环境因子调控植物花青素苷合成及呈色的机理

花青素苷（anthocyanin）是决定被子植物花、果实和种皮等颜色的重要色素之一。花青素苷的合成与积累过程往往与植物发育过程密切相关,由内外因子共同控制。环境因子通过诱导植物体内花青素苷合成途径相关基因的表达来调控花青素苷的呈色反应。该文追踪了国内外相关研究,认为光是影响花青素苷呈色的主要环境因子之一,光质和光强均能在一定程度上影响花青素苷的合成,其中光质起着更为关键的作用;低温能诱导花青素苷的积累,高温则会加速花青素苷的降解;不同的糖类物质均能影响花青素苷的合成,大部分结构基因和调节基因的表达均受糖调控。关于花发育与花青素苷呈色的关系、观赏植物花色对环境因子的响应以及花青素苷抵御逆境的机理尚待深入研究。因此,综合考察花发育与植物花青素苷合成及其呈色之间的关系,特别是光周期对花发育的影响导致花青素苷合成及呈色的机理是花色研究的一个重要课题。利用环境因子调控花色将会极大地提高花卉的观赏价值。     

环境因子调控植物花青素苷合成及呈色的机理

花青素苷(anthocyanin)是决定被子植物花、果实和种皮等颜色的重要色素之一。花青素苷的合成与积累过程往往与植物发育过程密切相关, 由内外因子共同控制。环境因子通过诱导植物体内花青素苷合成途径相关基因的表达来调控花青素苷的呈色反应。该文追踪了国内外相关研究, 认为光是影响花青素苷呈色的主要环境因子之一, 光质和光强均能在一定程度上影响花青素苷的合成, 其中光质起着更为关键的作用; 低温能诱导花青素苷的积累, 高温则会加速花青素苷的降解;不同的糖类物质均能影响花青素苷的合成, 大部分结构基因和调节基因的表达均受糖调控。关于花发育与花青素苷呈色的关系、观赏植物花色对环境因子的响应以及花青素苷抵御逆境的机理尚待深入研究。因此, 综合考察花发育与植物花青素苷合成及其呈色之间的关系, 特别是光周期对花发育的影响导致花青素苷合成及呈色的机理是花色研究的一个重要课题。利用环境因子调控花色将会极大地提高花卉的观赏价值。     

有关花青素色素发色机理研究介绍

花青素色素主要存在于高等植物中,能产生橙、红、紫、蓝等各种颜色,是重要的一类植物色素,对于花青素色素发色的机理,过去一般单纯地用花青甙的黄(钅羊)盐结构在不同溶液H离子浓度影响下,结构发生变化来解释的,如图1所示。     

基于类胡萝卜素着色的鸟类羽色多样性形成机制

人们对动物体色的研究由来已久。作为一类让生物呈现出多变色彩的重要色素, 类胡萝卜素可以在鸟类的羽毛、鸟喙和皮肤等体表组织中沉积, 产生红、橙、黄、粉、紫等颜色。类胡萝卜素不能在鸟类体内合成, 需从食物中摄取, 进而在体内完成吸收、运输、代谢和沉积等一系列过程, 才能用于羽毛着色。与类胡萝卜素着色相关的生理及遗传调控机制一直备受关注, BCO2、SCARB1和CYP2J19等影响类胡萝卜素在鸟类羽毛中着色的关键基因, 推动了对羽色遗传调控机制的深入认识。本文介绍了鸟类可利用类胡萝卜素的主要类型和基本特征, 综述了类胡萝卜素着色相关的生理过程以及调控基因研究的最新进展, 旨在增加对鸟类羽毛中类胡萝卜素着色过程和相关遗传机制的理解。     

