Structure of the apertural sporoderm of pollen grains inEuphorbia andChamaesyce (Euphorbiaceae)

The tricolporate pollen grains of 38 Mediterranean and Macaronesian species ofEuphorbia L. andChamaesyce S. F. Gray have a special apertural sporoderm not found in the other taxa of theEuphorbiaceae. At the apertural margo the ectexine is thinner because of shorter columellae and the thin, fragmented or even absent foot-layer. Ectexinous granules, mixed with endexinous material, are present near the ora. The endexine is homogeneous and thickened under the colpi (at the end and at the proximity of the end of colpus). Around the ora, the endexine is granulate and lamellar with irregular cavities. The apertural intine presents a characteristic structure with thickenings running along both sides of the colpi. The arrangement and structure of these intinous thickenings depend on the distance from the ora. This special morphology of the intine is present in all taxa studied here. The genusEuphorbia is considered to be the most evolved taxon of this family. The characteristic apertural sporoderm may be an adaptative modification to different physiological conditions, so it may present an apertural mechanism which is more adapted to harmomegathic changes and thus facilitate the germination and the formation of the pollen tube.     

A Phylogenetic Study of Tribe Euphorbieae (Euphorbiaceae)

A phylogenetic investigation of a monophyletic lineage of spurge plants, tribe Euphorbieae, was conducted to elucidate evolutionary relationships, to clarify biogeographic patterns, and to reexamine the previous classification of Euphorbieae. Cladistic analyses of the 52 morphological characters of 61 species resulted in 2922 equally most parsimonious trees of 193 steps with a consistency index of 0.34. The strict consensus tree indicates genus Anthostema of subtribe Anthosteminae as a likely sister group to all other members of tribe Euphorbieae. The morphological data support a monophyletic origin of subtribe Euphorbiinae, but the subtribes Anthosteminae and Neoquillauminiinae did not form monophyletic groups. Although the previous taxonomic treatments within tribe Euphorbieae have supported the generic status of Pedilanthus, Monadenium, Synadenium, Chamaesyce, and Elaeophorbia, the results of this analysis do not support generic placement of them based on cladistic principles. Recognition of these groups as genera results in Euphorbia becoming a paraphyletic group. One solution to this problem in Euphorbieae is to divide the largest genus Euphorbia into several monophyletic genera and to keep the generic ranks for previously recognized genera. The distribution of basal endemic genera in Euphorbieae showed African and east Gondwanan affinities and strongly indicated that the ancestor of Euphorbieae originated prior to the breakup of Gondwanaland from an old group in Euphorbiaceae. However, some recent African taxa of Euphorbia should be interpreted by transoceanic dispersal from the New World ancestors.     

A Phylogeny of Euphorbieae Subtribe Euphorbiinae (Euphorbiaceae) Based on Molecular Data

Phylogenetic relationships within Euphorbiinae were inferred from our analysis of the 3′; end of the chloroplast gene ndhF. A sampling of that subtribe covered 88 species; 3 closely related species from the subtribes Anthosteminae and Neoguillauminiinae and the tiribe Hippomaneae were included as outgroups. A phylogenetic assessment was carried out using the parsimony approach. The relationships revealed via these ndhF data supported the monophyly of subg.Esula, subg.Chamaesyce, subg.Euphorbia, and subg.Lacanthis. However, the polyphyly of subg.Agaloma, subg.Lyciopsis, and subg.Eremophyton also was strongly suggested. The African succulent Euphorbiinae can be divided into primarily two independent groups: 1) spiny succulents, which form a strongly supported clade with three subclades (subg.Euphorbia, subg.Lacanthis, andMonadenium+Synadenium); and 2) non-spiny succulents, which consist of sect.Meleuphorbia, sect.Medusae, sect.Anthacantha, sect.Trichadenia, sect.Pseudeuphorbium, sect.Treisia, and sect.Pseudacalypha. In the ndhF tree, the subg.Esula clade is placed as a sister to the rest of the Euphorbiinae. Thus, the origin of theEuphorbia s.I. should be sought within the herbaceous species of subg.Esula. The core North American endemicEuphorbia groups --Agaloma, Chamaesyce, andPoinsettia — are monophyletic and independent of the South American subg.Agaloma. Instead, they are derived from the AfricanEuphorbia subg.Lyciopsis andEremophyton. The Eurasian subg.Esula clade forms two subclades, which are concordant to sect.Esula and sect.Tithymalus.     

Evolutionary stasis in Euphorbiaceae pollen: selection and constraints

Although much attention has been paid to the role of stabilizing selection, empirical analyses testing the role of developmental constraints in evolutionary stasis remain rare, particularly for plants. This topic is studied here with a focus on the evolution of a pollen ontogenetic feature, the last points of callose deposition (LPCD) pattern, involved in the determination of an adaptive morphological pollen character (aperture pattern). The LPCD pattern exhibits a low level of evolution in eudicots, as compared to the evolution observed in monocots. Stasis in this pattern might be explained by developmental constraints expressed during male meiosis (microsporogenesis) or by selective pressures expressed through the adaptive role of the aperture pattern. Here, we demonstrate that the LPCD pattern is conserved in Euphorbiaceae s.s. and that this conservatism is primarily due to selective pressures. A phylogenetic association was found between the putative removal of selective pressures on pollen morphology after the origin of inaperturate pollen, and the appearance of variation in microsporogenesis and in the resulting LPCD pattern, suggesting that stasis was due to these selective pressures. However, even in a neutral context, variation in microsporogenesis was biased. This should therefore favour the appearance of some developmental and morphological phenotypes rather than others.     

