植物花色形成及其调控机理

综述了植物花色的表现、起源与进化、功能及其调控机制.植物花色主要表现为单色、变色和杂色,是长期进化的结果,主要功能是指示传粉者和保护花器官.花色素主要包括类黄酮、类胡萝卜素和生物碱.花色素的存在及其变化是植物花色表现的化学机制,色素在花瓣中的空间分布及其对光的作用是花色表现的解剖学和光学机制,细胞液pH值、花发育阶段和植物激素是花色表现的植物生理学机制.传粉者、真菌侵染、机械损伤、园艺措施、光、温度、水分、矿质营养和糖等是影响花色的外部因蝌素.花瓣彩斑主要由基因突变或病毒入侵而形成.     

植物花色的表现机理

从植物花色表现的化学机制、解剖学和光学机制、生理学机制以及基因控制或病毒侵染等方面阐述了植物花色的机理。花色素的存在及其变化是植物花色表现的化学机制,色素在花瓣中的空间分布及其对光的作用是花色表现的解剖学和光学机制,细胞液pH值、花发育阶段和植物激素是花色表现的植物生理学机制,基因控制或病毒侵染引起花色变异。     

蝶形花亚科植物花部适应机制与传粉系统

蝶形花亚科(Papilionoideae)植物种类丰富、繁育系统多样,与其传粉者关系密切,主要表现在花部适应与传粉系统的形成。蝶形花形态复杂,为完全花,花萼、花瓣5基数,花萼呈箭头状,相邻花瓣螺旋轮生,旗瓣较大位于最外边,两片翼瓣紧贴旗瓣着生并包裹两片合生的龙骨瓣,龙骨瓣内包裹着雌、雄蕊,雌蕊位于正中央,雄蕊轮生,构成二体雄蕊(多数9+1,少数5+5)。对蝶形花亚科植物的花部特征、传粉功能群、酬物与传粉系统构建进行了回顾,重点论述了蝶形花形态和化学组成与传粉系统的进化关系以及花粉呈现机制。其中,泛化的传粉系统以蜂媒传粉为主,同时存在鸟媒传粉、蝙蝠传粉和松鼠传粉等方式,花部结构和传粉者的相互选择和相互适应推动了传粉系统的演化。花粉释放是体现蝶形花植物与其传粉功能群相互作用的重要方面,二者协同进化形成了以下4种花粉呈现机制:弹花机制、活塞机制、瓣膜机制和毛刷机制。蝶形花的花部特征与传粉功能群的相互作用形成相应的传粉系统,而传粉系统是执行传粉功能的重要集合体,各个构件的相互协调和适应保证了蝶形花亚科植物传粉过程的顺利完成。     

石榴花瓣和雄蕊对其传粉过程与繁殖成功的影响

植物-传粉者相互关系中植物形成多种多样的视觉(花色)和嗅觉(花气味)信号来影响传粉者的访花过程,促进传粉成功。石榴(Punicagranatum)花瓣红色而雄蕊黄色,这种花内不同结构的颜色差异对于石榴吸引传粉者可能有不同的作用。本文比较了石榴花各部位发出的视觉信号(颜色、大小)和嗅觉信号(气味及含量)、花蜜体积、不同处理下的昆虫访花频率及坐果率,以探讨石榴花各部位颜色在传粉过程中的作用。结果发现:新疆喀什地区石榴主要传粉者为意大利蜜蜂(Apis mellifera)和食蚜蝇(Syrphidae sp.),雄蕊黄色及其分泌挥发性化合物种类和相对含量是吸引传粉者昆虫的主要因素。去除花瓣处理组与其他3种处理组(对照、去雄蕊、去雄蕊去花瓣)比较,意大利蜜蜂的访花频率(P <0.05)及停留时间(P <0.05)均显著提高,石榴坐果率(82.33%±4.45%)也显著提高。上述结果表明,石榴黄颜色的雄蕊可能是吸引传粉者的主要结构,而红色的花瓣对其传粉成功可能有负面影响;植物花内不同结构颜色差异可能有助于在变化的环境下吸引不同的传粉者,促进繁殖成功。     

