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1.

Background and Aims

The Borasseae form a highly supported monophyletic clade in the Arecaceae–Coryphoideae. The fruits of Coryphoideae are small, drupaceous with specialized anatomical structure of the pericarp and berries. The large fruits of borassoid palms contain massive pyrenes, which develop from the middle zone of the mesocarp. The pericarp structure and mode of its development in Borasseae are similar to those of Eugeissona and Nypa. A developmental carpological study of borassoid palms will allow us to describe the process of pericarp development and reveal the diagnostic fruit features of borassoid palms, determine the morphogenetic fruit type in Borasseae genera, and describe similarities in fruit structure and pericarp development with other groups of palms.

Methods

The pericarp anatomy was studied during development with light microscopy based on the anatomical sections of fruits of all eight Borasseae genera.

Key Results

The following general features of pericarp structure in Borasseae were revealed: (1) differentiation of the pericarp starts at early developmental stages; (2) the exocarp is represented by a specialized epidermis; (3) the mesocarp is extremely multilayered and is differentiated into several topographical zones – a peripheral parenchymatous zone(s) with scattered sclerenchymatous elements and vascular bundles, a middle zone (the stony pyrene comprising networks of elongated sclereids and vascular bundles) and an inner parenchymatous zone(s); (4) differentiation and growth of the pyrene tissue starts at early developmental stages and ends long before maturation of the seed; (5) the inner parenchymatous zone(s) of the mesocarp is dramatically compressed by the mature seed; (6) the endocarp (unspecialized epidermis) is not involved in pyrene formation; and (7) the spermoderm is multilayered in Hyphaeninae and obliterated in Lataniinae.

Conclusions

The fruits of Borasseae are pyrenaria of Latania-type. This type of pericarp differentiation is also found only in Eugeissona and Nypa. The fruits of other Coryphoideae dramatically differ from Borasseae by the pericarp anatomical structure and the mode of its development.  相似文献   

2.

Palm fruits show great structural complexity, and in-depth studies of their development are still scarce. This work aimed to define the developmental stages of the fruit of the neotropical palm Butia capitata and to characterize the ontogenesis of its pericarp. Biometric, anatomical, and histochemical evaluations were performed on pistillate flowers and developing fruits. The whole fruit develops in three phases: (I) histogenesis (up to 42 days after anthesis – DAA), when the topographic regions of the pericarp are defined; (II) pyrene maturation (42 to 70 DAA), when the sclerified zone of the pericarp is established; and (III) mesocarp maturation (70 to 84 DAA), when reserve deposition is completed. During pericarp ontogenesis (i) the outer epidermis and the outer mesophyll of the ovary give origin to the exocarp (secretory epidermis, collenchyma, parenchyma, sclerenchyma, and vascular bundles); (ii) the median ovarian mesophyll develops into the mesocarp, with two distinct topographical regions; (iii) the inner ovarian epidermis originates the endocarp; and in the micropylar region, it differentiates into the germination pore plate, a structure that protects the embryo and controls germination. (iv) Most of the inner region of the mesocarp fuses with the endocarp and, both lignified, give rise to the stony pyrene; (v) in the other regions of the mesocarp, carbohydrates and lipids are accumulated in a parenchyma permeated with fiber and vascular bundles. The development of the B. capitata pericarp presents high complexity and a pattern not yet reported for Arecaceae, which supports the adoption of the Butia-type pyrenarium fruit class.

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3.
The indehiscent fruitlets of the apparently basalmost extant angiosperm, Amborella trichopoda, have a pericarp that is differentiated into five zones, a thin one‐cell‐layered skin (exocarp), a thick fleshy zone of 25–35 cell layers (outer mesocarp), a thick, large‐celled sclerenchymatous zone (unlignified) of 6–18 cell layers (middle mesocarp), a single cell layer with thin‐walled (silicified?) cells (inner mesocarp), and a 2–4‐cell‐layered, small‐celled sclerenchymatous zone (unlignified) derived from the inner epidermis (endocarp). The border between inner and outer mesocarp is not even but the inner mesocarp forms a network of ridges and pits; the ridges support the vascular bundles, which are situated in the outer mesocarp. In accordance with previous observations by Bailey & Swamy, no ethereal oil cells were observed in the pericarp; however, lysigenous cavities as mentioned by these authors are also lacking; they seem to be an artefact caused by re‐expanding dried fruits. The seed coat is not sclerified. The fruitlets of Amborella differ from externally similar fruits or fruitlets in other basal angiosperms, such as Austrobaileyales or Laurales, in their histology. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 148 , 265–274.  相似文献   