植物花瓣呈色机理及花色分子育种

植物花瓣呈色的主要化学物质包括类黄酮/花青素苷、类胡萝卜素和甜菜色素。其中类黄酮/花青素苷是分布最广泛的色素,决定大多数植物花瓣的呈色;类胡萝卜素在一些植物黄色至橙红色花瓣中起着作用;而甜菜色素主要存在于石竹目植物,包含甜菜红素和甜菜黄素。目前,关于色素生物合成的分子网络已被解析,主要由一系列结构基因控制;一些与色素合成相关的调控因子在很多植物中被鉴定发现。另外,基于外源基因表达或内源基因编辑的分子育种在一些观赏植物的花色改良中被成功应用。本文系统性总结了植物中3种类型色素合成的分子基础、调控机制及分子育种应用等方面的研究进展;将有助于提高我们对植物色素合成分子调控网络的认识,并以期为今后开展花色分子设计育种提供理论支持。     

彩叶植物叶片呈色分子机制研究进展

彩叶植物叶片呈现不同的颜色主要是受遗传因素和外部环境的共同作用,揭示彩叶植物叶片呈色机制对选育彩叶植物新品种和彩叶植物的应用推广具有重要理论和实践意义。目前对彩叶植物呈色机制的研究主要集中于叶片中色素变化、光合特性、叶片结构和环境条件等方面。该文主要対近年来有关彩叶植物叶片中叶绿素代谢途径、类胡萝卜素代谢途径、次生代谢途径、光合作用和叶绿体发育相关结构基因和转录因子调控机制的研究进展进行综述,并对以后的研究方向进行了展望,为培育彩叶植物新品种提供了理论基础,也为人工调控叶色以及叶色的定向遗传改良提供了参考。     

二氢黄酮醇 4 还原酶在花青素合成中的功能及调控研究进展

植物色素主要有花青素、类胡萝卜素和生物碱类色素三大类,其中花青素是决定大部分被子植物组织或器官颜色的重要色素。花青素通过类黄酮途径合成,该途径是生物学上研究较多且较为清楚的代谢途径之一。近年来的研究表明,在该途径中除了查尔酮合成酶(chalcone synthase,CHS)、查尔酮异构酶(chalcone isomerase,CHI)和黄烷酮-3-羟化酶(flavanone-3-hydrolase,F3H)起着关键作用外,二氢黄酮醇-4-还原酶(dihydroflavonol 4-reductase,DFR)对花青素的合成也至关重要。DFR可催化3种二氢黄酮醇和2种黄烷酮生成5种不同的花青素前体,且DFR基因家族不同成员对各个底物的催化效率不同,因此它在一定程度上决定着植物中花青素的种类和含量,从而影响植物组织或器官的颜色。该文对近年来国内外有关DFR在花青素合成过程中的生物学功能与调控,包括DFR的特征、作用机制和系统进化以及环境、转录因子和一些结构基因与DFR的关系等方面的研究进展进行了综述,以期为DFR今后的研究和利用基因工程改变植物组织或器官的颜色提供理论依据。     

花青素代谢途径与植物颜色变异

花青素是种子植物呈色的重要色素, 由一系列结构基因编码的酶(CHS、CHI、F3H、F3'H、F3'5'H、DFR、ANS和3GT)催化而成, 随后经过各种修饰被转运至液泡等部位储存。各类器官中差异表达的MYB、bHLH和WDR三种调控因子通过形成MBW复合体直接正调控以上结构基因的表达。这个过程涉及的基因变异常会导致植物的各种颜色变异。在生活中人们广泛利用这些变异品种, 取其丰富色味。造成颜色变异的具体分子机制在很多情况下还不清楚, 但日益积累的个例研究为其中的规律性提供了基础数据。该文概述了花青素的合成、转运过程及其转录调控机制, 探讨了研究颜色变异品种的常用思路及方法。在总结近年工作的基础上, 对生活中常见蔬菜、水果和花卉的颜色变异品种的分子机制进行了综述。     