鼠尾草属不同物种的雄蕊杠杆机制对传粉者空间变异的进化响应

被子植物虫媒传粉植物的物种分化通常被认为是花性状响应传粉环境(传粉者)的空间变异而发生适应性分化的结果。通过对鼠尾草属(Salvia) 3个物种(共4个居群)传粉互作系统的比较, 探索了花性状对不同传粉环境的进化响应。结果表明: 各居群的传粉者组成、主要传粉者类型及其大小各不相同, 杠杆状雄蕊及相关花部性状大小在不同居群间具有显著差异; 各居群均表现出腹部传粉和背部传粉2种传粉模式, 但背部传粉仍然是最有效的传粉方式; 居群间杠杆状雄蕊长度与传粉者体长表现出极显著的正相关, 然而花冠长与传粉者体长表现出负相关; 花冠口高度和柱头高度与传粉者胸厚也表现出一定的协同变异。鼠尾草属植物的杠杆状雄蕊及相关花部性状在传粉系统的进化过程中表现出高度的可塑性, 表明雄蕊杠杆传粉机制对传粉环境的变异非常敏感, 在该属植物的物种分化过程中具有关键作用。     

番荔枝科(Annonaceae)传粉室类型及其演化意义

花部结构与传粉者协同进化是有花植物成功演化的关键原因,在番荔枝科的植物中传粉室是一个非常明显的花部结构。由于该科花瓣的大小、轮数及每轮花瓣数、花瓣间的连合方式等高度多样,导致其传粉室形态也高度多样。番荔枝科107属中68属具详细的花部结构研究,该文根据开花期传粉室的闭合情况将这68属的传粉室分为开放型、半闭合型和闭合型3类,研究其系统演化意义。结果表明:(1) 24属具开放型传粉室,38属具半闭合型传粉室,26属具闭合型传粉室;约17属具2种或3种传粉室类型。(2)不同传粉室类型的植物与传粉者之间具有一定的对应关系,蜂类和蝇类主要为开放型传粉室或传粉室空间较大的半闭合型和闭合型传粉室植物传粉;蓟马主要为半闭合型与闭合型传粉室植物传粉。(3)古热带和新热带是番荔枝科植物的分布中心。分布于古热带的植物种数最多,但传粉室类型和传粉者较单一;分布于新热带的植物种数相对较少,但传粉室类型与传粉者都高度多样性。已有研究表明非洲是番荔枝科植物的起源地,开放型传粉室的植物主要分布于热带非洲,普遍存在于各族早期分化的属中,且均由小甲虫进行传粉,是该科较原始的传粉室类型;半闭合型分布于热带非洲、热带亚洲及热带美洲,闭合型传粉室主要分布于热带亚洲,这两类传粉室植物的传粉者也较多样性,是番荔枝科植物中较进化的传粉室类型。     

神奇的“扳机”

大多数有花植物用艳丽的花瓣来吸引传粉者,然而竹芋科植物的花瓣并不显著,颜色暗淡,退化成带状。那么,它用什么来吸引传粉者呢？原来,竹芋科鲜艳的退化雄蕊部分替代了真正花瓣的作用。竹芋科植物的雄蕊是由退化雄蕊和可育雄蕊两部分组成的,退化雄蕊不仅结构功能复杂,     

花色多样性与变异的研究进展

花的颜色不仅在不同物种之间有着丰富的多样性,同一物种的不同居群或个体之间也有着花色的多态性,同一花中的不同器官甚至同一类型的器官也有颜色差异。了解花色多样性的形成和维持机制,有助于揭示花的演化。经典的观点认为,花色是植物提供给传粉者的视觉信号,能促进传粉和提高觅食效率。在分析花色多样性的基础上,本文介绍了4种不同的研究方法,并论述了当前解释花色多样性的3个主要假说。提出今后的研究有必要结合系统发育的分析方法,综合考虑传粉者、植食动物、物理环境等多个因子的选择作用,才能深入理解花色的多样性与演化。     

