天南星科植物的花多样性与传粉策略

天南星科植物具有特殊的佛焰苞花序及多样化的传粉策略, 是研究被子植物花的分化与动植物之间进化生态学联系的理想材料。本文简述了天南星科不同类型的花序结构及其传粉适应意义, 总结了天南星科传粉策略的基本类型与演化历史。天南星科的苞片结构主要包括原始型、外展平面型、直立宽佛焰苞型和直立狭佛焰苞4种类型, 呈现出从简单的片状与外展平面状结构向复杂的立体包裹状的佛焰苞结构演化的趋势。肉穗花序可分为两性花花序、单性花雌雄同序和单性花雌雄异序3种类型, 演化路线为两性花花序→单性花雌雄同序→单性花雌雄异序。天南星科的传粉者主要有鞘翅目、双翅目、膜翅目昆虫, 表现出5种主要传粉策略: 食物报酬型互利传粉、气味吸引型欺骗性传粉、交配场所型互利传粉、产卵场所型互利传粉和致死陷阱型欺骗性传粉。天南星科植物通过花序的形状、颜色、产热以及花部挥发物来吸引传粉者, 其中最主要的挥发物有二甲基硫化物、甲基吲哚化合物、萜类和苯类化合物, 模拟食物或产卵场所信号吸引鞘翅目甲虫和双翅目昆虫为其传粉。天南星科植物的佛焰苞被认为是促进该科物种分化的一个重要结构, 但该性状的演化历史及其与传粉系统分化之间的内在联系尚不明确。利用现代分子生物学技术以及模型模拟等手段, 结合生理生态学方法深入探究传粉事件与天南星科植物的花多样性以及物种分化之间的联系, 有望提升关于植物-传粉者互作与植物的花多样性分化之间关系的认识, 并丰富对被子植物多样性演化相关研究的理解。     

传粉网络的研究进展:网络的结构和动态

植物与传粉者之间相互作用,构成了复杂的传粉网络。近年来,社会网络分析技术的发展使得复杂生态网络的研究成为可能。从群落水平上研究植物与传粉者之间的互惠关系,为理解群落的结构和动态以及花部特征的演化提供了全新的视角。传粉网络的嵌套结构说明自然界的传粉服务存在冗余,而且是相对泛化的物种主导了传粉。在多年或者多季度的传粉网络中,虽然有很高的物种替换率,但是其网络结构仍然保持相对稳定,说明传粉网络对干扰有很强的抗性。尽管有关网络结构和动态的研究逐渐增多,但传粉网络维持的机制仍不清楚。网络结构可以部分由花部特征与传粉者的匹配来解释,也受到系统发生的制约,影响因素还包括群落构建的时间和物种多样性,以及物种在群落中的位置。开展大尺度群落动态的研究,为探索不同时间尺度、不同物种多样性水平上的传粉网络的生态学意义提供了条件。但已有的研究仍存在不足,比如基于访问观察的网络无法准确衡量传粉者的访问效率和植物间的花粉流动,以及结果受到调查精度区域研究不平衡的制约等。目前的研究只深入到传粉者携带花粉构成成分的水平,传粉者访问植物的网络不能代表植物的整个传粉过程。因此,研究应当更多地深入到物种之间关系对有性生殖的切实影响上。     

同域分布的紫堇属三种植物的繁育系统和传粉差异

为研究近缘物种之间繁育系统分化对传粉环境的适应性意义,本文针对湖北地区的3种紫堇属植物（紫堇Corydalis edulis Maxim.、尖距紫堇C.shearer S.Moore.和小花黄堇C.racemosa（Thunb.） Pers.）进行了传粉生态学研究,对比分析了它们在花部特征、分布模式、花期、交配系统、传粉系统等方面的差异。3种紫堇属植物常见伴生现象,花期有一定重叠;花色、距长、花蜜量等方面均有显著差异;尖距紫堇交配系统为自交不亲和、依靠传粉者异花授粉;而紫堇和小花黄堇交配系统为自交亲和,兼有自交和异交的混合交配系统。3个物种均由蜂类传粉,且花部性器官与传粉者的接触部位一致,但访花频率差异较大。在混合群落中,传粉者访花具有较高的忠实性,花部特征差异可能是传粉者选择性访花的原因。对于紫堇属3个物种,花部特征和交配系统的分化有助于其避免传粉过程的相互干扰,交配系统与传粉环境具有一定相关性,混合交配系统可能有利于提高植物对不同生境和气候的适应性。     

