COMPARATIVE ONTOGENY OF THE PERIANTH IN MIMOSOID LEGUMES

Comparative ontogeny of the perianth is reported for representative genera and species among mimosoid legumes in order to elucidate intertribal relationships and also relationships to the other two subfamilies of legumes. Initiation of the perianth is acropetal in two whorls. The calyx arises first followed by the corolla. Order of initiation of both calyx and corolla is determined during early ontogeny. Four different types of order of initiation have been found in the calyx: helical, simultaneous within one whorl, bidirectional, and ring meristem. Helical initiation is considered primitive; simultaneous within one whorl, bidirectional, and ring meristem are considered derived. Differences during early organogeny in the calyx among mimosoids result in similar morphologies of the mature calyx which indicates that parallel evolution may have played a major role in evolution of radial symmetry within the group. Order of initiation of the corolla is uniformly simultaneous whorled with one exception. Position of organs is a significant feature which separates mimosoids from caesalpinioids and papilionoids. In mimosoids the median sepal is located abaxially and the median petal adaxially in relation to the subtending bract. In both caesalpinioids and papilionoids the median sepal is located abaxially and the median petal adaxially in relation to the subtending bract. Fusion of the calyx in some taxa can be interpreted as an example of acceleration.     

Evolution of pollen,stigmas and ovule numbers at the caesalpinioid–mimosoid interface (Fabaceae)

In this study we examine the pollen, stigmas and ovaries from 62 collections of herbarium material representing 16 genera, using light and scanning electron microscopy. The caesalpinioid Dimorphandra group (Burkea, Dimorphandra, Erythrophleum, Mora, Pachyelasma, Stachyothyrsus and Sympetalandra) pollen grains are small, tricolporate monads, with perforate or psilate ornamentation. Dinizia, Pentaclethra and Aubrevillea have morphological characters that have suggested either a mimosoid or caesalpinioid placement. Dinizia pollen is in permanent tetrads with clavate ornamentation. Pentaclethra pollen grains are monads, two species have tricolporate pollen and the third is porate. Aubrevillea has tricolporate, finely reticulate monads. All ten genera have variable, non‐predictable stigma type and ovule number. The mimosoid Adenanthera group (Adenanthera, Tetrapleura, Amblygonocarpus, Pseudoprosopis, Calpocalyx and Xylia) pollen grains are in 8‐ to 16‐grain polyads. In all Adenanthera group species, the stigmatic cavity is only large enough to accommodate one polyad. In addition, the number of ovules present matches the number of pollen units in one polyad. Polyads have porate, operculate apertures that differ in layout, aperture morphology and development when compared with caesalpinioid and other eudicot pollen. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 594–615.     

Is nectar reabsorption restricted by the stalk cells of floral and extrafloral nectary trichomes?

Reabsorption is a phase of nectar dynamics that occurs concurrently with secretion; it has been described in floral nectaries that exude nectar through stomata or unicellular trichomes, but has not yet been recorded in extrafloral glands. Apparently, nectar reabsorption does not occur in multicellular secretory trichomes (MST) due to the presence of lipophilic impregnations – which resemble Casparian strips – in the anticlinal walls of the stalk cells. It has been assumed that these impregnations restrict solute movement within MST to occur unidirectionally and exclusively by the symplast, thereby preventing nectar reflux toward the underlying nectary tissues. We hypothesised that reabsorption is absent in nectaries possessing MST. The fluorochrome lucifer yellow (LYCH) was applied to standing nectar of two floral and extrafloral glands of distantly related species, and then emission spectra from nectary sections were systematically analysed using confocal microscopy. Passive uptake of LYCH via the stalk cells to the nectary tissues occurred in all MST examined. Moreover, we present evidence of nectar reabsorption in extrafloral nectaries, demonstrating that LYCH passed the stalk cells of MST, although it did not reach the deepest nectary tissues. Identical (control) experiments performed with neutral red (NR) demonstrated no uptake of this stain by actively secreting MST, whereas diffusion of NR did occur in plasmolysed MST of floral nectaries at the post‐secretory phase, indicating that nectar reabsorption by MST is governed by stalk cell physiology. Interestingly, non‐secretory trichomes failed to reabsorb nectar. The role of various nectary components is discussed in relation to the control of nectar reabsorption by secretory trichomes.     

