Floral anatomy of Paepalanthoideae (Eriocaulaceae, Poales) and their Nectariferous structures

BACKGROUND AND AIMS: Eriocaulaceae (Poales) is currently divided in two subfamilies: Eriocauloideae, which comprises two genera and Paepalanthoideae, with nine genera. The floral anatomy of Actinocephalus polyanthus, Leiothrix fluitans, Paepalanthus chlorocephalus, P. flaccidus and Rondonanthus roraimae was studied here. The flowers of these species of Paepalanthoideae are unisexual, and form capitulum-type inflorescences. Staminate and pistillate flowers are randomly distributed in the capitulum and develop centripetally. This work aims to establish a floral nomenclature for the Eriocaulaceae to provide more information about the taxonomy and phylogeny of the family. METHODS: Light microscopy, scanning electron microscopy and chemical tests were used to investigate the floral structures. KEY RESULTS: Staminate and pistillate flowers are trimerous (except in P. flaccidus, which presents dimerous flowers), and the perianth of all species is differentiated into sepals and petals. Staminate flowers present an androecium with scale-like staminodes (not in R. roraimae) and fertile stamens, and nectariferous pistillodes. Pistillate flowers present scale-like staminodes (except for R. roraimae, which presents elongated and vascularized staminodes), and a gynoecium with a hollow style, ramified in stigmatic and nectariferous portions. CONCLUSIONS: The scale-like staminodes present in the species of Paepalanthoideae indicate a probable reduction of the outer whorl of stamens present in species of Eriocauloideae. Among the Paepalanthoideae genera, Rondonanthus, which is probably basal, shows vascularized staminodes in their pistillate flowers. The occurrence of nectariferous pistillodes in staminate flowers and that of nectariferous portions of the style in pistillate flowers of Paepalanthoideae are emphasized as nectariferous structures in Eriocaulaceae.     

First molecular phylogenetic insights into the evolution of Eriocaulon (Eriocaulaceae,Poales)

Eriocaulon is a genus of c. 470 aquatic and wetland species of the monocot plant family Eriocaulaceae. It is widely distributed in Africa, Asia and America, with centres of species richness in the tropics. Most species of Eriocaulon grow in wetlands although some inhabit shallow rivers and streams with an apparent adaptive morphology of elongated submerged stems. In a previous molecular phylogenetic hypothesis, Eriocaulon was recovered as sister of the African endemic genus Mesanthemum. Several regional infrageneric classifications have been proposed for Eriocaulon. This study aims to critically assess the existing infrageneric classifications through phylogenetic reconstruction of infrageneric relationships, based on DNA sequence data of four chloroplast markers and one nuclear marker. There is little congruence between our molecular results and previous morphology-based infrageneric classifications. However, some similarities can be found, including Fyson’s sect. Leucantherae and Zhang’s sect. Apoda. Further phylogenetic studies, particularly focusing on less well sampled regions such as the Neotropics, will help provide a more global overview of the relationships in Eriocaulon and may enable suggesting the first global infrageneric classification.

     

Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.     

荔枝花蜜腺发育解剖学研究

荔枝花蜜腺呈盘状,位于子房和花萼之间的花托上。花盘蜜腺由表皮、产蜜组织、维管束组成。蜜腺的原始细胞由花托表面的2~3层细胞脱分化产生。成熟蜜腺产蜜组织细胞含有淀粉粒,为淀粉型蜜腺,表皮细胞内无淀粉粒。产蜜组织出现分化:PAS反应颜色深的细胞成网状分布,与表皮下方的1~2层细胞相连,构成蜜汁的运输通道;颜色浅的细胞分布于网眼处。蜜腺表皮上的角质层波状皱折,有泌蜜孔。表皮毛主要起保护作用,大部分蜜汁通过泌蜜孔排出。     

小花糖芥花蜜腺的解剖学研究

小花糖芥（Erysimum cheiranthoides L.)花蜜腺位于雄蕊部花托上,仅有2枚侧蜜腺,属十字花科侧蜜腺类型中的侧半环亚型,其2枚密腺均由分泌表皮,产蜜组织和维管束组成。分泌表皮上有变态气孔器,产蜜组织中有维管束分布,属较进化的十字花科花蜜腺亚型类型。蜜腺原基是在花的各部分原基基本分化完成后,由花托表面及雄蕊基部区域的1－2层细胞,经反分化形成,在蜜腺发育过程中,蜜腺组织中的液泡和多糖物质都发生了有规律的变化,其原蜜汁在产蜜组织中加工合成,最后由表皮细胞和变态气孔泌出。     