植物叶色形成调控机制研究进展

彩叶植物具有色彩鲜艳、观赏期长等特点,有助于提高城市绿化的观赏性。叶绿素、类胡萝卜素和花青素等天然色素的含量变化使叶片产生绿色、黄色、白色和紫红色等颜色,3种色素在光反应、响应生物和非生物胁迫中发挥重要作用。本文对影响叶绿素、类胡萝卜素和花青素生物合成途径遗传调控和外部环境因子综述,为解释叶片呈色机制提供理论基础。现有研究表明,光(光周期、光照强度及光质)、温度、干旱和盐等环境因子及激素变化均会刺激HY5、PIFs、DELLA等转录因子和结构基因转录,同时甲基化、乙酰化等染色质修饰和miRNAs、lncRNAs等转录后表观遗传修饰也会直接或间接调控3种色素生物合成途径基因的表达。虽然目前3种色素生物合成途径已较清晰,但有关彩叶林木3种色素代谢与环境信号、体内激素的具体调控模式仍有待进一步研究。未来可构建彩叶植物杂交群体和种质资源库,并利用基因组、转录组、蛋白组、代谢组和表型组等多组学技术为创制彩叶新种质提供可能。     

基于花青素代谢培育蓝色花卉的研究进展

花色是植物吸引昆虫传播花粉的主要因素,对于植物在自然界的生存必不可少,也是观赏植物最重要的性状之一。在蓬勃发展的花卉产业中,色彩各异花卉的培育,可以弥补自然花色的匮乏,但是令人垂涎的蓝色花比较难培育。花色的多样性主要是由花青素及其衍生物的种类和含量等因素决定的,飞燕草色素的合成是形成蓝色花的关键因素,许多植物体内缺少合成飞燕草色素的结构基因。近年来,利用基因工程技术培育蓝色花的研究也时有报道。文中以常见的观赏植物为例,基于花青素代谢调控,从影响飞燕草色素合成的关键因素和不同分子改良途径培育蓝色花等几个方面对植物花朵呈色的机制进行了综述,并展示不同分子育种策略可能在其他领域的应用,为其他植物或经济作物的色泽改良如彩色棉蓝色纤维的培育等提供参考和技术支持。     

Melanin coloration in New World orioles II: ancestral state reconstruction reveals lability in the use of carotenoids and phaeomelanins

Although many animals use carotenoids to produce bright yellow, orange, and red colors, an increasing number of studies have found that other pigments, such as melanins, may also be used to produce bright colors. Yet, almost nothing is known about the evolutionary history of this colorful melanin use. We used reflectance spectrometry to determine whether colors in New World orioles were predominantly due to carotenoids, colorful melanins, or a mixture of both. We then used ancestral state reconstruction to infer the directionality of any pigment changes and to test for phylogenetic signal. We found that three oriole taxa likely switched from carotenoid- to melanin-based colors. Several other oriole taxa apparently gained localized melanin coloration, or had coloration that seemed to be produced by a mixture of carotenoids and melanins. We also found little phylogenetic signal on the use of carotenoids or melanins to produce color. However, all pigment changes occurred within one of three major clades of the oriole genus, suggesting there may be signal at deeper phylogenetic levels. These repeated independent switches between carotenoid and melanin colors are surprising in light of the important signaling role that color pigments (especially carotenoids) are thought to play across a wide range of taxa.     

Chemical Characterization of Melanins in Sheep Wool and Human Hair

The color of hair and wool in mammals and feathers in birds is mostly determined by the quantity and quality of melanins that are synthesized in follicular melanocytes and transferred to keratinocytes. There are two chemically distinct types of melanin pigments: the black to brown eumelanins and the yellow to reddish pheomelanins. Melanins in sheep wool and human hair of various colors were characterized by HPLC methods to estimate 5,6-dihydroxyindole-2-carboxylic acid (DHICA)-derived units in eumelanins and benzothiazine units in pheomelanins. Melanins were also characterized by spectrophotometric methods after differential solubilization in alkalies. It was demonstrated that 1) black wool in Asiatic sheep contains eumelanin with the DHICA content similar to black mouse melanin, while black to brown melanins from human hair contain much lower ratios of DHICA-derived units, comparable to the slaty mutation in mice, 2) dark brown to brown hair in human contains eumelanin whose chemical properties are indistinguishable from those of black hair, 3) dark red wool and red human hair contain pheomelanic pigments whose chemical properties are rather different from those of yellow pheomelanins in mice, and 4) light brown, blonde, and red hairs in human can be differentiated from each other with this methodology.     