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。     

Structure and development of bracteal nectary glands in Aphelandra (Acanthaceae)

A survey of bracteal (extrafloral) nectaries in species of Aphelandra (Acanthaceae) reveals substantial diversity. Each bracteal nectary is an aggregate of individual glands that vary in number, size, and structure among species. Glands contain three cell layers: a palisade-like secretory cell layer, a one-to-many-celled intermediate layer with thickened cell walls, and a foot layer. Members of the A. pulcherrima complex have one of two distinct gland types: relatively small glands with a single-celled intermediate layer or larger glands that have a multicellular intermediate layer. Nectaries composed of small glands are patches of many (>50) glands, whereas those composed of large glands are patches of < 10 glands. Four outgroup species have bracteal nectaries of numerous small glands with pluricellular intermediate layers. Glands of all three types are initiated as single enlarged protodermal cells, and all undergo similar early periclinal divisions; the large-gland type shows greater subsequent enlargement with many more anticlinal divisions. The bracteal nectar glands are interpreted to be homologous with simpler glandular trichomes, and mark a monophyletic lineage within Aphelandra. Comparisons with outgroup species show that both nectary types in the A. pulcherrima complex have diverged from an ancestral condition of numerous small glands with pluricellular intermediate layers. Use of the ontogenetic criterion to polarize gland type within the A. pulcherrima complex would yield erroneous results because evolution has apparently involved a developmental truncation with loss of cell divisions in the intermediate layer of small glands. Comparable nectar glands in more distant taxa are interpreted as remarkable cases of convergent evolution, perhaps from similar trichome precursors.     

PRIMITIVE MIMOSOID FLOWERS FROM THE PALEOCENE-EOCENE AND THEIR SYSTEMATIC AND EVOLUTIONARY IMPLICATIONS

Compressed mimosoid inflorescences from a Paleocene-Eocene boundary locality in western Tennessee are the earliest fossil evidence of the subfamily. The discovery confirms the antiquity of a suite of characters that has been considered primitive based on the comparative morphology of modern mimosoids. The fossil characters are also consistent with the suggested close relationship (ancestral or sister group) between the subfamily Mimosoideae and the Dimorphandra group of the tribe Caesalpinieae (subfamily Caesalpinioideae). These flowers show little change in morphology or size in the basal to Upper Eocene interval.     

Chelonian mental glands

A survey of 69 of the 74 currently recognized chelonian genera revealed that 21 genera in three families (Emydidae, Platysternidae and Testudinidae) possess paired integumentary glands or gland vestiges in the anterior throat skin. These glands are here termed mental glands; they are holocrine and may be classified morphologically as follows: Class I mental glands are large, complex, multilobed, have specialized ducts, and are found only in the genus Gopherus: Class II mental glands are small, simple sacklike invaginations containing secretory cells or keratinizing cells. The structure of Class II glands varies from distinctive and saccular to shallow keratinized invaginations having no glandular tissue; they are found only in the families Platysternidae and Emydidae. Mental glands occur in 17 of the 22 genera in the subfamily Batagurinae (sensu McDowell, 64); only 2 of 9 genera in the subfamily Emydinae have these glands. The taxonomic occurrence of mental glands suggests that they are primitive structures. The loss of mental glands in most emydines is interpreted as a subfamilial trend toward integumentary simplification.     

Two structures and functions in the nectary of Frankincense tree (Boswellia sacra Flueck.)

The flowers of Boswellia sacra Flueck. (Burseraceae) present a showy nectariferous ring which changes color from yellow to brilliant red in a few days. In this paper, the structure and development of this peculiar nectary were studied using light microscopy as well as scanning and transmission electron microscopy. The nectary presents a double way of secretion, since it releases nectar through both glandular trichomes and nectarostomata. A direct vascular supply is lacking; however, a large quantity of starch was stored in glandular parenchyma cells at the early secretory stage, while it disappeared at the senescent stage. The nectary, besides showing the typical secretory parenchyma cells, is characterized by the occurrence of highly osmiophilic cell rows. Experimental evidence shows that these cells are involved in nectarostoma secretion. The different secreting structures are described and their role is discussed.     

Novel reports of glands in Neotropical species of Indigofera L. (Leguminosae,Papilionoideae)