垂花青兰花蜜腺的发育解剖学研究

垂花青兰（Dracocephalum nutans Linn.）花蜜腺分布于子房基部的花托上,盘状蜜腺的上部裂成三小一大的四枚裂片,基部在膨大的花托外环绕一圈。蜜腺组织由分泌表皮、产蜜组织和维管束三部分组成,是典型的结构蜜腺;组织化学染色显示淀粉粒动态明显,因此又属淀粉蜜腺。在发育的过程中细胞液泡化动态明显,且淀粉粒和蛋白质具有明显的消长变化,蜜汁通过气孔器和表皮细胞的角质层泌出。     

Floral nectary morphology and evolution in Pedicularis (Orobanchaceae)

Intricate associations between floral morphology and pollinator foraging behaviour are common. In this context, the presence and form of floral nectaries can play a crucial role in driving floral evolution and diversity in flowering plants. However, the reconstruction of the ancestral state of nectary form is often hampered by a lack of anatomical studies and well‐resolved phylogenetic trees. Here, we studied 39 differentially pollinated Pedicularis spp., a genus with pronounced interspecific variation in colour, shape and size of the corolla. Anatomical and scanning electron microscopy observations revealed two nectary forms [bulged (N = 27) or elongated (N = 5)] or the absence of nectaries (N = 7). In a phylogenetic context, our data suggest that: (1) the bulged nectary should be the ancestral state; (2) nectaries were independently lost in some beaked species; and (3) elongated nectaries evolved independently in some clades of beakless species. Phylogenetic path analysis showed that nectary presence is indirectly correlated with beak length/pollinator behaviour through an intermediate factor, nectar production. No significant correlation was found between nectary type and nectar production, beak length or pollinator behaviour. Some beaked species had nectary structures, although they did not produce nectar. The nectary in beaked species may be a vestigial structure retained during a recent rapid radiation of Pedicularis, especially in the Himalaya–Hengduan Mountains of south‐western China. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 592–607.     

Gondwanan evolution of the grass alliance of families (Poales)

Phylogenetic interrelationships among all 18 families of Poales were assessed by cladistic analysis of chloroplast DNA rbcL and atpB sequences from 65 species. There are two well-supported main clades; the graminoid clade with Poaceae (grasses), Anarthriaceae, Centrolepidaceae, Ecdeiocoleaceae, Flagellariaceae, Joinvilleaceae, and Restionaceae; and the cyperoid clade with Cyperaceae, Juncaceae, and Thurniaceae. A sister group relationship between Poaceae and Ecdeiocoleaceae is identified with strong support. The sister group of this pair is Joinvilleaceae. These relationships help in elucidating the evolution of grasses and the grass spikelet. Dating of the tree was done by nonparametric rate smoothing of rbcL molecular evolution. Most Poales families date back to the Cretaceous >65 million years ago (mya). Dispersal-vicariance analysis indicates that the Poales originated in South America, the cyperoid clade in West Gondwana (South America or Africa), and the graminoid clade in East Gondwana (Australia). The Trans-Antarctic connection between South America and Australia, and its breakup about 35 mya, probably influenced the evolution of the Poales and the graminoid clade in particular, leading to vicariance between the continents, but the separation of Africa from the other Gondwanan areas, completed about 105 mya, is too old for such a relation.     

