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。     

Genetic Modification in Floriculture

An important driving force for the floriculture industry is the development of novel plants and flowers. New varieties provide marketing opportunities for retailers and judicious selection can increase productivity for growers, as well as improving the quality of the final product in the consumer's hands. While plant exploration and conventional breeding programs have been very successful in achieving these goals, genetic modification offers additional routes for the generation of new varieties of important floricultural plants. This can be achieved by the incorporation of genes from outside of the normally available gene pool. This paper provides a summary of the potential applications of gene technology in floriculture and reviews progress to date, with a particular emphasis on the manipulation of flower color. The manipulation of the anthocyanin biosynthesis pathway in carnation to produce novel-colored flowers is so far the only commercial application of genetic modification in floriculture. This progress is in stark contrast to the widespread cultivation of genetically modified broad-acre crops. The commercial use of gene technology requires adherence to regulatory regimes specific to genetically modified plants, and compliance with intellectual property laws. These added complexities are a significant cost, which may be hampering the use of gene technology by breeders of floricultural crops. Another factor may be a perception that the public and retail trade may not accept genetically modified floricultural products. Experience in the real marketplace with the Florigene Moon-series? of genetically modified carnation suggests that these concerns are unwarranted.     

Production of red-flowered plants by genetic engineering of multiple flavonoid biosynthetic genes

Orange- to red-colored flowers are difficult to produce by conventional breeding techniques in some floricultural plants. This is due to the deficiency in the formation of pelargonidin, which confers orange to red colors, in their flowers. Previous researchers have reported that brick-red colored flowers can be produced by introducing a foreign dihydroflavonol 4-reductase (DFR) with different substrate specificity in Petunia hybrida, which does not accumulate pelargonidin pigments naturally. However, because these experiments used dihydrokaempferol (DHK)-accumulated mutants as transformation hosts, this strategy cannot be applied directly to other floricultural plants. Thus in this study, we attempted to produce red-flowered plants by suppressing two endogenous genes and expressing one foreign gene using tobacco as a model plant. We used a chimeric RNAi construct for suppression of two genes (flavonol synthase [FLS] and flavonoid 3′-hydroxylase [F3′H]) and expression of the gerbera DFR gene in order to accumulate pelargonidin pigments in tobacco flowers. We successfully produced red-flowered tobacco plants containing high amounts of additional pelargonidin as confirmed by HPLC analysis. The flavonol content was reduced in the transgenic plants as expected, although complete inhibition was not achieved. Expression analysis also showed that reduction of the two-targeted genes and expression of the foreign gene occurred simultaneously. These results demonstrate that flower color modification can be achieved by multiple gene regulation without use of mutants if the vector constructs are designed resourcefully. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Flower color modulations of Torenia hybrida by downregulation of chalcone synthase genes with RNA interference

Suppression of biosynthetic genes involved in flower color formation is an important approach for obtaining target flower colors. Here we report that flower color of the garden plant Torenia hybrida was successfully modulated by RNA interference (RNAi) against a gene of chalcone synthase (CHS), a key enzyme for anthocyanin and flavonoid biosynthesis. By using each of the coding region and the 3'-untranslated region of the CHS mRNA as an RNAi target, exhaustive and gene-specific gene silencing were successfully induced, and the original blue flower color was modulated to white and pale colors, respectively. Our results indicate that RNAi is quite useful for modulations of flower colors of commercially important garden plants.     

观赏植物花色基因工程研究进展

花色是观赏植物的重要性状,创造新花色是花卉育种的主要目标之一。基因工程技术 在观赏植物花色育种上可弥补传统育种技术的缺陷,因此它在花色育种方面的研究和应用发 展迅速。本文从花的成色作用和花色素种类入手,介绍了花色苷的生物合成,并从花色基因 的种类和克隆、花色基因工程操作的策略和方法等角度综述了近年来观赏植物花色基因工程 的研究进展。同时对我国观赏植物花色基因工程的前景作一展望。     

观赏植物花色基因工程研究进展

花色是观赏植物的重要性状,创造新花色是花卉育种的主要目标之一。基因工程技术在观赏植物花色育种上可弥补传统育种技术的缺陷,因此它在花色育种方面的研究和应用发展迅速。本文从花的成色作用和花色素种类人手,介绍了花色苷的生物合成,并从花色基因的种类和克隆、花色基因工程操作的策略和方法等角度综述了近年来观赏植物花色基因工程的研究进展。同时对我国观赏植物花色基因工程的前景作一展望。     