4.
All Illicium spp. have explosive fruits, which is a unique character among the basal grade of angiosperms. Illicium fruits consist of several ventrally dehiscing follicles developing from conduplicate carpels, with a prominent, slightly postgenitally fused ventral slit. The closure of the ventral slit is also secured by two mirror‐symmetrical massive longitudinal sclerenchymatous bands in the mesocarp along the edges and by turgor pressure. The pericarp differentiates into a fleshy (or coriaceous) peripheral zone (exocarp and mesocarp) with numerous ethereal‐oil‐containing cells and a sclerenchymatous (single‐layered, palisade) inner zone (endocarp). Dehydration of the fleshy zone of the pericarp and partial compression of the epidermal sclereids with U‐shaped wall thickenings lining the ventral suture are instrumental in explosive fruitlet dehiscence. Generally, the fruit structure of Illicium differs dramatically from those in other early diverging angiosperms. Gynoecium and fruit structure (and a probable early Cretaceous divergence from the SchisandraKadsura clade) provide evidence for treatment of Illicium as separate from Schisandraceae s.s. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 640–654.  相似文献   

5.
Lepidocaryum tenue, Mauritia flexuosa and Mauritiella armata belong to the subtribe Mauritiinae, one early divergent lineage of the Arecaceae and one of the few of Calamoideae that occur in South America. These species occur in swampy environments and have fruits that are characteristically covered with scales. The objective of this study was to describe the formation of the layers of the pericarp within this subtribe and attempt to correlate fruit structure with the environment where species typically occur. Toward this goal, flowers in pre-anthesis and anthesis and fruits throughout development were analyzed using standard methods for light microscopy. The ontogeny of the layers of the pericarp of all three species was found to be similar. The scales were formed from non-vascularized emergences composed of exocarp and mesocarp. The median mesocarp accumulates lipids only in M. flexuosa and M. armata. The inner mesocarp together with the endocarp becomes papyraceous and tenuous in all species. This internal region of pericarp showed collapsed cells due to seed growth at the end of fruit development. Fruits of Mauritiinae are baccate, and the characters of the pericarp, especially the inner mesocarp and endocarp, help to maintain moisture. On the other hand, many species close to Mauritiinae show pericarp with sclerenchyma adjacent to the seed. This variation can contribute to understand the importance of this striking character in dispersal, germination and colonization in Arecaceae.  相似文献   

6.
The structure and late development of the flowers of the South‐East Asian bee‐pollinated palm Licuala peltata are described with special focus on the architecture of the unusual labyrinthine nectaries. The nectaries are derived from septal nectaries by extensive convolution of the carpel flank surfaces below the ovary throughout the inner floral base, thus also encompassing the inner surface of the corolla–androecium tube. A comparison with septal nectaries elsewhere in Arecaceae and with labyrinthine nectaries in other monocots shows that labyrinthine nectaries situated below the ovary, as described here, are not known from any other palms, but are similar to those of a few Bromeliaceae and, less strongly convoluted, some Haemodoraceae and Xanthorrhoeaceae. In addition, the substantial participation of parts other than the gynoecium in the nectary architecture of Licuala appears unique at the level of monocots. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 66–77.  相似文献   