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。     

昆虫植食对植物有性生殖的影响

植物在个体发育的各个阶段都与不同的群落成员相互作用,如竞争的植物、有益的传粉者和敌对的植食动物。昆虫植食在各类生态系统中普遍存在,并可能对植物有性生殖产生各种影响。植食昆虫可通过对植物有性生殖结构的消耗直接对植物生殖产生影响,也可通过影响植物资源分配和花性状等改变传粉者服务,从而间接对植物有性生殖带来正面、负面或中性的影响。同一植物的植食昆虫和传粉者往往对植物的吸引性状 （如花大小、气味、颜色等）有相同的偏好,因此植食者与传粉者均能对植物有性生殖性状施加选择压力。本文从昆虫植食对植物有性生殖的直接影响、间接影响以及植食昆虫对植物有性生殖性状选择的影响三个方面进行综述,以期为昆虫植食和生物资源多样性保护相关研究提供参考。     

管花秦艽的传粉生态学研究——兼与同域分布近缘种的比较

在青藏高原东北部连续两年观察了晚秋开花植物管花秦艽Gentiana siphonantha的传粉生态学特征,并在此基础上进一步比较分析了与该物种同域分布且亲缘关系较近、但开花较早的麻花艽G straminea之间的传粉生态学特征.管花秦艽的花发育过程表现出雌雄异熟和雌雄异位的特点,不存在花内的自花传粉,套袋隔离的花不结实也支持这一结论;株内自交的高结实率表明该物种是自交亲和的.盛花期每植株平均有15朵开放的花,雄性和雌性阶段的花比例为1.2:1;自然条件下产生种子必须依赖传粉媒介;苏氏熊蜂是最有效的传粉昆虫,且访花过程中埘雄性和雌性阶段花不具明显的偏向性;株内连续访花的频率高达87.8%,从而导致同株异花传粉自交的广泛存在.与同域分布的麻花艽相比,管花秦艽的单花花期、雄性和雌性期持续时间缩短.但盛花期开花数量明显增加.令人感兴趣的是尽管两个近缘种的花形态特征存在显著差异,但都是由同一种熊蜂传粉.这一特点与过去认为花颜色和花管长度是物种分化过程中与不同传粉昆虫协同进化导致牛殖隔离的假说不相符合.管花秦艽单花的访花频率和同株异花连续访花的比例都明显高于麻花艽.两个物种不同花序设计导致访花昆虫行为的改变可能是造成这一差异的主要原因.两个物种具有不同的开花时间,但仍然存在一定的花期重叠,表现出不完全的传粉生殖隔离状态.     

自交与异交象牙参属植物的蜜腺结构及其生物学意义

蜜腺是有花植物与传粉昆虫构建互惠关系的关键花部结构,解析不同繁殖特性物种间蜜腺结构的差异可为理解有花植物繁殖特性的演化提供理论依据。以传粉系统高度特化、异交实现有性生殖的早花象牙参（Roscoea cautleoides）和缺乏有效传粉者、主动自交实现有性生殖的无柄象牙参（R. schneideriana）为材料,通过野外测量2个物种蜜腺特性、超景深显微镜观察和石蜡切片染色,探究早花象牙参与无柄象牙参在蜜腺分布位置、外部形态及内部结构的异同。结果表明：2个物种均具有结构蜜腺,从外部形态来看,早花象牙参蜜腺体积较大、分泌糖浓度较高的花蜜,而无柄象牙参蜜腺体积较小、无花蜜分泌;从内部结构来看,早花象牙参蜜腺结构特化、各个组成部分有明显的区分,而无柄象牙参蜜腺出现一定程度的退化。该研究结果为揭示早花象牙参与无柄象牙参在花部特征、传粉机制及繁育系统的差异以及无柄象牙参蜜腺无花蜜分泌的可能机理提供了重要的形态学依据,也为深入地探讨植物与传粉动物间的协同进化关系以及理解蜜腺多样性的起源与维持机制奠定了科学基础。     