Pollinator-dependent evolution of floral trait combinations in an orchid herb

作为驱动花性状进化的重要媒介之一,传粉者塑造了被子植物多样的花部特征。目前,大部分研究集中于传粉者驱动被子植物单个性状的进化,而较少涉及对传粉者驱动被子植物组合性状进化的研究。本研究以兰科植物绶草(Spiranthes sinensis)为材料,分别鉴别和估算了传粉者对植物单个性状和组合性状施加的定向选择和关联选择压力。研究结果表明,传粉者对开花时间和花大小施加定向选择压力,传粉者选择早开花、花更大的个体。传粉者对绶草花展示的组合性状(花大小与花数目)施加正向的关联选择压力,而对开花时间与花展示的组合性状(开花时间与株高、开花时间与花数目)施加负向的关联选择压力。同时,传粉者的选择强度在年际间存在差异。该研究证实了传粉者在驱动植物组合性状进化过程中的作用,同时强调在认识和理解被子植物表型性状进化轨迹时,需要考虑植物组合性状的影响。     

马先蒿属花冠形态的多样性与传粉式样的关系

马先蒿属(Pediculais)是有花植物中花冠形态多样化最为集中的属。该属主要的传粉者是熊蜂属(Bormbus)昆虫;在北美,熊蜂和蜂鸟是马先蒿植物一些种类有效的传粉者;也发现壁蜂(Osmia)为其传粉。不同的传粉机制要求某一特定的取食式样储藏和释放花粉。本文讨论了花冠类型的进化趋势与传粉式样和花粉形态的关系。传粉者的选择压力是决定花冠多样化的重要因素之一;花冠类型与传粉者和传粉行为紧密相关。马先蒿植物和传粉者的相互依赖与其花冠类型、功能和物候互相适应,但花冠类型与花粉形态两者之间似乎没有明显的一一对应关系。通过北美、日本和喜马拉雅不同地理分布马先蒿种类的比较研究表明,具有相同花冠类型的种类有着相同的传粉方式,花冠形态与传粉式样存在紧密的协同进化关系。     

Arctic mustard flower color polymorphism controlled by petal-specific downregulation at the threshold of the anthocyanin biosynthetic pathway

Intra- and interspecific variation in flower color is a hallmark of angiosperm diversity. The evolutionary forces underlying the variety of flower colors can be nearly as diverse as the colors themselves. In addition to pollinator preferences, non-pollinator agents of selection can have a major influence on the evolution of flower color polymorphisms, especially when the pigments in question are also expressed in vegetative tissues. In such cases, identifying the target(s) of selection starts with determining the biochemical and molecular basis for the flower color variation and examining any pleiotropic effects manifested in vegetative tissues. Herein, we describe a widespread purple-white flower color polymorphism in the mustard Parrya nudicaulis spanning Alaska. The frequency of white-flowered individuals increases with increasing growing-season temperature, consistent with the role of anthocyanin pigments in stress tolerance. White petals fail to produce the stress responsive flavonoid intermediates in the anthocyanin biosynthetic pathway (ABP), suggesting an early pathway blockage. Petal cDNA sequences did not reveal blockages in any of the eight enzyme-coding genes in white-flowered individuals, nor any color differentiating SNPs. A qRT-PCR analysis of white petals identified a 24-fold reduction in chalcone synthase (CHS) at the threshold of the ABP, but no change in CHS expression in leaves and sepals. This arctic species has avoided the deleterious effects associated with the loss of flavonoid intermediates in vegetative tissues by decoupling CHS expression in petals and leaves, yet the correlation of flower color and climate suggests that the loss of flavonoids in the petals alone may affect the tolerance of white-flowered individuals to colder environments.     

矮牵牛新种质的花色表型及花色素分析

以27个上海交通大学自育矮牵牛新种质为研究材料,对花色这一重要观赏性状及其花色素进行了系统研究。用RHSCC比色和色差仪测色方法描述了矮牵牛的花色表型,通过特征显色反应初步判断了矮牵牛的花色素类型,以标准曲线法和pH示差法等方法测定了矮牵牛3类花色素的含量。研究表明：这27个矮牵牛种质的花色可归于5个色系,以紫红色和红色为主;矮牵牛花色在CIELab表色系统中分布较广,而且不同色系花色参数的区分度较大。矮牵牛花瓣中含有类黄酮和花色苷,不含或含少量类胡萝卜素。13个被测种质的花瓣类黄酮含量在2.5～12.2 mg·/g–1 ·FW之间,花色苷含量在0.08～3.88 mg·g–1 FWmg/g·FW之间,而类胡萝卜素在矮牵牛花瓣中含量很低,远远低于类黄酮含量,在7个被测种质中,最高仅为0.216 mg·g–1 FWmg/g·FW,最低为0.004 mg·g–1 FWmg/g·FW。以上结果显示,5个色系矮牵牛所含花色素种类不尽相同,含量也有明显差异,其中紫红色系和红色系花瓣大多不含或含极少量类胡萝卜素,黄色系、白色系和紫色系花瓣的类黄酮含量较高,紫色系和紫红色系花瓣花色苷含量较高。     