By considering controversial discussions in the literature with regard to gland denomination in Indigofera species, as well as the taxonomic value of secretory structures in Leguminosae, we aim to morphologically detail glands that had been previously observed in I. microcarpa and I. sabulicola, and to investigate the occurrence of glands in vegetative and reproductive organs of other six Neotropical species that belong to the genus. Glands analyzed through scanning electronic microscopy (SEM) in combination with anatomic analyses correspond to secretory trichomes that are classified into seven types. Main variations in relation to types occurred with regard to head shape and peduncle size. Trichome heads were multicellular, with a thin cuticle. Hollow heads with conspicuous inner space characterized only one type (type 1); the other trichome types had massive heads. Peduncles, which varied from biseriate to multiseriate, had thick, pecto-cellulosic cell walls. Trichomes were found on stems, stipules, petioles, rachis, petiolules, leaflets, bracteoles, sepals, standards and fruits, more commonly along the margins. Each of the eight Indigofera species analyzed had at least two different trichome types out of the seven types that occurred in reproductive and vegetative organs of these taxa. Various types of secretory trichomes were found in I. campestris, I. lespedezioides, I. microcarpa, I. spicata, I. suffruticosa and I. truxillensis. Stems and rachis were the vegetative organs in which a greater variety of trichomes occurred, and sepals were parts of reproductive organs with the same status. Five out of the seven secretory trichome types occurred on both vegetative and reproductive organs. Distribution and gland types differed between species and these gland distribution patterns can be used as diagnostic characters. Reports of glands in Indigofera campestris, I. hirsuta, I. lepedezioides, I. suffruticosa, I. spicata and I. truxillensis, their recognition as secretory trichomes, and the morphological variety of types found for such trichomes are novel data for Indigofera.     

Distribution and structural aspects of extrafloral nectaries in Cedrela fissilis (Meliaceae)

The occurrence of extrafloral nectaries (EFNs) in Meliaceae has been reported for some genera, but little anatomical data are available. Therefore, to determine the distribution and structural aspects of EFNs, Cedrela fissilis Vell. leaves in different stages of development were collected, fixed, and processed for light and scanning electron microscopy. On the petiole, rachis and petiolule, EFNs were found to be arranged predominantly towards the abaxial surface, while their occurrence in leaflet blades was restricted to the abaxial surface of the major veins, noticeably on the midrib. Basal leaflets displayed few EFNs; however, we observed an increase towards the leaf's apex. The leaf can contain more than 300 inconspicuous EFNs, which show secretory activity throughout the leaf's life. Two EFN morphotypes were visible: flattened or elevated, both circular or slightly elliptical and similar in origin and tissue composition. The secretory tissue is embedded in the rachis cortex or in the major veins of the leaf blade and EFNs are not vascularized. The EFN secretory pole shows a uniseriate epidermis with compactly arranged cells and a thin cuticle; stomata and trichomes are absent. The observation of ant visits at these structures reinforces the assumption that EFNs mediate ant–plant interactions and play a protective role against herbivores throughout the life of a leaf.     

Stipular extranuptial nectaries new to Polygala: morphology and ontogeny

Although taxonomic studies indicate that approximately one‐third of the genera of Polygalaceae have nodal glands, few anatomical data are available on the structure and ontogeny of these secretory organs. We studied the as yet unknown origin, structure and function of such glands in Polygala laureola. During field observations, we detected glucose in the secretion using Glicofita Plus® and visitors were recorded. Vegetative shoot apices and nodal glands were examined using light microscopy and scanning electron microscopy. The presence of glucose in the secretion allowed us to identify these nodal glands as extranuptial nectaries. Secretory cells occupy a medullary position and are surrounded by phloem. Vascular bundles are concentric, and xylem is only observed at the basal region of the nectary. Nectar is released during the daytime through a pore at the top of the nectary. A stipular origin was confirmed by the fact that the procambial strand is connected to the leaf trace, opposite the leaf gap. The occurrence of stipular extranuptial nectaries in a nodal position is new to Polygala. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 40–50.     

Structure of the receptacular nectary and circadian metabolism of starch in the ant‐guarded plant Ipomoea cairica (Convolvulaceae)

Nectaries occur widely in Convolvulaceae. These structures remain little studied despite their possible importance in plant–animal interactions. In this paper, we sought to describe the structure and ultrastructure of the receptacular nectaries (RNs) of Ipomoea cairica, together with the dynamics of nectar secretion. Samples of floral buds, flowers at anthesis and immature fruits were collected, fixed and processed using routine methods for light, scanning and transmission electron microscopy. Circadian starch dynamics were determined through starch measurements on nectary sections. The secretion samples were subjected to thin layer chromatography. RNs of I. cairica were cryptic, having patches of nectar‐secreting trichomes, subglandular parenchyma cells and thick‐walled cells delimiting the nectary aperture. The glandular trichomes were peltate type and had typical ultrastructural features related to nectar secretion. The nectar is composed of sucrose, fructose and glucose. Nectar secretion was observed in young floral buds and continued as the flower developed, lasting until the fruit matured. The starch content of the subglandular tissue showed circadian variation, increasing during the day and decreasing at night. The plastids were distinct in different portions of the nectary. The continuous day–night secretory pattern of the RNs of I. cairica is associated with pre‐nectar source circadian changes in which the starch acts as a buffer, ensuring uninterrupted nectar secretion. This circadian variation may be present in other extrafloral nectaries and be responsible for full daytime secretion. We conclude that sampling time is relevant in ultrastructural studies of dynamic extranuptial nectaries that undergo various changes throughout the day.     

