台闽苣苔（苦苣苔科）花部器官的形态发生

在扫描电镜下对台闽苣苔 (T .oldhamii (Hemsl.)Solereder)进行了花部器官形态发生的观察 ,为探索该类群的个体发育、类群间的系统发育关系和进化趋势提供依据。研究发现该属植物萼片、花冠和雄蕊发生式样均为五数花类型 ,它们各自来源于花原基上分化出来的萼片原基、花冠原基和雄蕊原基 ;花冠与雄蕊的两侧对称性与花冠上唇生长稍快和退化雄蕊原基发育迟滞相关 ;萼片原基的发生和发育的顺序是不一致的 :萼片原基发生的式样为近轴中原基—远轴 2原基— 2侧原基 ,发育式样则为近轴中萼片— 2侧萼片—远轴 2萼片 ,花蕾时为镊合状排列。花冠裂片原基的发生和发育式样是一致的 ,即远轴中裂原基 (下唇中裂片 )—远轴 2侧裂原基 (下唇 2侧裂片 )—近轴 2裂原基 (上唇 2裂片 )。花蕾期卷迭式为覆瓦状排列 ,从外向内 :下唇中裂片—下唇 2侧裂片—上唇 2裂片或下唇 2侧裂片—上唇 2裂片—下唇中裂片。雄蕊原基与花冠裂片原基互生 ,前方雄蕊原基在发生上稍迟于后方雄蕊原基 ,后者与退化雄蕊原基几乎同时发生 ,但较小 ,并与近轴心皮 (或柱头上唇 )对生。将该属与玄参科 (Scrophulari aceae)的地黄属 (Rehmannia)、苦苣苔科 (Gesneriaceae)的异叶苣苔属 (Whytockia)和尖舌苣苔属 (Rhynchoglossum)的花部器官比较发现     

台闽苣苔(苦苣苔科)花部器官的形态发生

在扫描电镜下对台闽苣苔 (T. oldhamii (Hemsl.) Solereder)进行了花部器官形态发生的观察,为探索该类群的个体发育、类群间的系统发育关系和进化趋势提供依据.研究发现该属植物萼片、花冠和雄蕊发生式样均为五数花类型,它们各自来源于花原基上分化出来的萼片原基、花冠原基和雄蕊原基;花冠与雄蕊的两侧对称性与花冠上唇生长稍快和退化雄蕊原基发育迟滞相关;萼片原基的发生和发育的顺序是不一致的:萼片原基发生的式样为近轴中原基-远轴2原基-2侧原基,发育式样则为近轴中萼片-2侧萼片-远轴2萼片,花蕾时为镊合状排列.花冠裂片原基的发生和发育式样是一致的,即远轴中裂原基(下唇中裂片)-远轴2侧裂原基(下唇2侧裂片)-近轴2裂原基(上唇2裂片).花蕾期卷迭式为覆瓦状排列,从外向内:下唇中裂片-下唇2侧裂片-上唇2裂片或下唇2侧裂片-上唇2裂片-下唇中裂片.雄蕊原基与花冠裂片原基互生,前方雄蕊原基在发生上稍迟于后方雄蕊原基,后者与退化雄蕊原基几乎同时发生,但较小,并与近轴心皮(或柱头上唇)对生.将该属与玄参科(Scrophulariaceae)的地黄属( Rehmannia )、苦苣苔科(Gesneriaceae)的异叶苣苔属( Whytockia)和尖舌苣苔属(Rhynchoglossum )的花部器官比较发现,这四个属在这方面呈现出多样性和交叉.过去一直按子房室数和胎座类型划分玄参科(子房2室、中轴胎座)和苦苣苔科(子房1室、侧膜胎座)这一做法受到了质疑.     

广西的苦苣苔科植物

位于我国南部边题的广西壮族自治区,自然条件十分优越,物产丰富,动植物种类繁多,仅维管束植物就多达8354种,仅次于云南和四川而位居全国第三位,其中有许多珍稀、奇异和特有的种类。苦苣苔科就是这一大家庭中的一个新成员。说它是新成员,主要是指广西的诊科植物有60％以上的种类都是在八十年代以后才发现的新种,而且现在每年都还有不少新种陆续发表,其比例超过了任何一科的植物。对于这些刚露面不久的“山野隐士”,人们普遍感到陌生和好奇。一、种类十分丰富,特有现象极为突出苦芭苔科可分为两个亚科即苦芭苔亚科和大岩桐亚科,苦…     