7.
Observations on the vascular floral anatomy, carpel morphology and floral biology ofHeloniopsis orientalis are presented. The lower flowering pedicel has six large bundles which lack an enclosing sclerenchymatous sheath. At mid-pedicel, branch bundles originate via radial divisions from each of these bundles. Subsequently, there is a vascular ring of 12 bundles below the receptacle. The six smaller bundles which are derived from alternate pedicel bundles eventually establish all of the ventral gynoecium supply. The six larger bundles supply the tepals, stamens and dorsal gynoecial vasculature. The simple dorsals do not branch or fuse in their vertical ascent. The ventral and placental supplies are far more complex. Fusion occurs between paired sets of the six smaller pedicel bundles along the septal radii and results in a submarginal laminal ventral network. An independent ventral plexus is formed in each septum and from each plexus two septal axials, of which the innermost has a reversed xylem-phloem disposition, and four placental bundles are derived. Two placental bundles are associated with each septal axial. Basally the septa are fused centrally, but are freed at mid-gymoecial height. The broadly tri-lobed, tri-carpellate gynoecium is depressed terminally where the erect, hollow style with its capitate stigma is attached. Dorsal grooves are present: the fruit is loculicidally dehiscent. There are no septal glands due to complete lateral fusion of the septal wings. Basally each of the six equal tepals has a saccate nectary. The similarity in vascular anatomy and carpel morphology of the AsianHeloniopsis and eastern North American endemic,Helonias bullata, justifies their position in the same tribe. Research and publication supported in part by the M. Graham Netting Research Fund through a grant from the Cordelia Scaife May Charitable Trust, the U. S.—Japan Cooperative Science Program Grant GF-41367, the Japan Society for the Promotion of Science, and Grant-in-Aid No. 934053 from the Ministry of Education, Japan.  相似文献   

8.
Female flower and fruit anatomy, including vasculature, are studied for the first time in Tetroncium (Juncaginaceae: Alismatales). Other members of Juncaginaceae (and the relatively close Maundiaceae) possess a peculiar type of gynoecium with pronounced carpel fusion via the floral centre. Their carpels are supplied by individual vascular traces and can be interpreted either as synascidiate (if viewed as horizontally inserted) or free and plicate (if viewed as obliquely inserted on an elongated receptacle). In Tetroncium, the gynoecium is tetracarpellate and clearly has a well‐developed synascidiate zone with septa formed by united flanks of adjacent carpels. The gynoecium of Tetroncium is supplied by a common ring of vascular tissue that splits into dorsal and heterocarpellary ventral (synventral) bundles, a condition that can be expected in a typical syncarpous gynoecium. The fruit is indehiscent and contains one or two seeds. The syncarpy of Tetroncium is of functional significance for fruit formation, as it allows the thin septa to be distorted, thus providing more space for the developing seed(s). The occurrence of typical syncarpy in Tetroncium provides further evidence for the highly homoplastic evolution of gynoecium characters in the early‐divergent monocot order Alismatales. Either the similarity between gynoecia of Maundiaceae and Triglochin (Juncaginaceae) is due to parallel evolution or the syncarpy of Tetroncium should be viewed as secondarily derived. In the latter scenario, fusion via the floral centre is probably a synapomorphy of core Alismatales (Helobiae) and more typical syncarpy evolved independently in several lineages, such as Scheuchzeria, Tetroncium and Butomus/Hydrocharitaceae. In total, Tetroncium differs from other Juncaginaceae in 13 structural characters, including ensiform leaves that are similar to those of Tofieldiaceae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 712–724.  相似文献   

9.
Floral structure is compared in Pelagodoxa and Sommieria (Arecaceae, Arecoideae). Male flowers have three free, imbricate sepals, three basally congenitally united and apically valvate petals, and six stamens. Anthers are dorsifixed and dehiscence introrse. The sterile gynoecium is tricarpellate. Female flowers have three free, imbricate sepals and three free, imbricate petals, which are slightly fused with the sepals at the base. Four to six staminodes are congenitally united at the base and fused with the ovary for a short distance. The gynoecium is syncarpous. Carpels are almost equal in early development; later the gynoecium becomes pseudomonomerous. The three stigmatic branches are equally developed, apical and sessile. The carpels are (syn-)ascidiate up to the level of the placenta and (sym-)plicate above. Each carpel has one ovule, in the sterile carpels it is aborted at anthesis. The fertile ovule is erect up to anthesis and pendant afterwards because of the bulging out of the ovary. Pollen tube transmitting tracts (PTTT) encompass the secretory epidermis of the ventral slits of each carpel. Floral structure in Pelagodoxa and Sommieria supports the sister group relationship between the two genera suggested in recent molecular phylogenies and reflects their close relationships to a major clade of pseudomonomerous arecoid palms from the Indo-Pacific region.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 27–39.  相似文献   