The uncommon cavitated secretory trichomes in Bauhinia s.s. (Fabaceae): the same roles in different organs

Cavitated secretory trichomes are characterized by a short or absent stalk that is connected to a secretory hollow head. They are rare structures in angiosperms; in Fabaceae, they have been recorded in only seven genera, including Bauhinia s.s. Because B. curvula and B. rufa exhibit glands that are responsible for attracting pollinators to flowers, this study aimed to test whether the cavitated secretory trichomes present in the flowers of these species have an attraction function. As leaf trichomes are commonly related to plant defence, comparative analyses of the morphology, ontogeny, ultrastructure and chemical profile of the secretory trichomes present in flowers and leaves were conducted. It was found that cavitated secretory trichomes are similar in their external morphology and development, regardless of the organ or species analysed. However, interspecific differences were found in the secretion process and chemical profile of the exudate. The differences found in the cavitated secretory trichomes between species indicate that they secrete distinct compounds, whereas the similarities found in these structures between vegetative and reproductive organs indicate that the cavitated trichomes have equivalent ecological functions within a species, probably in plant defence during organ development. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 104–122.     

The extrafloral nectaries of cowpea (Vigna unguiculata (L.) Walp): I. Morphology,anatomy and fine structure

The cowpea bears two distinctive types of extrafloral nectaries. One, on the stipels of trifoliolate leaves, consists of a loosely demarcated abaxial area (1–2 mm diameter) of widely-spaced trichomes (papillae) borne on a stomata-free epidermis, and lacking a specific vascular supply. Each trichome has up to eight apical (head) cells, two to four intermediate cells, and a single large stalk cell. The secretory faces of the apical cells bear wall ingrowths and an easily detached cuticle. The wall separating the stalk cell and the underlying epidermal cell(s) has a mean plamodesmatal frequency of 25/m². The second type of nectary consists of a large elliptical mound of tissue (short and long axes about 2 mm and 4 mm) formed between a pair of flowers on an inflorescence stalk. It comprises four to eight cone-shaped subnits of secretory tissue, each with a circular secretory orifice and an individual supply of phloem, but not of xylem. Cells of the secretory tissue of the nectary subunits separate as they mature, and nectar flows to the orifice through the resulting intercellular spaces. Intact secretory cells and cellular debris are extruded into the nectar. Some of the sieve elements terminating in the inner secretory tissue exhibit open sieve pores. Each mature secretory cell contains many small (2 m diameter) spherical protein bodies and one to three large (up to 2–3 m diameter 15 m long), paracrystalline bodies. These inclusions are absent or not fully developed in inner, less mature regions of the secretory tissue. Mechanisms of secretion are proposed for the two classes of nectary, including estimates of flux of sugar into the trichomes of the stipel nectary.     

Anatomy and histochemistry of the nectaries of Rodriguezia venusta (Lindl.) Rchb. f. (Orchidaceae)

Orchidaceae is one of the largest angiosperm families. Although extensively studied, reports of anatomy of secretory structures of orchids are relatively scarce. Rodriguezia venusta is an epiphytic orchid occurring in Brazil and Peru that has floral and extrafloral nectaries. This study describes the structure and the histochemistry of these secretory structures. Floral and extrafloral nectary samples were obtained from R. venusta plants that were collected in a gallery forest in the State of Bahia, Brazil, and grown in a greenhouse. Theses samples were fixed and processed according to routine procedures in plant anatomy and histochemistry or for scanning electron microscopy. The extrafloral nectaries occur on the edge and sub-edge of young leaves and at the basal portion of bracts that subtend the floral buds. They are structurally very similar, being formed by a nectary parenchyma and a simple epidermis with stomata (“non-structured nectaries”). The floral nectary is inserted at the floral receptacle fused with the labellum base, between this structure and the two inferior connate sepals. This nectary consists of an epidermis with numerous specific nectar secreting trichomes, a subnectary and a nectary parenchyma abundantly supplied by vascular terminations. Its structure is complex and distinct from other floral nectaries described for Orchidaceae.     