中国苦苣苔科的研究(十)

<正> 多年生草本。根状茎,粗。叶约10枚,均基生,具长或短柄;叶片纸质,卵形或卵状菱形,长2—3.4厘米,宽1.6—2.9厘米,顶端钝或圆形,基部宽楔形或圆形,边缘有钝重牙齿,上面多少泡状,在细脉形成的小空间中密被短柔毛,下面脉上密被白色短柔毛,在隆起的中脉和侧脉上还被暗褐色长柔毛,侧脉每侧4—5条;叶柄长1—3厘米,密被暗褐色长     

石山苣苔属(苦苣苔科)花形态演化及分类学意义

石山苣苔属(苦苣苔科)约30种,主要分布于我国南部的石灰岩地区.目前该属已知物种数虽少但花形态极其多样,是该科中分类最为困难的类群之一.基于分子证据,其它8个属中花形态迥异的一些物种被并入石山苣苔属.然而,该属花形态的演化趋势缺乏系统性的研究,传统分类对属的界定与分子系统学研究结果相矛盾的原因,以及是否有形态特征支持新界定的石山苣苔属还不清楚.该研究中,总共编码了19种石山苣苔属植物和9种报春苣苔属植物的35个形态特征,其中包括26个花部形态特征,在分子系统树上追踪了它们的演化路径.结果表明:无论属内还是属间,多数花部形态特征,尤其以往属的分类界定特征,在演化过程中变化频繁且发生了高度同塑性演化,这是导致传统形态分类不自然的关键因素.此外,在观察研究的所有特征中,花丝和柱头的差异可能在石山苣苔属植物共同祖先中经历了演变,或可用于区分石山苣苔属与其姐妹报春苣苔属的大多数种类.因此,在苦苣苔科植物的分类学研究中应当慎用这些花部性状作为分类依据,而且应对形态特征进行广泛地观察研究,在密集的取样和分辨率更高、更可靠的系统树上追踪它们的演化规律.更为重要的是,需要进一步研究导致复杂形态性状演化的内在分子调控机理和外在的自然选择动力,最终更加深入地理解石山苣苔属等典型喀斯特植物的演化过程和机理.     

苦苣苔科吊石苣苔属一新异名

2017年,印度学者Joe等基于采自中国藏东南地区的植物标本,发表了合萼吊石苣苔的一新变种Lysionotus gamosepalus W. T. Wang var.biflorus A. Joe, Hareesh&M. Sabu,并且认为该新变种与原变种的主要区分特征包括叶缘稍具锯齿、每个腋生花序仅有2朵花、花冠被毛、具2枚卷曲的退化雄蕊。然而,近年来,笔者基于查阅大量标本和野外考察工作,认为所谓变种的形态特征是在合萼吊石苣苔(Lysionotus gamosepalus W. T. Wang)的变异范围之内,加之分布区完全重叠,生长环境类似,物候期一致,不宜作为变种等级处理。故在此将该变种处理为合萼吊石苣苔的异名。     

The pollination ecology of Paraboea rufescens (Gesneriaceae): a buzz-pollinated tropical herb with mirror-image flowers

BACKGROUND AND AIMS: Gesneriaceae is a pantropical plant family with over 3000 species. A great variety of pollination mechanisms have been reported for the neotropical members of the family, but the details of buzz-pollination and enantiostyly for the family have not been described. We investigated the floral biology and pollination ecology of Paraboea rufescens in Xishuangbanna, south-west China, considering three aspects: (1) the type of enantiostyly exhibited; (2) whether the species is self-compatible; and (3) whether pollinator behaviour could enhance the precision of pollen transfer between flowers of contrasting stylar orientation. METHODS: Flowering phenology was monitored once a month during vegetative growth, and once a week during flowering both in the field and under cultivation. Pollination manipulations and pollinator observation in the field were conducted. KEY RESULTS: Anthesis occurred early during the morning, and flowers remained open for 1-5 d, depending on weather conditions. Controlled pollinations revealed that P. rufescens is self-compatible, and exhibited inbreeding depression in seed set. Plants were pollinator limited in natural populations. The similar stylar deflection among flowers within a plant limits autonomous self-pollination as well as pollination between flowers. Two species of bumble bees (Bombus spp.), Amegila malaccensis and Nomia sp. effectively pollinated P. rufescens. These pollinators visited flowers in search of pollen with almost the same frequency. None of the pollinators appeared to discriminate between left- or right-handed flowers. CONCLUSIONS: Paraboea rufescens exhibits monomorphic enantiostylous flowers and a buzz-pollination syndrome. Floral morphology in P. rufescens and pollinator foraging behaviour seems likely to reduce self-pollination and pollinations between flowers of the same stylar deflection.     