10.
The comparative vegetative and reproductive morphology and anatomy of the Malagasy endemic family Sphaerosepalaceae is examined in light of two current competing hypotheses of relationship from recent molecular studies. Sphaerosepalaceae are similar to Thymelaeaceae on the basis of leaf architecture, calyx vasculature and in having endostomal micropyles. Comparisons with Tepuianthus and Thymelaeaceae subfamily Octolepidoideae are drawn on the basis of seed structure, indument type, perianth structure and pollen. Resin-filled, sclerenchymatous idioblasts, floral (positional) monosymmetry, a single series of stamen trunk bundles and a well-developed bixoid chalaza in the seed of Dialyceras parvifolium link Sphaerosepalaceae with its other putative sister group: a clade containing Bixaceae, Cochlospermaceae and Diegodendraceae. Synapomorphies of Sphaerosepalaceae include: fused, intrapetiolar stipules, embryo structure, pollen with endoapertures encompassing the ectoapertures and a tetramerous perianth. The extremely well-developed apical septum in the eusyncarpous gynoecium of Rhopalocarpus suggests that the gynoterminal style present in this genus has been secondarily derived from an ancestor with a fully syncarpous, basistylous gynoecium, as in Dialyceras . The morphological and evolutionary nature of basistylous and apically septate gynoecia is discussed. A rosette arrangement of ovules in each carpel coccus of D. coriaceum expands the bauplan concept of Sphaerosepalaceae and is probably unique among angiosperms as a whole.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 1–40.  相似文献   

11.
Tiquilia is very different from the other members of the Ehretiaceae (Boraginales) in many aspects of morphology and ecology. Because detailed knowledge about flower and fruit traits is necessary to reliably infer character evolution of and within Tiquilia, we investigated flower to fruit ontogeny in eight species of Tiquilia using light and electron microscopy. Tiquilia accumulated a number of autapomorphies such as the prostrate growth form, the lack of lateral and ventral bundles in the gynoecium, and the formation of nutlet‐like mericarpids as dispersal units instead of more or less succulent drupes. The internal architecture of the superior bicarpellate ovary resulted from the development of several secondary septa including apical, basal and false septa, as it has been reported also from other Boraginales. However, no character found in Tiquilia can be regarded as synapomorphic with any other taxon of the Ehretiaceae. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 520–534.  相似文献   

12.
Functional traits play a key role in driving biodiversity effects on ecosystem functioning. Here, we examine the geographical distributions of three key functional traits in New World palms (Arecaceae), an ecologically important plant group, and their relationships with current climate, soil and glacial–interglacial climate change. We combined range maps for the New World (N = 541 palm species) with data on traits (leaf size, stem height and fruit size), representing the leaf–height–seed plant strategy scheme of Westoby, to estimate median trait values for palm species assemblages in 110 × 110‐km grid cells. Spatial and non‐spatial multi‐predictor regressions were used with the Akaike Information Criterion to identify minimum adequate models. Present‐day seasonality in temperature and precipitation played a major role in explaining geographical variation of all traits. Mean annual temperature and annual precipitation were additionally important for median leaf size. Glacial–interglacial temperature change was the most important predictor for median fruit size. Large‐scale soil gradients played only a minor role overall. These results suggest that current climate (larger median trait values with increasing seasonality) and glacial–interglacial temperature change (larger median fruit size with increasing Quaternary temperature anomaly) are important drivers for functional trait distributions of New World palms. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 602–617.  相似文献   