Anatomical and ultrastructural changes of the floral nectary ofPisum sativum L. during flower development

Summary The floral nectary ofPisum sativum L. is situated on the receptacle at the base of the gynoecium. The gland receives phloem alone which departed the vascular bundles supplying the staminal column. Throughout the nectary, only the companion cells of the phloem exhibited wall ingrowths typical of transfer cells. Modified stomata on the nectary surface served as exits for nectar, but stomatal pores developed well before the commencement of secretion. Furthermore, stomatal pores on the nectary usually closed by occlusion, not by guard-cell movements. Pore occlusion was detected most frequently in post-secretory and secretory glands, and less commonly in pre-secretory nectaries. A quantitative stereological study revealed few changes in nectary fine structure between buds, flowers secreting nectar, and post-secretory flowers. Dissolution of abundant starch grains in plastids of subepidermal secretory cells when secretion commenced suggests that starch is a precursor of nectar carbohydrate production. Throughout nectary development, mitochondria were consistently the most plentiful organelle in both epidermal and subepidermal cells, and in addition to the relative paucity of dictyosomes, endoplasmic reticulum, and their associated vesicles, the evidence suggests that floral nectar secretion inP. sativum is an energy-requiring (eccrine) process, rather that granulocrine.Abbreviations ER endoplasmic reticulum - GA glutaraldehyde - SEM scanning electron microscopy     

荇菜花蜜腺的发育研究

荇菜花蜜腺的发育过程可分为：起源期、生长期、分泌期以及泌蜜停止期等４个时期。荇菜的５枚花蜜腺均起源于子房基部的表皮及表皮内的２－４层细胞。这些细胞经反分化后分别成为蜜腺的原分泌表皮及原泌蜜组织,两部分细胞径不断地分裂分化,最冬成为成熟蜜腺。在蜜腺发育过程中,蜜腺的分泌表皮及蜜腺组织内的内质网、质体、线粒体、液泡等细胞器结构均发生了有规律的变化,内质网在蜜腺分泌期最为发达,且产生大量的分泌小泡。质体     

INVESTIGATIONS OF TERTIARY ANGIOSPERMS: A NEW FLORA INCLUDING EOMIMOSOIDEA PLUMOSA FROM THE OLIGOCENE OF EASTERN TEXAS

A new fossil flower and inflorescence-bearing locality has been discovered in the Oligocene of the Texas Gulf Coast. The new flora is similar to the Middle Eocene Claiborne Flora of the southeastern USA, but the quality of preservation is sometimes better in the Oligocene fossils. One component of the new flora, a mimosoid legume inflorescence, appears identical with Eomimosoidea plumosa, first reported from the Claiborne Formation of western Tennessee. Investigations of these younger specimens indicate that the taxon had changed little during the Middle Eocene-Oligocene interval, and the better quality of preservation of the Texas specimens has provided further insights into the structure of the fossils. Comparisons of the fine structural details of the pollen of Eomimosoidea with similar pollen of extant mimosoids has confirmed that the fossil genus is indeed extinct and suggests that tetrahedral tetrads of columellate, tricolporate pollen grains are ancient, possibly primitive, in the Mimosoideae.     

Floral nectar production and carbohydrate composition and the structure of receptacular nectaries in the invasive plant <Emphasis Type="Italic">Bunias orientalis</Emphasis> L. (Brassicaceae)

The data relating to the nectaries and nectar secretion in invasive Brassicacean taxa are scarce. In the present paper, the nectar production and nectar carbohydrate composition as well as the morphology, anatomy and ultrastructure of the floral nectaries in Bunias orientalis were investigated. Nectary glands were examined using light, fluorescence, scanning electron and transmission electron microscopy. The quantities of nectar produced by flowers and total sugar mass in nectar were relatively low. Total nectar carbohydrate production per 10 flowers averaged 0.3 mg. Nectar contained exclusively glucose (G) and fructose (F) with overall G/F ratio greater than 1. The flowers of B. orientalis have four nectaries placed at the base of the ovary. The nectarium is intermediate between two nectary types: the lateral and median nectary type (lateral and median glands stay separated) and the annular nectary type (both nectaries are united into one). Both pairs of glands represent photosynthetic type and consist of epidermis and glandular tissue. However, they differ in their shape, size, secretory activity, dimensions of epidermal and parenchyma cells, thickness of secretory parenchyma, phloem supply, presence of modified stomata and cuticle ornamentation. The cells of nectaries contain dense cytoplasm, plastids with starch grains and numerous mitochondria. Companion cells of phloem lack cell wall ingrowths. The ultrastructure of secretory cells indicates an eccrine mechanism of secretion. Nectar is exuded throughout modified stomata.