可以通过光来补偿,所以存在这种可能性。这个步骤,由一个假定的 Z-ISO 介导,被预测发生在与电子传递链偶联的氧化还原反应序列中,将无色 15-顺式-phytoene 转化为红色全反式-lycopene。在没有光的情况下(例如在胚乳中,这是提高营养含量的目标)进行类胡萝卜素生物合成的谜团,这证明了 Z-ISO 是途径的要求。因此,对植物类胡萝卜素生物合成的理解显然是不完整的。为了证明 Z-ISO 的存在,分离了玉米(Zea mays)和拟南芥(Arabidopsis thaliana)突变体,并鉴定了该基因。对该基因产物在大肠杆菌中的功能测试表明,9,15,9'-三-cis-ζ-胡萝卜素中的 15-顺式双键发生异构化,证明 Z-ISO 编码了缺失的步骤。发现 Z-ISO 对暴露在光下和“黑暗”组织都很重要。比较基因组学阐明了在高等植物、藻类、硅藻和蓝藻中发现的 Z-ISO 的起源。Z-ISO 是从与细菌反硝化所需的 NnrU(用于亚硝酸盐和一氧化氮还原酶 U)基因相关的祖先进化而来的,反硝化是一种产生氮氧化物作为厌氧生长的替代电子受体的途径。因此,植物类胡萝卜素的生物合成是通过从非类胡萝卜素产生的细菌中招募基因而进化而来的。