13.
Heterocoma is a Brazilian endemic genus resulting from the dismemberment of Sipolisiinae, in which only representatives with fruit containing phytomelanin were included in the genus. As the fruits of Asteraceae are known to be systematically important at various taxonomic levels and Heterocoma fruit has not been described previously, we studied the morphology and anatomy of the cypselas of all species of the genus, comparing them with other fruits in the family containing phytomelanin and evaluating the systematic potential at the specific and tribal levels. The fruits were analysed by scanning electron microscopy (SEM) and light microscopy. The morphological features of the fruit, including the carpopodium, ribs and pappi, varied in the genus and demonstrated potential for species discrimination. The anatomy showed a pattern for the genus with a uniseriate exocarp, the outer mesocarp composed of fibres arranged in several layers, the inner mesocarp composed of several layers of parenchyma, the endocarp, and phytomelanin deposited between the inner and outer mesocarp. This anatomical pattern of phytomelanin deposition differs from that of other Asteraceae with phytomelanin in their fruit. Heterocoma is also the only genus in Vernonieae that has phytomelanin deposition in the cypselas. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 255–265.  相似文献   

14.
With the aim of correlating the pericarp structure with current phylogenies of Myrteae, this study describes the ontogeny in five species included in five out of the six South American clades of the tribe. In these taxa, the outer and inner ovarian epidermis gives rise to the exocarp and the endocarp, respectively, both with 1 layer. In the mesocarp, derived from the ovarian mesophyll, secretory cavities are arranged into a circle just below the exocarp and near the endocarp in Campomanesia adamantium; only below the exocarp in Eugenia pitanga and Myrcia multiflora; more internally in Myrciaria cuspidata, and below the exocarp and throughout the mesophyll in Myrceugenia alpigena. The promising traits for phylogenetic studies in the group include: direction of elongation of pericarp layers, regions that develop most in relation to the circle of larger vascular bundles, differentiation of spongy and sclerenchymatous tissues and position of secretory cavities.  相似文献   

15.
New comparative data are presented on the reproductive morphology and anatomy of two genera closely related to grasses, Flagellaria and Joinvillea, in which the flowers are superficially similar, especially in stamen morphology. This investigation demonstrates some anatomical differences between the two genera. For example, both genera depart from the ‘typical’ condition of tepal vasculature (three‐traced outer tepals and one‐traced inner tepals): in Flagellaria, each tepal receives a single vascular bundle and, in Joinvillea, each tepal is supplied by three vascular bundles. Joinvillea possesses supernumerary carpel bundles, as also found in the related family Ecdeiocoleaceae, but not in Flagellaria or grasses. In the anther, the tapetum degenerates early in Flagellaria, and is relatively persistent in Joinvillea, in which the pollen grains remain closely associated with the tapetum inside the anther locule, indicating a correlation between peripheral pollen (a feature that is common in grasses) and a persistent tapetum. This study highlights the presence of a pollen‐tube transmitting tissue (PTTT) or solid style in the gynoecium of Flagellaria, as also in many Poaceae, but not in Joinvillea or Ecdeiocoleaceae. We speculate that the presence of a PTTT could represent one of the factors that facilitated the subsequent evolution of the intimately connected gynoecia that characterize grasses. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 393–404.  相似文献   

16.
Xyridaceae belongs to the xyrid clade of Poales, but the phylogenetic position of the xyrid families is only weakly supported. Xyridaceae is divided into two subfamilies and five genera, the relationships of which remain unclear. The development of the ovule, fruit and seed of Abolboda spp. was studied to identify characteristics of taxonomic and phylogenetic value. All of the studied species share anatropous, tenuinucellate and bitegmic ovules with a micropyle formed by the inner and outer integuments, megagametophyte development of the Polygonum type, seeds with a tanniferous hypostase, a helobial and starchy endosperm and an undifferentiated embryo, seed coat derived from both integuments with a tanniferous tegmen and a micropylar operculum, and fruits with a parenchymatous endocarp and mesocarp and a sclerenchymatous exocarp. Most of the ovule and seed characteristics described for Abolboda are also present in Xyris and may represent a pattern for the family. Abolboda is distinguished by the ovule type, endosperm formation and the number of layers in the seed coat, in agreement with its classification in Abolbodoideae. The following characteristics link Xyridaceae to Eriocaulaceae and Mayacaceae, supporting the xyrid clade: tenuinucellate, bitegmic ovules; seeds with a tanniferous hypostase, a starchy endosperm and an undifferentiated embryo; and a seed coat with a tanniferous tegmen. A micropylar operculum in the seeds of Abolboda is described for the first time here and may represent a synapomorphy for the xyrids. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 144–154.  相似文献   