数珠珊瑚(商陆科)的花器官发生

对数珠珊瑚的花器官发生和子房的发育过程进行了观察。结果表明：(1)数珠珊瑚花被呈2／5螺旋状发生,远轴侧的1枚先发生,其次为近轴侧的1枚发生,最后侧方的2枚花被几乎同时发生,第3枚花被在靠近第1枚的位置发生,第2枚和第3枚之间有1个空隙;(2)4枚雄蕊是同时发生的;(3)心皮发生于分生组织的远轴侧,心皮原基形成后,向上向轴生长,在子房成熟前在近轴侧非正中位形成1个孔,该孔为心皮最终愈合前的残迹,到子房成熟时．因子房的生长孔被挤压缩小,在进一步的生长过程中愈合。子房由1枚心皮构成;(4)从子房发育过程的切片看,该植物的胚珠是在子房发生后不久发生的,子房上的圆孔形成时,从近轴侧的分生组织发生胚珠原基,由胚珠原基分化出珠被与珠心。     

Floral development in Scutellaria pinnatifida (Lamiaceae): the ontogenetic basis for sepal reduction

The Scutellaria is a Labiatae genus (subfamily Scutellarioideae) with a highly specialised floral structure. The genus is characterised by a peculiar two‐lobed calyx with a projecting appendage, named the scutellum. Here, we present a detailed analysis of floral development, using epi‐illumination light microscopy, to clarify open questions about its floral organisation. Floral whorls appeared in an acropetal sequence, with a marked temporal overlap of petal and stamen appearance. Organ appearance in each whorl proceeded unidirectionally from the abaxial to the adaxial side. Significant developmental features included the formation of the scutellum, reduction of sepal lobes and formation of a three‐lobed nectary disc. Our study revealed that both loss of organ initiation and fusion of primordia are responsible for the reduction in sepal members in Scutellaria. The nectary structure was markedly different from most other studied Lamiaceae.     

唐古特白刺花蜜腺的发育解剖学研究

唐古特白刺（Nitraria tangutorum Bobr.）蜜腺位于花瓣内侧,按Fahn蜜腺分类法,属花被蜜腺;其由分泌表皮细胞构成,从植物解剖学的角度来看,又属典型的非结构蜜腺;经组织化学染色显示,淀粉粒的动态不明显,因此又属非淀粉蜜腺。唐古特白刺的分泌表皮细胞,在蜜腺发育过程中特化为分泌表皮毛,分泌腔原始细胞在蜜腺发育过程中裂解成分泌腔,分泌表皮上具有特殊的角质层纹理,在分泌表皮细胞发育过程中,液泡呈现一定的变化规律,其变化与蜜汁的合成和分泌规律相关,液泡是参与了多糖物质的降解、蜜汁的转运等物质的循环而发生着有规律的变化,淀粉和糖原的动态不明显。最后形成的蜜汁经分泌腔,由分泌表皮细胞特化为单细胞的表皮毛中泌出,在败花期时,分泌表皮毛萎缩并随花瓣一起脱落,泌蜜由此停止。     

马先蒿属植物花冠分化与繁殖适应的研究进展

结合已有的研究报道和作者近年来的工作,对马先蒿属（Pedicularis）植物的花冠多样化成因与繁殖适应特性进行了总结和探讨。通过对该属4种进化花冠型的花器官发生和分化的研究发现,花部各器官在发生和发育初期基本一致,后期上唇形态的分化是导致成熟花形态结构产生较大差异的重要阶段。孢粉学研究认为,花冠类型与花粉萌发孔类型之间具有显著相关性;萌发沟的演化可能与繁殖适应有一定的关系。分子系统学研究表明,多样化的花冠类型在不同的谱系内经过若干次的独立进化而表现出了高度的平行演化（parallelism）。传粉生物学研究证实,该属植物花冠多样化与其主要传粉者熊蜂属（Bombus）昆虫的传粉行为存在较为密切的关系。具有相同（似）花冠类型的马先蒿可能被同种或不同种的熊蜂以相同的方式访问,但在花粉落置位置上存在显著差异,这可能有助于同域分布重叠的物种间在生殖上的机械隔离,而花冠的分化在一定程度上促进了新的物种形成。     

地椒花蜜腺发育的解剖学观察

通过显微和亚显微观察对地椒花蜜腺的发育进行了研究。地椒花蜜腺位于子房基部的花盘上,属于盘状蜜腺,新鲜时呈绿色。蜜腺由分泌表皮和泌蜜组织组成,分泌表皮为一层细胞,表皮细胞角质膜较厚,表皮上分布着大量的气孔器,气孔器突出于表面;泌蜜组织细胞多层。花盘中央有维管束通向子房,在维管束和泌蜜组织之间有４￣５层大型的薄壁细胞。蜜腺由花盘的表皮及其内侧相邻的细胞发育而来,在发育过程中,其细胞中的液泡和贮藏的淀粉     