Light induces petal color change in Quisqualis indica (Combretaceae)

Petal color change, a common phenomenon in angiosperms, is induced by various environmental and endogenous factors. Interestingly, this phenomenon is important for attracting pollinators and further reproductive success. Quisqualis indica L. (Combretaceae) is a tropical Asian climber that undergoes sequential petal color change from white to pink to red. This color changing process is thought to be a good strategy to attract more pollinators. However, the underlying physiological and biochemical mechanisms driving this petal color change phenomenon is still underexplored. In this context, we investigated whether changes in pH, pollination, light, temperature or ethylene mediate petal color change. We found that the detected changes in petal pH were not significant enough to induce color alterations. Additionally, pollination and temperatures of 20-30℃ did not alter the rate of petal color change; however, flowers did not open when exposed to constant temperatures at 15℃ or 35℃. Moreover, the application of ethylene inhibitor, i.e., silver thiosulphate, did not prevent color change. It is worth mentioning here that in our study we found light as a strong factor influencing the whole process of petal color change, as petals remained white under dark conditions. Altogether, the present study suggests that petal color change in Q. indica is induced by light and not by changes in petal pH, pollination, ethylene, or temperature, while extremely low or high temperatures affect flower anthesis. In summary, our findings represent the probable mechanism underlying the phenomenon of petal color change, which is important for understanding flower color evolution.     

Flower color alteration in <Emphasis Type="Italic">Lotus japonicus</Emphasis> by modification of the carotenoid biosynthetic pathway

To establish a model system for alteration of flower color by carotenoid pigments, we modified the carotenoid biosynthesis pathway of Lotus japonicus using overexpression of the crtW gene isolated from marine bacteria Agrobacterium aurantiacum and encoding β-carotene ketolase (4,4′-β-oxygenase) for the production of pink to red color ketocarotenoids. The crtW gene with the transit peptide sequence of the pea Rubisco small subunit under the regulation of the CaMV35S promoter was introduced to L. japonicus. In most of the resulting transgenic plants, the color of flower petals changed from original light yellow to deep yellow or orange while otherwise exhibiting normal phenotype. HPLC and TLC analyses revealed that leaves and flower petals of these plants accumulated novel carotenoids, believed to be ketocarotenoids consisting of including astaxanthin, adonixanthin, canthaxanthin and echinenone. Results indicated that modification of the carotenoid biosynthesis pathway is a means of altering flower color in ornamental crops.     

蒙农红豆草不同花色表型及色素成分特征分析

蒙农红豆草不仅是良好的饲草作物,还可以用作庭院观赏及蜜源植物。该研究以蒙农红豆草浅色花瓣突变体与对照群体中的粉红色、紫红色花瓣为试验材料,通过对花瓣颜色的表型和色素种类及含量的综合分析,明确影响花色形成的主要物质。结果表明:(1)蒙农红豆草浅色花突变体与对照的粉红色花和紫红色花为3种不同的色系,根据黄度(b*)和色相角(h°)将浅色花突变体的花色定义为黄白色花。(2)在3种花色中共检测到10种类黄酮和5种花青素,其中6种山奈酚衍生物、2种矮牵牛素衍生物、2种飞燕草素衍生物和1种锦葵素衍生物为首次在蒙农红豆草中报道;同时还发现山奈酚-3-芸香苷、山奈酚-3-葡萄糖苷和飞燕草素-3-羧基修饰芸香苷在3种花色中含量(36%~50%、21%~35%和27%~65%)最多。研究推测:芦丁、山奈酚-3-芸香苷-5-鼠李糖苷和山奈酚-3-p-香豆酰葡萄糖苷为影响蒙农红豆草花色变化的主要成分。     