圆唇苣苔属(Gyrocheilos)花柱侧偏弯折现象及其传粉适应机制

圆唇苣苔属(Gyrocheilos)是苦苣苔科的中国特有属,有5种,全部狭域分布在我国西南及广东的高海拔山区。圆唇苣苔属所有物种的花柱侧偏且花柱顶端呈90°弯折,使得柱头位于花开口的中央位置。这种独特的侧偏弯折花柱结构,说明圆唇苣苔属可能有着特殊的演化历史和适应机制。为揭示这种特殊的花柱侧偏弯折现象的发生范围、发育过程及其传粉适应机制,该研究在圆唇苣苔(Gyrocheilos chorisepalus)、折毛圆唇苣苔(G. retrotrichus)和微毛圆唇苣苔(G. microtrichus)3个物种中开展了花部综合征观察,并研究了广东大雾岭保护区内的折毛圆唇苣苔花发育过程、花部特征和繁育系统以及传粉过程。结果表明:(1)微毛圆唇苣苔只有花柱左偏弯折现象,而圆唇苣苔和折毛圆唇苣苔虽然大部分花是花柱左偏弯折,但在部分个体中出现了少量的花柱右偏弯折现象(占种群总花数的2%～3%)。(2)传粉观察发现,折毛圆唇苣苔在花蕾期即出现了花柱弯折现象,2个可育雄蕊的花药合生、位于花冠筒喉部中央位置,与侧偏花柱不存在左右镜像对称关系。(3)折毛圆唇苣苔的花粉胚珠比(P/O)为456.98±15.55,属于兼性异交繁育系统。折毛圆唇苣苔存在一定的传粉限制,自交授粉可以结实,但异交种子萌发率更高,可能存在近交衰退。(4)折毛圆唇苣苔的访花昆虫较少,访花频率较低,主要访花昆虫有隧蜂、熊蜂、食蚜蝇等; 熊蜂体型较大,访花时降落在弯折花柱和花瓣下唇,胸部侧面及下部能有效接触到柱头。(5)反射率结果显示,折毛圆唇苣苔花瓣反射波长范围集中在紫光和蓝紫光区域,花冠的反射波长范围与蜂类视觉范围一致且花冠筒外侧和花瓣下唇的反射强度最大,更容易吸引蜂类落置在花冠宽大的下唇; 圆唇苣苔属的花柱侧偏弯折现象可能来自近缘的长蒴苣苔属(Didymocarpus)的花柱下弯现象或镜像花(mirror-image flowers)。综上认为,这种侧偏弯折的花柱,可能通过提供昆虫降落平台,使得柱头位于花开口中央和花瓣下唇的上方位置,提高了柱头接触访花昆虫的概率,是适应高海拔地区低频率访花者的一种机制。     

中国苦苣苔科植物的多样性与地理分布（附表）

物种多样性编目是开展生物多样性保护的重要基础,该研究结合最新分子系统学研究成果以及近年来发表的新资料,对中国苦苣苔科植物多样性和地理分布数据进行了统计和分析.结果表明:中国苦苣苔科植物共有44属671种(含种下单位,下同),其中特有属11个;特有种573种,占总种数的85.39%.种数最多的10个属依次为广义报春苣苔属(180种)、广义马铃苣苔属(122种)、石蝴蝶属(39种)、半蒴苣苔属(39种)、芒毛苣苔属(38种)、长蒴苣苔属(35种)、石山苣苔属(31种)、吊石苣苔属(31种)、蛛毛苣苔属(28种)、汉克苣苔属(22种).在地理分布上,种数排名前10的省份(区)有广西(260种,33属)、云南(236种,30属)、贵州(96种,28属)、广东(93种,17属)、四川(85种,21属)、湖南(58种,13属)、西藏(39种,9属)、湖北(29种,15属)、福建(26种,13属)、江西(25种,9属).含中国特有苦苣苔科植物的属中排前10位的分别为广义报春苣苔属(178种)、广义马铃苣苔属(119种)、石蝴蝶属(37种)、半蒴苣苔属(35种)、石山苣苔属(30种)、长蒴苣苔属(29种)、吊石苣苔属(23种)、蛛毛苣苔属(19种)、芒毛苣苔属(19种)、汉克苣苔属(11种).这表明中国南部和西南部是苦苣苔科植物的一个分布中心,特别是石灰岩地区有着高度的物种多样性和特有性,广义报春苣苔属、广义马铃苣苔属、石蝴蝶属、半蒴苣苔属、石山苣苔属、吊石苣苔属等为我国典型的优势属.此外,根据目前的研究现状,还对我国苦苣苔科植物资源的调查、分类学和系统发育研究、保护和可持续利用等进行了讨论.     