17.
Vegetative and reproductive anatomy and morphology are described for the first time for Tetracarpaea tasmannica Hook., a small shrub endemic to Tasmania. Tetracarpaea, a monotypic genus, has many characteristics of other woody Saxifragaceae, such as wood with solitary pores, scalariform perforation plates, sparse axial xylem parenchyma, tracheids, spiral thickenings in tracheary elements, and perforated ray cells. The tracheary elements of Tetracarpaea are much smaller than those characteristic of the Escallonioideae, a feature probably related to its montane forest habitat. Other features of Tetracarpaea inconsistent with most Escallonioideae include dark-staining deposits in the ray cells; a unilacunar, one-trace nodal pattern; lack of unicellular foliar trichomes; simple craspedodromous venation; areole development that is lacking or incomplete; straight and tapered veinlets; abaxial fibers associated with the foliar vascular bundles; and lack of bundle sheaths. The genus is further characterized by complete, hypogynous, tetramerous flowers. The essentially apocarpous gynoecium has multiovulate carpels, each supplied by three veins that reach the stigma. Ovules are anatropous, bitegmic, and crassinucellate. Lateral sepal bundles are derived either from the sepal midrib or from the petal-plane bundles; stamens are supplied by independent traces or by bundles originating from compound traces in both sepal- and petal-planes. The follicular fruits possess a sclerenchymatous endocarp and contain winged seeds that have a membranous testa, a ridged surface, and a cellular endosperm. Reproductive morphological and anatomical features are more consistent with features of the Saxifragoideae than with the Escallonioideae or the Cunoniaceae, although the essentially apocarpous gynoecium with multiovulate carpels is not found in these groups. Vegetative and reproductive characteristics indicate that Tetracarpaea is more closely related to Saxifragaceae than to Cunoniaceae. It is possible this isolated genus should have separate familial status.  相似文献   

18.
核桃果皮的发育解剖学研究   总被引:6,自引:0,他引:6  
核桃果皮的发育过程可分为3个阶段,发育时期:中、内三层果皮的界线不清,维管束处于发育初期;发育中期:随着中果皮最外侧两层石细胞的出现和薄壁组织细胞体积的迅速扩大以及维管束轮数的增加,使三层果皮具较明显的界面,发育后期:中果皮的维管束递增到4-5轮,  相似文献   

19.
A histological study was made of the systemic growth of Botrytis cinerea from styles, stamens and sepals to the flower receptacle and mesocarp of immature pear fruit. In most styles, hyphal growth ceased in the upper portion at the onset of stylar senescence, which occurred at about 1 wk after full bloom. Hyphae never passed through styles into the carpel. Unlike the styles, hyphae in filaments grew without restriction and progressed within 4 days, via vascular tissue, through sepals into tissues of the upper end of the flower receptacle, or of the mesocarp adjoining the sepals, without causing symptoms. Filaments remained green to partly green until harvest. B. cinerea entered filaments and spread into the receptacle or mesocarp at any time between blossoming and harvest and then became latent in these tissues. Filaments were, however, more susceptible at the flowering stage. After 2 months floral tubes were closed, and the stamens protected from infection. Careful inspection of ripe, cold–stored fruit showed that decay invariably spreads from mesocarp tissue adjoining the sepals, outward along the vascular bundles, but not from secondary inoculum in the floral tube. The behaviour of the pathogen suggests that control of blossom–end rot could be achieved if pears are sprayed with fungicide at 75—100% petal fall (when most stamens are exposed) and a month later (before floral tubes started to close).  相似文献   

20.
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