植物气味腺研究进展

气味腺是花器官上能够产生和释放花气味的特殊腺体结构,在传粉过程中与其他花部性状结合能够吸引传粉者访花,气味腺的研究有助于揭示动物与植物之间的协同进化机制,此外,气味腺外部形态特征及细胞微形态可作为分类依据之一。对气味腺的结构、类型和检测方法,及其在植物科属中的分布情况进行了归纳总结,并对气味腺在传粉过程中和分类学上的意义进行了分析。最后提出只有结合分子实验技术手段,全面综合考虑繁殖生物学和植物化学的分析方法,才能深入理解气味腺的多样性与演化。     

滇西北特有植物蓝果杜鹃的花色多态性研究(英文)

蓝果杜鹃(Rhododendron cyanocarpum)为大理苍山特有的濒危植物,有粉色和白色两种花冠类型。为了探讨该物种花色多态性的意义,本研究调查了粉色花和白色花植株在已知的各居群的分布频率、花冠的反射光谱及其它的花部特征、有效传粉者及其访花频率与结实情况。结果表明:粉色花植株在所有调查的居群中占优势(77%~100%)。粉色花的花冠反射光谱在430 nm和650 nm有两个峰,而白色花只在430 nm有一个反射峰。同时,花特征如:花柱与柱头颜色、花冠长度、花萼长度、花梗长度以及雌雄蕊最短距离,两种花冠存在显著差异。另外,尽管熊蜂作为这两种花冠的主要传粉者,但粉红花的访花频率以及自然条件下的结实情况显著高于白色花。本研究结果推测粉红色花可能受到了稳定性选择的作用。     

滇西北特有植物蓝果杜鹃的花色多态性研究?

蓝果杜鹃( Rhododendron cyanocarpum)为大理苍山特有的濒危植物,有粉色和白色两种花冠类型。为了探讨该物种花色多态性的意义,本研究调查了粉色花和白色花植株在已知的各居群的分布频率、花冠的反射光谱及其它的花部特征、有效传粉者及其访花频率与结实情况。结果表明：粉色花植株在所有调查的居群中占优势(77％~100％)。粉色花的花冠反射光谱在430 nm和650 nm有两个峰,而白色花只在430 nm有一个反射峰。同时,花特征如：花柱与柱头颜色、花冠长度、花萼长度、花梗长度以及雌雄蕊最短距离,两种花冠存在显著差异。另外,尽管熊蜂作为这两种花冠的主要传粉者,但粉红花的访花频率以及自然条件下的结实情况显著高于白色花。本研究结果推测粉红色花可能受到了稳定性选择的作用。     

COMPETITION FOR HUMMINGBIRD POLLINATION SHAPES FLOWER COLOR VARIATION IN ANDEAN SOLANACEAE

One classic explanation for the remarkable diversity of flower colors across angiosperms involves evolutionary shifts among different types of pollinators with different color preferences. However, the pollinator shift model fails to account for the many examples of color variation within clades that share the same pollination system. An alternate explanation is the competition model, which suggests that color divergence evolves in response to interspecific competition for pollinators, as a means to decrease interspecific pollinator movements. This model predicts color overdispersion within communities relative to null assemblages. Here, we combine morphometric analyses, field surveys, and models of pollinator vision with a species‐level phylogeny to test the competition model in the primarily hummingbird‐pollinated clade Iochrominae (Solanaceae). Results show that flower color as perceived by pollinators is significantly overdispersed within sites. This pattern is not simply due to phylogenetic history: phylogenetic community structure does not deviate from random expectations, and flower color lacks phylogenetic signal. Moreover, taxa that occur in sympatry occupy a significantly larger volume of color space than those in allopatry, supporting the hypothesis that competition in sympatry drove the evolution of novel colors. We suggest that competition among close relatives may commonly underlie floral divergence, especially in species‐rich habitats where congeners frequently co‐occur.     