Contributions of pterin and carotenoid pigments to dewlap coloration in two anole species

Animals can acquire bright coloration using a variety of pigmentary and microstructural mechanisms. Reptiles and amphibians are known to use two types of pigments - pterins and carotenoids - to generate their spectrum of colorful red, orange, and yellow hues. Because both pigment classes can confer all of these hues, the relative importance of pterins versus carotenoids in creating these different colors is not always apparent. We studied the carotenoid and pterin content of red and yellow dewlap regions in two neotropical anole species - the brown anole (Norops sagrei) and the ground anole (N. humilis). Pterins (likely drosopterins) and carotenoids (likely xanthophylls) were present in all tissues from all individuals. Pterins were more enriched in the lateral (red) region, and carotenoids more enriched in the midline (yellow) region in N. humilis, but pterins and carotenoids were found in similar concentrations among lateral and midline regions in N. sagrei. These patterns indicate that both carotenoid and pterin pigments are responsible for producing color in the dichromatic dewlaps of these two species, and that in these two species the two pigments interact differently to produce the observed colors.     

Melanin coloration in New World orioles I: carotenoid masking and pigment dichromatism in the orchard oriole complex

Carotenoids produce the brilliant red, orange, and yellow colors of many animals. However, melanin pigments can also confer some of these same hues. Because carotenoid and melanin colors are produced in different ways and may serve different signaling functions, either within or between species, it is important to establish whether one or both types of pigment are responsible for coloration. We have discovered what appears to be an evolutionary switch from carotenoid- to melanin-based color in two sexually dichromatic New World orioles. Using a combination of reflectance spectrometry and chromatographic analyses of plumage pigments, we found that the chestnut plumage of adult male orchard orioles Icterus spurius is produced predominantly by phaeomelanins. Orchard oriole feathers also contain carotenoids, which appear to be masked by the high concentration of phaeomelanins. In contrast, both carotenoids and phaeomelanins appear to contribute to color in adult male Fuertes's orioles I. fuertesi . Moreover, yellow yearling male and female plumage in both species is produced by carotenoids alone. The masking of carotenoids with phaeomelanins in orchard orioles is interesting in light of the signaling roles that carotenoids are thought to play. In addition, these plumage differences produce a unique case of age and sexual pigment dimorphism in orchard and Fuertes's orioles.     

Transcript profiling of genes involved in carotenoid biosynthesis among three carrot cultivars with various taproot colors

Protoplasma - Carotenoids are a group of natural pigments that are widely distributed in various plants. Carrots are plants rich in carotenoids and have fleshy roots with different colors....     

The influence of carotenoid acquisition and utilization on the maintenance of species-typical plumage pigmentation in male American goldfinches (Carduelis tristis) and northern cardinals (Cardinalis cardinalis).

Birds display a tremendous variety of carotenoid-based colors in their plumage, but the mechanisms underlying interspecific variability in carotenoid pigmentation remain poorly understood. Because vertebrates cannot synthesize carotenoids de novo, access to pigments in the diet is one proximate factor that may shape species differences in carotenoid-based plumage coloration. However, some birds metabolize ingested carotenoids and deposit pigments that differ in color from their dietary precursors, indicating that metabolic capabilities may also contribute to the diversity of plumage colors we see in nature. In this study, we investigated how the acquisition and utilization of carotenoids influence the maintenance of species-typical plumage pigmentation in male American goldfinches (Carduelis tristis) and northern cardinals (Cardinalis cardinalis). We supplemented the diet of captive goldfinches with red carotenoids to determine whether males, which are typically yellow in color, were capable of growing red plumage. We also deprived cardinals of red dietary pigments to determine whether they could manufacture red carotenoids from yellow precursors to grow species-typical red plumage. We found that American goldfinches were able to deposit novel pigments in their plumage and develop a striking orange appearance. Thus, dietary access to pigments plays a role in determining the degree to which goldfinches express carotenoid-based plumage coloration. We also found that northern cardinals grew pale red feathers in the absence of red dietary pigments, indicating that their ability to metabolize yellow carotenoids in the diet contributes to the bright red plumage that they display.     