Nectarless flowers: ecological correlates and evolutionary stability

In animal-pollinated flowers, the pollinators cannot detect the presence of nectar before entering flowers, and therefore flowers may cheat by not producing nectar. An earlier model suggested that a mixed strategy of producing nectarful and nectarless flowers would be evolutionarily stable. Here we compare nectarless flowers as a cheating strategy with three competing hypotheses namely "visit-more-flowers", "cross-pollination enhancement" and "better contact". We collected field data on 28 species of plants to test some of the differential predictions of the hypotheses. Nectarless flowers were detected in 24 out of 28 plant species. Correlations of percent nectarless flowers with floral and ecological variables support the cheater flower hypothesis. We further model the cost-benefits of cheating and show that an evolutionary stable ratio of nectarless to nectarful flowers can be reached. The equilibrium ratio is mainly decided by factors associated with pollinator density and pollinator learning.     

Evolution of funnel-revolver flowers and ornithophily in nasa (loasaceae)

Floral morphology, distribution, and flower visitors for 60 taxa of Nasa are investigated and compared to molecular trees inferred both from a combined marker analysis (ITS1 and trnL (UAA)) and from a single marker (ITS1). Flowers conform to two different floral types: Firstly, "tilt-revolver flowers", with spreading to reflexed, white to yellow petals and small, brightly coloured floral scales contrasting with the petals and firmly enclosing the nectar (Saccatae and Carunculatae); secondly, "funnel-revolver flowers", with half-erect to erect, orange to red petals and floral scales not contrasting with the petals, or enclosed in the corolla, and nectar freely accessible by funnel-shaped floral scales ( Alatae, Grandiflorae, and N. venezuelensis species group). Phylogenetic analysis shows that "tilt-revolver flowers" represent the plesiomorphic condition by outgroup comparison. The two groups with tilt-revolver flowers in Nasa are not monophyletic ( Saccatae are paraphyletic, Carunculatae are polyphyletic). Most Saccatae fall into two monophyletic assemblages, the N. poissoniana species group and the N. triphylla species group. The remainder of Saccatae group either with Grandiflorae ( N. insignis species group) or with Alatae ( N. laxa species group). The clades retrieved in the molecular analysis contradict the traditional classification, but are congruent with vegetative morphology, details of the flower morphology, and biogeography. "Funnel-revolver flowers" represent the derived condition, but molecular data suggest a convergent development (at least twice independently), since the corresponding species do not constitute a monophyletic group. "Tilt-revolver flowers" are visited and pollinated by bees (especially Colletidae), whereas "funnel-revolver flowers" are mostly visited by hummingbirds. The transition from melittophily to ornithophily may have been the license for the colonization of, and the diversification in, both cloud forest and high Andean habitats.     

Reversal versus specialization in floral morphological evolution in Petrocosmea (Gesneriaceae)

A widely held hypothesis in evolution is that adaptive specialization constrains the potential direction of future evolutionary change and thus may be irreversible, also known as Dollo's law. However, this hypothesis has long been subject to debate in evolutionary biology. Floral specialization is intriguing as it is usually linked to reproductive isolation and could affect speciation. Here, following the discovery of four new taxa, we observed some interesting phenomena of reversal versus specialization in morphology in a clade with the most specialized flowers in the genus Petrocosmea. In the phylogenetic tree based on sequences of multiple DNA regions, the morphological reversals, especially the regain of a long corolla tube, are nested within the branches characteristic of normally specialized flowers with a short corolla tube and highly specialized zygomorphy. Our results indicate that the highly specialized floral organ of this clade is still actively evolving in multiple branches toward specialization while reversals to different ancestral states occur in some branches. Great disturbance of ecological environment is likely a crucial factor affecting trait reversibility, such as the rapid uplift of the Himalayan–Tibetan plateau. The four new taxa are treated herein taxonomically. The flowers of this clade represent an interesting model to explore the genetic basis underlying the evolutionary reversal versus specialization and the interplay between genetic factors and environmental variables.     