Variation in context‐dependent foraging behavior across pollinators

Pollinator foraging behavior has direct consequences for plant reproduction and has been implicated in driving floral trait evolution. Exploring the degree to which pollinators exhibit flexibility in foraging behavior will add to a mechanistic understanding of how pollinators can impose selection on plant traits. Although plants have evolved suites of floral traits to attract pollinators, flower color is a particularly important aspect of the floral display. Some pollinators show strong innate color preference, but many pollinators display flexibility in preference due to learning associations between rewards and color, or due to variable perception of color in different environments or plant communities. This study examines the flexibility in flower color preference of two groups of native butterfly pollinators under natural field conditions. We find that pipevine swallowtails (Battus philenor) and skippers (family Hesperiidae), the predominate pollinators of the two native Texas Phlox species, Phlox cuspidata and Phlox drummondii, display distinct patterns of color preferences across different contexts. Pipevine swallowtails exhibit highly flexible color preferences and likely utilize other floral traits to make foraging decisions. In contrast, skippers have consistent color preferences and likely use flower color as a primary cue for foraging. As a result of this variation in color preference flexibility, the two pollinator groups impose concordant selection on flower color in some contexts but discordant selection in other contexts. This variability could have profound implications for how flower traits respond to pollinator‐mediated selection. Our findings suggest that studying dynamics of behavior in natural field conditions is important for understanding plant–pollinator interactions.     

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.     

Pollinator response to flower color polymorphism and floral display in a plant with a single-locus floral color polymorphism: consequences for plant reproduction

Variation in flower color, particularly polymorphism, in which two or more different flower color phenotypes occur in the same population or species, may be affected or maintained by mechanisms that depend on pollinators. Furthermore, variation in floral display may affect pollinator response and plant reproductive success through changes in pollinator visitation and availability of compatible pollen. To asses if flower color polymorphism and floral display influences pollinator preferences and movements within and among plants and fitness-related variables we used the self-incompatible species Cosmos bipinnatus Cav. (Asteraceae), a model system with single-locus flower color polymorphism that comprises three morphs: white (recessive homozygous), pink (heterozygous co-dominate), and purple (dominant homozygous) flowers. We measured the preferences of pollinators for each morph and constancy index for each pollinator species, pollination visitation rate, floral traits, and female fitness measures. Flower color morphs differed in floral trait measures and seed production. Pollinators foraged nonrandomly with respect to flower color. The most frequent morph, the pink morph, was the most visited and pollinators exhibited the highest constancy for this morph. Moreover, this morph exhibited the highest female fitness. Pollinators responded strongly to floral display size, while probed more capitulums from plants with large total display sizes, they left a great proportion of them unvisited. Furthermore, total pollinator visitation showed a positive relation with female fitness. Results suggest that although pollinators preferred the heterozygous morph, they alternate indiscriminately among morphs making this polymorphism stable.     

POLLINATION OF ANGIOSPERMS IN CONTRASTING CONIFEROUS FORESTS

We studied the distributions of flower color, flower morphology, and putative pollinators in eight communities in the western Cascades of Washington and 14 communities in the eastern Cascades. These two forested regions differ in the proportions of species in each flower color and morphological type such that the eastern region is more complex. The putative pollinators of each species were determined from morphological and color properties of the flowers and from the recent, extensive literature. The floristic complement of the western region appears to be serviced by a collection of pollinators that is less diverse than that of the east. In each region there is a strong positive correlation between species richness and flower color diversity, floral morphology diversity, and putative pollinator diversity, respectively. These comparisons suggest the competition for pollinators has led to greater niche differentiation in the eastern regions and that this factor, combined with greater habitat complexity, results in the far greater species richness of the eastern region.     

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素。花瓣彩斑主要由基因突变或病毒入侵而形成。     

The birds, the bees, and the virtual flowers: can pollinator behavior drive ecological speciation in flowering plants?