Mechanisms underlying flower color variation in Asian species of Meconopsis: A preliminary phylogenetic analysis based on chloroplast DNA and anthocyanin biosynthesis genes

The Qinghai–Tibet Plateau (QTP) harbors the highest species diversity of alpine plants in the world, with a spectacular diversity of flower colors. Among these QTP plants, the genus Meconopsis comprises more than 50 species, for which flower color is a key diagnostic character. However, the mechanisms underlying flower color variation have rarely been investigated. In the present study, we used three chloroplast (cp) DNA fragments and two anthocyanin biosynthesis genes (F3H andF3′H) for phylogenetic reconstruction of Meconopsis. We revealed the presence of three well-supported clades and/or subclades in the cpDNA and nuclear gene trees; further, flower color transition occurred in each lineage. The results of selection tests and preliminary expression analyses of the anthocyanin biosynthesis genes indicate that the pigment pathway leading to cyanidin is active in blue and red flowers of Meconopsis; further, a blue–red color shift is not attributable to an on/off switching of the anthocyanin biosynthetic pathway (ABP) branches. Together with the results of previous flower pigment analyses, our findings suggest that blue–red flower color transitions in Meconopsis are attributable to modification of cyanidin. Our molecular dating results indicate that the lineage diversification inMeconopsis is closely related to the QTP uplift; thus, it is likely that environmental changes arising from the QTP uplift have played important roles in driving the diversification of flavonoids, through which species of Meconopsis have adapted physiologically to diverse habitats.     

Flower Color Modification by Engineering of the Flavonoid Biosynthetic Pathway: Practical Perspectives

The status quo of flavonoid biosynthesis as it relates to flower color is reviewed together with a success in modifying flower color by genetic engineering. Flavonoids and their colored class compounds, anthocyanins, are major contributors to flower color. Many plant species synthesize limited kinds of flavonoids, and thus exhibit a limited range of flower color. Since genes regulating flavonoid biosynthesis are available, it is possible to alter flower color by overexpressing heterologous genes and/or down regulating endogenous genes. Transgenic carnations and a transgenic rose that accumulate delphinidin as a result of expressing a flavonoid 3′,5′-hydroxylase gene and have novel blue hued flowers have been commercialized. Transgenic Nierembergia accumulating pelargonidin, with novel pink flowers, has also been developed. Although it is possible to generate white, yellow, and pink-flowered torenia plants from blue cultivars by genetic engineering, field trial observations indicate difficulty in obtaining stable phenotypes.     

蓝色花形成分子机理研究进展

花色是观赏植物的重要特征, 在自然界中蓝色花占比很少, 很多观赏植物都缺少蓝色种质。因此, 研究蓝色花形成的分子机理对于蓝色花定向育种具有重要意义。研究表明, 花色的形成主要是通过花青苷积累, 花青素通过糖基化形成花青苷, 再通过酰基、甲基化修饰及金属离子络合反应, 在特定的液泡pH环境中呈现出稳定的蓝色。该文从花青苷合成与代谢途径入手, 对蓝色花形成关键基因功能、花青苷各位点酰化的影响、金属离子的作用、液泡pH值相关基因研究及蓝色花分子育种等方面进行综述。     

欧报春（Primula vulgaris）不同花色与色素关系及花色遗传初步分析

为了掌握欧报春各花色遗传规律服务于良种生产,通过对欧报春各色花进行色素吸收光谱和薄层层析分析,进行不同花色杂交研究,分析了欧报春各色花所含色素类型及各花色遗传规律。结果显示欧报春群体含多种花色素,单株也可含有多种花色素,形成多变的粉色、红色及蓝色花。黄色深浅主要由类胡萝卜素含量决定。白色对粉色及黄色为隐性遗传,黄色、粉色为显性遗传并有数量遗传特征,黄色与粉色独立遗传。蓝色为多基因控制的隐性遗传,并具有数量遗传特征。     

花色改造基因工程

自1987年世界首例成功运用转基因技术改造矮牵牛花色以来,花色改造基因工程技术不断展现它在培育新花色品系上的无穷魅力。介绍了近年来运用基因工程技术成功改造花色的3种主要策略:(1)采用反义RNA及共抑制的方法来改变花颜色的深浅;(2)通过导入新基因产生新奇花色;(3)利用转座子构建特殊表达载体,随机激活花色合成的基因来产生嵌合花色。此外,还对转基因株花色不稳定原因进行了讨论。     