Development of the gametophytes, flower, and floral vasculature in Dichorisandra thyrsiflora (Commelinaceae)

The flowers of Dichorisandra thyrsiflora (Commelinaceae) are monosymmetric and composed of three sepals, three petals, six stamens, and three connate carpels. The anthers are poricidal and possess a wall of five cell layers (tapetum included). This type of anther wall, not previously observed in the Commelinaceae, is developmentally derived from the monocotyledonous type via an additional periclinal division and the persistence of the middle layers through anther dehiscence. Secondary endothecial thickenings develop in the cells of the two middle layers only. The tapetum is periplasmodial and contains raphides. Microsporogenesis is successive and yields both decussate and isobilateral tetrads. Pollen is shed as single binucleate grains. The gynoecium is differentiated into a globose ovary, hollow elongate style, and trilobed papillate stigma. Each locule contains six to eight hemianatropous to slightly campylotropous crassinucellar ovules with axile (submarginal) placentation. The ovules are bitegmic with a slightly zig-zag micropyle. Megagametophyte development is of the Polygonum type. The mature megagametophyte consists of an egg apparatus and fusion nucleus; the antipodals having degenerated. The floral vasculature is organized into an outer and inner system of bundles in the pedicel. The outer system becomes ventral carpellary bundles. All other floral vascular traces originate from the inner system.     

鹅掌柴花蜜腺的发育解剖学研究

对鹅掌柴(Scheffler octophylla Harms．)花蜜腺的发育进行解剖结构观察。鹅掌柴花盘蜜腺位于下位子房上方环绕花柱基部。蜜腺由分泌表皮、产蜜组织组成,心皮维管束与其相邻并发出一些伸入蜜腺基部的短分枝。蜜腺起源于心皮原基基部外侧的几层细胞。鹅掌柴花蜜腺为淀粉型蜜腺,淀粉粒为许多微小颗粒聚集成的复粒。原蜜汁由蜜腺基部维管束的筛管提供,达产蜜组织细胞和表皮细胞后以淀粉粒的形式贮藏。泌出的蜜汁一部分来自淀粉粒的降解,一部分来自泌蜜期输入的原蜜汁。表皮和产蜜组织细胞均具泌蜜功能。泌出的蜜汁大部分通过气孔排出,还有部分由角质层渗出。     

Changes in the range of the medicinal herb Eriocaulon buergerianum Körnicke. (Eriocaulaceae) under climate change

• Eriocaulon buergerianum Körnicke. (Eriocaulaceae) is one of the most common and least expensive herbal medicines for eye disease. This species is facing potential threats from climate change. Insufficient biogeographic knowledge of this plant species can hinder its effective management for long‐term population survival.
• We integrated ecological niche modelling (Biomod2) with 70 records of E. buergerianum and eight environmental variables to estimate changes in distribution over time. A core area Zonation algorithm was introduced to identify conservation priority areas.
• Our results indicate that the range of E. buergerianum will likely decrease in the future: the overall range change on average is ?44.36 ± 21.56% (?3.70% to ?77.73%); values of range loss and range gain are 45.79 ± 20.30% (9.29–78.19%) and 1.43 ± 1.53% (0.18–5.59%), respectively. According to conservation priority analysis, the mandatory reserve (top 5%), negotiable reserve (0.95–0.9) and partial reserve (0.9–0.8) areas are 19,799, 19,799 and 39,597 km², respectively. The areas identified as conservation priority are located in the southeast, especially in northern Taiwan and the Wuyi Mountains.
• Based on these results, we suggest a re‐evaluation of the threatened status of this species, with a potential upgrade to the vulnerable (VU) category. To overcome the adverse conditions faced by populations of E. buergerianum in China, we propose a multi‐faceted conservation strategy involving more complete resource assessment, a monitoring system, medical research focused on revealing medicinal components or substitutes, and a regional development plan that considers both wildlife and socio‐economic issues.