Additional betalain accumulation by genetic engineering leads to a novel flower color in lisianthus (Eustoma grandiflorum)

Betalains, comprising violet betacyanins and yellow betaxanthins, are pigments found in plants belonging to the order Caryophyllales. In this study, we induced the accumulation of betalains in ornamental lisianthus (Eustoma grandiflorum) by genetic engineering. Three betalain biosynthetic genes encoding CYP76AD1, dihydroxyphenylalanine (DOPA) 4,5-dioxygenase (DOD), and cyclo-DOPA 5-O-glucosyltransferase (5GT) were expressed under the control of the cauliflower mosaic virus (CaMV) 35S promoter in lisianthus, in which anthocyanin pigments are responsible for the pink flower color. During the selection process on hygromycin-containing media, some shoots with red leaves were obtained. However, most red-colored shoots were suppressed root induction and incapable of further growth. Only clone #1 successfully acclimatized and bloomed, producing pinkish-red flowers, with a slightly greater intensity of red color than that in wild-type flowers. T₁ plants derived from clone #1 segregated into five typical flower color phenotypes: wine red, bright pink, pale pink, pale yellow, and salmon pink. Among these, line #1-1 showed high expression levels of all three transgenes and exhibited a novel wine-red flower color. In the flower petals of line #1-1, abundant betacyanins and low-level betaxanthins were coexistent with anthocyanins. In other lines, differences in the relative accumulation of betalain and anthocyanin pigments resulted in flower color variations, as described above. Thus, this study is the first to successfully produce novel flower color varieties in ornamental plants by controlling betalain accumulation through genetic engineering.     

Flower color change accelerated by bee pollination in Tibouchina (Melastomataceae)

Floral color changes are common among Melastomataceae and have been interpreted as a warning mechanism for bees to avoid old flowers, albeit increasing long-distance flower display. Here the reproductive systems of Tibouchina pulchra and T. sellowiana were investigated by controlled pollinations. Their pollinators were identified, and experiments on floral color and fragrance changes were conduced to verify if those changes affect the floral visitation. Both Tibouchina species are self compatible. The flowers lasted three days or more, and the floral color changed from white in the 1st day to pink in the following days. Pollen deposition on stigma induced floral color change. The effectiveness of the pollination is dependent on bees’ size; only large bees were regarded as effective pollinators. In experimental tests, the bees in T. pulchra preferred the natural white flowers while the visitors of T. sellowiana were attracted by both natural and mimetic 1st-day flowers (2nd-day flowers with experimentally attached 1st-day flower petals). During the experiments on floral fragrance, the bees visited both natural and mimetic 1st-day flowers (2nd-day flowers with 1st-day flower scents). In both experiments, the bees avoided natural 2nd-day flowers, but seldom visited modified 2nd-day flowers. The attractiveness of T. pulchra and T. sellowiana flowers cannot be attributed exclusively to the color or the fragrance separately, both factors seemingly act together.     

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。     

Geographic patterns in the reproductive ecology of Agave lechuguilla (Agavaceae) in the Chihuahuan desert. I. Floral characteristics, visitors, and fecundity

Floral characteristics such as morphology and flower color have been interpreted as adaptive traits that evolved through selective pressures generated by pollinators. Differences among populations in the expression of floral characters could result from natural selection for their adaptive value to local conditions. We describe the patterns of variation of flower morphology, color, and fecundity of Agave lechuguilla in 11 populations along a latitudinal gradient encompassing the whole range of the species in the Chihuahuan desert. We found a latitudinal pattern in flower shape and color. Flowers tended to be shorter, more open, and colorful toward the northern part of the gradient. We also recorded flower visitation, discriminating between pollinators and floral robbers. The main pollinators seems to be nocturnal hawk moths (Hyles lineata) and diurnal large bees (Bombus pennsylvanicus and Xylocopa californica). In all populations large bees were the most abundant potential pollinators. However, the abundance of the potential pollinators varied along the gradient. We observed no bat visits along the gradient. The number of visits by all potential pollinators decreased significantly with latitude as did fruit set.