Composition of Carotenoids and Flavonoids in Narcissus Cultivars and their Relationship with Flower Color

Narcissus is widely used for cut flowers and potted plants, and is one of the most important commercial bulbous flowers in the floricultural industry. In this study, ten carotenoid and eighteen flavonoid compounds from the perianths and coronas of fifteen narcissus cultivars were measured by HPLC–APCI-MS/MS and UPLC-Q-TOF-MS/MS. Among these, six carotenoids, a total of seventeen flavonols and chlorogenic acid were identified in narcissus for the first time. A multivariate analysis was used to explore the relationship between flower color and pigment composition. We found that all-trans-violaxanthin and total carotenoid content were the main factors that affected flower color. These investigations could provide a global view of flower color formation and a theoretical basis for hybridization breeding in narcissus.     

Genetic engineering of the anthocyanin biosynthetic pathway with flavonoid-3′,5′-hydroxylase: specific switching of the pathway in petunia

Flavonoid-3',5'-hydroxylase (F3'5'H) is the key enzyme in the synthesis of 3',5'-hydroxylated anthocyanins, which are generally required for the expression of blue or purple flower color. It has been predicted that the introduction of this enzyme into a plant species that lacks it would enable the production of blue or purple flowers by altering the anthocyanin composition. We present here the results of the genetic engineering of petunia flower color, pigmentation patterns and anthocyanin composition with sense or antisense constructs of the F3'5'H gene under the control of the CaMV 35S promoter. When sense constructs were introduced into pink flower varieties that are deficient in the enzyme, transgenic plants showed flower color changes from pink to magenta along with changes in anthocyanin composition. Some transgenic plants showed novel pigmentation patterns, e.g. a star-shaped pattern. When sense constructs were introduced into blue flower petunia varieties, the flower color of the transgenic plants changed from deep blue to pale blue or even pale pink. Pigment composition analysis of the transgenic plants suggested that the F3'5'H transgene not only created or inhibited the biosynthetic pathway to 3',5'-hydroxylated anthocyanins but switched the pathway to 3',5'-hydroxylated or 3'-hydroxylated anthocyanins.     

Are pollinators and seed predators selective agents on flower color in <Emphasis Type="Italic">Gentiana lutea</Emphasis>?

Animals which interact with plants often cause selective pressures on plant traits. Flower color variation within a species might be shaped by the action of animals feeding on the plant species. Pollinators might exert natural selection on color if flower color is related to their foraging efficiency. For example, some pollinator species might require more time to detect particular colors. If that is the case, flower color might have evolved as a pollination exploitation barrier—ensuring that flowers are more visited by the most efficient pollinators. In addition, non-pollinator agents such as predispersal seed predators may select on flower color, if color indicates food resources (seeds) or if color is related to deterrent compounds. We address selection on flower color in a population of Gentiana lutea where color varies among individuals from yellow to orange. We hypothesize that opposed selection from mutualists (pollinators) and antagonists (predispersal seed predators) maintains flower color variation in this population. By means of path analysis we addressed the role of both interactors in flower color selection. We found that selection acts on flower color, mediated by both pollinators and seed predators. Both agents favored yellow-flowered individuals, thus selection by pollinators and seed predators does not maintain flower color variation in this population.     

植物花青素生物合成相关基因的研究及应用

花青素是决定植物花色的主要色素,使大多数花呈现从红到蓝的系列变化,是花色研究和开发的重点,并具有重要的营养和药用作用。目前花青素生物合成途径已日益清楚,并已分离到大量的相关酶和基因,并获得了一批具有商业价值的转基因植物新品种。本文重点介绍了花青素合成途径中关键基因的研究成果,并概述了国内外花青素基因在植物基因工程中的应用研究进展情况,同时对花青素基因的研究应用前景和发展趋势作一展望。     

Are pan traps colors complementary to sample community of potential pollinator insects?