植物LYCs的特性功能及其相互关系

植物番茄红素环化酶（LYC）是将线性番茄红素分子转化为具环类胡萝卜素的关键酶,本文从番茄红素环化酶cDNA的核苷酸和氨基酸序列,底物特异性以及作用特点等方面,阐述了两种重要的番茄红素环化酶即β－环化酶和ε－环化酶的异同,并对这两种酶的作用特点,对植物具环类胡萝卜素种类及总量的调节作用以及对其它环化酶的相互关系进行了讨论。     

萱草花瓣中类胡萝卜素分析样品的制备及UPCC-MS检测方法研究

以大苞萱草（Hemerocallis middendorfii Trautv.et Mey.）和‘原谅’萱草（H.‘Pardon Me’）为研究对象,对萱草属植物花瓣中类胡萝卜素的样品制备方法以及UPCC-MS定性和定量检测方法进行了研究。结果表明：（1）萱草花瓣类胡萝卜素样品制备过程中,不同的提取试剂、振荡方法及皂化方法对类胡萝卜素的提取效率均有显著影响,经过对提取结果的方差分析,确定最佳的样品制备方案为：提取试剂B丙酮：正己烷（3：5/V：V）、温控摇床振荡提取,常温皂化16 h;（2） UPCC-MS技术能在10 min内高效分离萱草花瓣中的类胡萝卜素,且使用的有毒化学试剂少,是检测类胡萝卜素的较好选择;（3）大苞萱草和原谅萱草花瓣中共含有20种类胡萝卜素物质,两者颜色不同,类胡萝卜素的组成和含量也存在差异。     

Catecholamine melanins. Structural changes induced by copper ions

Melanins synthesized from adrenaline and dopamine in the presence or absence of copper ions were characterized by pyrolysis-gas chromatography-mass spectrometry and by IR and ESR methods. It was shown that Cu2+ are able to induce changes in the melanin structure. Melanins obtained from adrenaline-Cu2+ and dopamine-Cu2+ complexes are composed mainly from monomeric units of the indole type. Melanins synthesized from these catecholamines without Cu2+ contain additionally large amounts of unindolized monomeric units. The structure differences in both types of melanins are reflected in their sorptive abilities and spectroscopic characteristics.     

Biosynthesis of plant pigments: anthocyanins, betalains and carotenoids

Plant compounds that are perceived by humans to have color are generally referred to as 'pigments'. Their varied structures and colors have long fascinated chemists and biologists, who have examined their chemical and physical properties, their mode of synthesis, and their physiological and ecological roles. Plant pigments also have a long history of use by humans. The major classes of plant pigments, with the exception of the chlorophylls, are reviewed here. Anthocyanins, a class of flavonoids derived ultimately from phenylalanine, are water-soluble, synthesized in the cytosol, and localized in vacuoles. They provide a wide range of colors ranging from orange/red to violet/blue. In addition to various modifications to their structures, their specific color also depends on co-pigments, metal ions and pH. They are widely distributed in the plant kingdom. The lipid-soluble, yellow-to-red carotenoids, a subclass of terpenoids, are also distributed ubiquitously in plants. They are synthesized in chloroplasts and are essential to the integrity of the photosynthetic apparatus. Betalains, also conferring yellow-to-red colors, are nitrogen-containing water-soluble compounds derived from tyrosine that are found only in a limited number of plant lineages. In contrast to anthocyanins and carotenoids, the biosynthetic pathway of betalains is only partially understood. All three classes of pigments act as visible signals to attract insects, birds and animals for pollination and seed dispersal. They also protect plants from damage caused by UV and visible light.