Biologists have long assumed that pollinator behavior is an important force in angiosperm speciation, yet there is surprisingly little direct evidence that floral preferences in pollinators can drive floral divergence and the evolution of reproductive (ethological) isolation between incipient plant species. In this study, we expose computer-generated plant populations with a wide variation in flower color to selection by live and virtual hummingbirds and bumblebees and track evolutionary changes in flower color over multiple generations. Flower color, which was derived from the known genetic architecture and phenotypic variance of naturally occurring plant species pollinated by both groups, evolved in simulations through a genetic algorithm in which pollinator preference determined changes in flower color between generations. The observed preferences of live hummingbirds and bumblebees were strong enough to cause adaptive divergence in flower color between plant populations but did not lead to ethological isolation. However, stronger preferences assigned to virtual pollinators in sympatric and allopatric scenarios rapidly produced ethological isolation. Pollinators can thus drive ecological speciation in flowering plants, but more rigorous and comprehensive behavioral studies are required to specify conditions that produce sufficient preference levels in pollinators.     

Can indirect selection and genetic context contribute to trait diversification? A transition‐probability study of blossom‐colour evolution in two genera

Darwin recognized that biological diversity has accumulated as a result of both adaptive and nonadaptive processes. Very few studies, however, have addressed explicitly the contribution of nonadaptive processes to evolutionary diversification, and no general procedures have been established for distinguishing between adaptive and nonadaptive processes as sources of trait diversity. I use the diversification of flower colour as a model system for attempting to identify adaptive and nonadaptive causes of trait diversification. It is widely accepted that variation in flower colour reflects direct, adaptive response to divergent selective pressures generated by different pollinators. However, diversification of flower colour may also result from the effects of nonadaptive, pleiotropic relationships with vegetative traits. Floral pigments that have pleiotropic relationships to vegetative pigments may evolve and diversify in at least two nonadaptive ways. (1) Indirect response to selection on the pleiotropically related nonfloral traits may occur (indirect selection). (2) Divergent evolution in response to parallel selective pressures (e.g. selection by pollinators for visually obvious flowers) may occur because populations are at different genetic starting points, and each population follows its own genetic `line of least resistance.' A survey of literature suggests that pleiotropic relationships between flower colour and vegetative traits are common. Phylogenetically informed analyses of comparative data from Dalechampia (Euphorbiaceae) and Acer (Aceraceae), based on trait‐transition probabilities and maximum likelihood, indicated that floral and vegetative pigments are probably pleiotropically related in these genera, and this relationship better explains the diversification of floral colour than does direct selection by pollinators. In Dalechampia pink/purple floral bract colour may have originated by indirect response to selection on stem and leaf pigments. In Acer selection by pollinators for visually obvious flowers may to have led to the evolution of red or purple flowers in lineages synthesizing and deploying red anthocyanins in leaves, and pale‐green or yellow flowers in species not deploying red anthocyanins in vegetative structures. This study illustrates the broader potential of indirect selection and parallel selection on different genetic starting points to contribute to biological diversity, and the value of testing directly for the operation of these nonadaptive diversifying processes.     

Flower color microevolution in wild radish: evolutionary response to pollinator-mediated selection

Evolutionary ecologists are fundamentally interested in how species interactions affect evolutionary change. We tested the degree to which plant-pollinator interactions affect the frequency of flower color morphs of Raphanus sativus. Petal color in R. sativus is determined by two independently assorting loci, producing four petal colors (yellow, white, pink, and bronze). We assessed the impact of pollinator discrimination on changes in flower color variation by comparing the frequency of colors produced in the presence (open pollination) versus absence (null pollination) of pollinator discrimination. We also assessed the impact of postpollination and developmental effects on progeny colors using equal pollinations with all four color morphs. Our results from open pollinations found an overrepresentation of yellow progeny in the next generation, when compared with both null pollinations and cumulative ratios based on Hardy-Weinberg and linkage equilibria assumptions. When these results were combined with those from equal pollinations, the overrepresentation of yellow could be attributed to selection from pollinators. Yet, surveys in the field the following year found no flower color frequency changes in the next generation. These results illustrate that flower color microevolution can be driven by both pollinator discrimination and other nonpollinator selective forces acting during the seed-to-adult transition, countering selection imposed by pollinators.     

The genetic dissection of floral pollination syndromes

A major factor in the evolution of the angiosperms is the adaptation of plants to animal pollinators. The specific morphology of a flower, its color, nectar composition and scent production can all contribute to reproductive success by attracting pollinators and by limiting out-crossing with other species. It has now become feasible to dissect the genetic basis of plant adaptation to different pollinators.