The global initiatives of monitoring and conserving pollinators require worldwide assessments with comparable data sets collected through standardized methods. The use of pan traps is a passive method widely applied to sample flower visitors, standing out for its simplicity. Despite its wide use to sample pollinator diversity, the influence of color on trap efficiency is not well understood. The available studies are particularly scarce in the tropics and have generated divergent results. The main goal of the present study was to assess whether blue, yellow and white pan traps are complementary to sample Hymenoptera community. For this, we placed 49 sample units of blue, white and yellow pan traps in agricultural and natural (savanna-like) areas in Chapada Diamantina, Bahia, Brazil. We found that the species richness from blue and yellow pan traps were not significantly different, but both were significantly greater than the species richness from white pan traps. However, bees were significantly more attracted to the blue pan traps and wasps to the yellow ones; thus, color attractiveness was group-specific. Pan traps of different color showed low species composition overlap with 61 % of species collected exclusively in one of the three pan trap colors, and the species composition in the blue traps differed consistently from that in the traps of the other colors. In the article we discuss the implication of the results and defend the combined use of pan traps with different colors as a solution for the differential variable sample bias.     

Floral color change and the attraction of insect pollinators in lungwort (Pulmonaria collina)

We studied the effect of floral color change on long- and short-distance attraction of insect pollinators to the herb lungwort, Pulmonaria collina. Lungwort flowers change color with age from red to blue. Young red flowers had a significantly greater pollen and nectar reward and were significantly more often unpollinated than old blue ones. Red and blue flowers both influenced long-distance attractiveness of plants, defined as the number of insect approaches towards an individual plant. After reaching a plant, flower visitors preferred to visit young red flowers. Therefore, short-distance attractiveness, defined as the number of flowers visited successively on an individual plant, was influenced mainly by the number of young red flowers. The co-occurrence of the change in reproductive ability, in amount of reward, and in flower color enabled lungwort plants to direct pollinators to reproductive, highly rewarding red flowers. The data suggest that by maintaining changed flowers lungwort plants can increase their long-distance attraction and simultaneously enhance the probability of flower visits to pre-changed flowers. Thus, we propose floral color change as a mechanism that can increase the efficiency of pollen transfer to enhance plant fitness. Received: 2 November 1998 / Accepted: 14 July 1999     

梅花‘南京红须’、‘南京红’花色的呈现特征(英文)

梅花‘南京红须’、‘南京红’的花色主要存在着花发育阶段导致的时间变化,反映其花色受花发育控制。二者的花色都在蕾期最浓艳,在初花期略淡,在盛花期又稍浓,在末花期最淡,尽管花瓣在花开放时便开始衰老;在整个花发育时期,同一朵花不同层次花瓣的颜色浓淡均为:外层花瓣>中层花瓣>内层花瓣,即花瓣在花冠中的具体排列位置决定着该片花瓣的特定颜色深浅;但不同层次花瓣颜色的变化趋势不完全一致。同时,两个品种外层花瓣的总黄酮含量变化与外层花瓣的色度变化成正相关。而花朵在树冠的着生部位导致的花色差异极不显著,表明‘南京红须’、‘南京红’的花色的空间变化极微。本文可为梅花红色花色的机理探索和花色色素生物合成关键酶基因cDNA克隆中的花朵选择提供参考。     

The Flavonoid Pathway Regulates the Petal Colors of Cotton Flower

Although biochemists and geneticists have studied the cotton flower for more than one century, little is known about the molecular mechanisms underlying the dramatic color change that occurs during its short developmental life following blooming. Through the analysis of world cotton germplasms, we found that all of the flowers underwent color changes post-anthesis, but there is a diverse array of petal colors among cotton species, with cream, yellow and red colors dominating the color scheme. Genetic and biochemical analyses indicated that both the original cream and red colors and the color changes post-anthesis were related to flavonoid content. The anthocyanin content and the expression of biosynthesis genes were both increased from blooming to one day post-anthesis (DPA) when the flower was withering and undergoing abscission. Our results indicated that the color changes and flavonoid biosynthesis of cotton flowers were precisely controlled and genetically regulated. In addition, flavonol synthase (FLS) genes involved in flavonol biosynthesis showed specific expression at 11 am when the flowers were fully opened. The anthocyanidin reductase (ANR) genes, which are responsible for proanthocyanidins biosynthesis, showed the highest expression at 6 pm on 0 DPA, when the flowers were withered. Light showed primary, moderate and little effects on flavonol, anthocyanin and proanthocyanidin biosynthesis, respectively. Flavonol biosynthesis was in response to light exposure, while anthocyanin biosynthesis was involved in flower color changes. Further expression analysis of flavonoid genes in flowers of wild type and a flavanone 3-hydroxylase (F3H) silenced line showed that the development of cotton flower color was controlled by a complex interaction between genes and light. These results present novel information regarding flavonoids metabolism and flower development.