A theoretical investigation of the evolution and maintenance of mirror-image flowers

Enantiostyly is the deflection of the female sex organ either to the left or to the right side of the floral axis, resulting in mirror-image flowers. Two types of enantiostyly occur: in monomorphic enantiostyly, individuals exhibit both flower forms, whereas in dimorphic enantiostyly, the forms occur on separate plants. Monomorphic enantiostyly is known from at least 10 families, whereas dimorphic enantiostyly is reported in only three. Phylogenetic evidence suggests that monomorphic enantiostyly has evolved from a straight-styled ancestor and that dimorphic enantiostyly is derived from monomorphic enantiostyly. Here, we use theoretical models to investigate the role of pollen transfer in influencing these evolutionary transitions. We used numerical calculations to examine the evolution of monomorphic and dimorphic enantiostyly under different conditions of pollen transfer, inbreeding depression, and pollinator visitation. Our results demonstrate that in comparison to a putative straight-styled ancestor, both monomorphic and dimorphic enantiostyly function to reduce geitonogamous pollen transfer with a concomitant increase in pollen export. Our calculations suggest that the first stage in the evolution of monomorphic enantiostyly involves the deflection of the style only, followed by selection for reciprocity in anther position to promote more precise cross-pollination. Constraints associated with the developmental genetics of left-right asymmetries may account for the low number of evolutionary transitions from monomorphic to dimorphic enantiostyly, despite the evolutionary stability of this condition once it arises.     

Darwin's legacy: the forms,function and sexual diversity of flowers

Charles Darwin studied floral biology for over 40 years and wrote three major books on plant reproduction. These works have provided the conceptual foundation for understanding floral adaptations that promote cross-fertilization and the mechanisms responsible for evolutionary transitions in reproductive systems. Many of Darwin''s insights, gained from careful observations and experiments on diverse angiosperm species, remain remarkably durable today and have stimulated much current research on floral function and the evolution of mating systems. Here I review Darwin''s seminal contributions to reproductive biology and provide an overview of the current status of research on several of the main topics to which he devoted considerable effort, including the consequences to fitness of cross- versus self-fertilization, the evolution and function of stylar polymorphisms, the adaptive significance of heteranthery, the origins of dioecy and related gender polymorphisms, and the transition from animal pollination to wind pollination. Post-Darwinian perspectives on floral function now recognize the importance of pollen dispersal and male outcrossed siring success in shaping floral adaptation. This has helped to link work on pollination biology and mating systems, two subfields of reproductive biology that remained largely isolated during much of the twentieth century despite Darwin''s efforts towards integration.     

Non-Euglossine bees also function as pollinators of Sinningia species (Gesneriaceae) in southeastern brazil

Pollination by male and female Euglossini bees, euglossophily, was suggested for a number of neotropical Gesneriaceae species. Information on bee species other than Euglossini as pollinators of neotropical members of this family is limited, and in the tribe Sinningieae data about bee pollination are still lacking. Here, we report on floral biology and bee pollination of four Sinningia species: S. schiffneri, S. eumorpha, S. villosa, and Sinningia "canastrensis". The flower features, such as corolla size, shape, and colour, are very different among the four species, but all conform to the melittophilous syndrome. The average nectar volume and sugar amount is low in S. schiffneri, S. eumorpha, and Sinningia "canastrensis", when compared to that of S. villosa, but low nectar amounts is a general feature of Sinningia species. The main pollinators of the four species are: small Tapinotaspidini (Trigonopedia ferruginea) of S. schiffneri, large Bombini (Bombus morio) and large Centridini (Epicharis morio) of S. eumorpha, large Euglossini (Eulaema cingulata and Eufriesea surinamensis) of S. villosa, and large Euglossini (Eufriesea violascens) and Megachilini (Megachile sp.) of Sinningia "canastrensis". Out of the four species, only S. villosa is exclusively Euglossini-pollinated. The marked differences in flower features and nectar production of these Sinningia species may reflect their pollination by distinct groups of bees. These results strengthen the idea of multiple origins for the pollination systems involving bees within this genus, which is highly supported by molecular phylogenetic analyses.