     

Morphology and anatomy of inflorescence and inflorescence axis in Paepalanthus sect. Diphyomene Ruhland (Eriocaulaceae,Poales) and its taxonomic implications

Paepalanthus sect. Diphyomene has inflorescences arranged in umbels. The underlying bauplan seems however to be more complex and composed of several distinct subunits. Despite appearing superficially very similar, the morphology and anatomy of the inflorescences can supply useful information for the understanding of the phylogeny and taxonomy of the group. Inflorescences of Paepalanthus erectifolius, Paepalanthus flaccidus, Paepalanthus giganteus, and Paepalanthus polycladus were analyzed in regard to branching pattern and anatomy. In P. erectifolius, P. giganteus and P. polycladus the structure is a tribotryum, with terminal dibotryum, and with pherophylls bearing lateral dibotrya. In P. flaccidus, the inflorescence is a pleiobotryum, with terminal subunit, and without pherophylls. Secondary inflorescences may occur in all species without regular pattern. Especially when grown in sites without a pronounced seasonality, the distinction between enrichment zone (part of the same inflorescence) and new inflorescences may be obscured. The main anatomical features supplying diagnostic and phylogenetic information are as follows: (a) in the elongated axis, the thickness of the epidermal cell walls and the cortex size; (b) in the bracts, the quantity of parenchyma cells (c) in the scapes, the shape and the presence of a pith tissue. Therefore, P. sect. Diphyomene can be divided in two groups; group A is represented by P. erectifolius, P. giganteus and P. polycladus, and group B is represented by P. flaccidus. The differentiation is based in both, inflorescence structure and anatomy. Group A presents a life cycle and anatomical features similar to species of Actinocephalus. Molecular trees also point that these two groups are closely related. However, inflorescence morphology and blooming sequence are different. Species of group B present an inflorescence structure and anatomical features shared with many genera and species in Eriocaulaceae. The available molecular and morphology based phylogenies still do not allow a precise allocation of the group in the bulk of basal species of Paepalanthus collocated in P. sect. Variabiles. The characters described and used here supply however important information towards this goal.     

拟南芥花蜜腺筛分子及蜜腺组织发育过程中的细胞学研究

应用高压冷冻和低温替代技术,对拟南芥(Arabidopsis thaliana L.)花蜜腺发育过程中细胞的超微结构变化进行了研究.蜜腺组织中深色细胞的超微结构与筛分子早期分化的超微结构十分相似:细胞核中染色质逐渐出现凝集并且边缘化;细胞器分布异常;细胞质浓稠.这些超微结构特征与近年来报道的动植物细胞程序性死亡的超微结构相似.在筛分子和深色细胞分化中,细胞核及一些细胞器的逐渐解体与原蜜汁的运输、加工和蜜汁的分泌有直接联系.这反映了蜜腺发育过程中筛分子和蜜腺组织的细胞学变化是与蜜腺的生长、发育和生理功能的完善联系在一起的.     

Floral ontogeny in Dipterygeae (Fabaceae) reveals new insights into one of the earliest branching tribes in papilionoid legumes

Flowers of Dipterygeae (Fabaceae, Papilionoideae) exhibit an unusual petaloid calyx. The two adaxial sepals are large and petaloid, and the three abaxial sepals form a three‐toothed lobe. The goal of this study was to elucidate the ontogenetic pathways of this peculiar calyx in light of the floral development of the three genera that comprise the tribe. Floral buds of Dipteryx alata, Pterodon pubescens and Taralea oppositifolia were analysed using scanning electron microscopy and light microscopy. The order of bracteole and sepal initiation varies among the species. The androecium is asymmetric. The carpel cleft is positioned to the right or to the left, and is opposite the adaxial antepetalous stamen. The peculiarity of the calyx becomes noticeable in the intermediate stages of floral development. It results from the differential growth of the sepal primordia, in which the abaxial and lateral primordia remain diminutive during floral development, compared with the adaxial ones that enlarge and elongate. Bracteoles, abaxial sepals, petals and anthers are appendiculate, except in T. oppositifolia, in which the appendices were not found in bracteoles or anthers. These appendices comprise secretory canals or cavities. Considering that the ontogenetic pathway for the formation of the petaloid calyx is similar and exclusive for Dipterygeae, it might be a potential synapomorphy for the group, with the presence of secretory canals in the appendices of abaxial and lateral sepals and petals. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 529–550.