Hydration, sporoderm breaking and germination of Cupressus arizonica pollen

I n vitro and in vivo rehydration and germination in Cupressus arizonica pollen were examined using light and scanning electron microscopy. Shed pollen has 12.6% water content, which reduced to 8.2% after dispersal, and this latter pollen survived for some months at room temperature and for years at −10 °C. Rehydration requires breaking of the sporoderm walls and depends on the composition and pH of the rehydration medium. Acidity restrains the breakage, while alkalinity promotes it. Pollen division follows exine shedding and requires the persistence of the mucilaginous layer; hence, pH values countering these outcomes prevent division. Division results in a large and a small cell separated by a callosic wall. A pollen tube develops from the innermost intine of the large cell, which is callosic, and extends into the mucilaginous middle intine. The percentage germination never exceeded 17% in all tested media. In vivo , pollen rehydrates and casts off the exine in the micropylar drop. Drop withdrawal brings pollen to the apical nucellar cells that degenerate in the meantime, and it leaves a deposit on the surface of the micropylar canal. After contaction of the nucellar cells, the pollen flattens and its mucilaginous layer shrinks and disappears. This occurs simultaneously with sealing of the micropylar canal. During this time, pollen divides asymmetrically without the callosic wall, and the larger cell develops a tube in the interface with the nucellus. Only some pollen grains accomplish adhesion to the nucellus and germinate. The in vitro and in vivo developmental stages are discussed.     

凤仙花属总状花序组的花粉形态及其分类学意义

总状花序组隶属于凤仙花亚属凤仙花属,以具多花的总状花序、侧生萼片2枚、子房5室、果实为线状圆柱形、种子多数、迈卵形、种皮表面具指状突起等特征区别于该亚属的其他类群。前人对凤仙花花粉形态的研究表明凤仙花属的花粉形态对于属下分类及种间界定具有重要的分类学价值。而对于总状花序组来说,有关花粉形态方面的研究少有报道,为了进一步探讨该组花粉S的形态特征及其分类学意义,该研究应用扫描电镜技术对总状花序组的21种植物的花粉进行了观察。结果表明:总状花序组植物的花粉粒均具4条萌发沟,两侧对称,极面观为矩圆形、长矩圆形或近方形;长赤道面观多为椭圆形至矩圆形,短赤道面观为椭圆形;长、短赤道轴比值多在1.4~2.4范围内;表面具网状纹饰,网眼在花粉粒表面均匀分布或赤道面的网眼较极面观的小,网脊边缘光滑或具波状突起,网眼内具不同密度的颗粒状突起。该组的花粉形态与宏观形态性状相关性较小,但花粉极面观形状、长短赤道轴的比值、网眼的分布、网脊边缘形状、网眼内颗粒物密度等特征具有种水平上的稳定性和特异性,对该组植物种间界定具有重要的意义。     

Taxonomic significance of the distribution of constituents of leaf cuticular waxes of Croton species (Euphorbiaceae)

Foliar epicuticular waxes of specimens of 13 Croton species native in Brazil were extracted. The fractions containing alkanes and primary alcohols were isolated by preparative thin layer chromatography. Derivatized n-primary alcohols were identified by gas chromatography (GC) coupled with mass spectrometry and n-alkanes by GC and comparison with known standards. Relative abundances were estimated by GC coupled with flame ionization detector. The distribution of constituents of both classes was analyzed by cluster analysis, using the UPGMA method and Euclidean distances. The chemical affinities among species were compared with published data of molecular phylogenetic relationships. The distribution of n-alkanes and primary alcohols were shown to be useful markers of Croton species. Primary alcohols were more consistent than n-alkanes for species fingerprinting.     

Taxonomic significance of seed sculpture and pollen ecto-mycoflora at the infraspecific level: Brassica tournefortii case study

Several populations of Brassica tournefortii (Brassicaceae) occurring in Egypt are investigated from the micromorphological point of view (seed ornamentations). The species is known to show a notable phenotypic plasticity and five morphotypes was identified in the past. Furthermore, a soil analysis as well as a study of the fungal species from anthers were carried out. The aim of the study is to verify the taxonomic value of the morpholotypes of B. tournefortii and their ecologic relationship with soil variables. The results obtained demonstrated that the five morphological forms can be distinguished based on the seed sculpture. The Canonical Correspondence Analysis (CCA) exhibited a clear correlation between the soil variables and the identified forms. Six species of fungi were detected from the ecto-anthers in the Forms (F2-F4), while F1 was lacking the fungal species. The study revealed that the morphological plasticity of studied B. tournefortii depends on ecological factors.     

Taxonomic significance of pollen types in the Guyana Highland-centred composite genera of Mutisioideae (Asteraceae)

The Guyana Highland-centred genera, or Stenopadus group, are a complex of species that belong to the tribes Mutisieae and Stifftieae of Asteraceae. The pollen morphologies of 29 species, from 12 of 13 genera of this complex, are described and illustrated using light and scanning electron microscopy. The exine sculpture and structure are highly significant. Four exine types, previously characterized, were distinguished: Gongylolepis , Wunderlichia , Mutisia , and Stenopadus . The characterization of the Stenopadus exine type is enlarged here. These exine types led to the recognition of four well-defined pollen types, whereas the spine length and exine thickness characterized six subtypes. Pollen types circumscribe genera or groups of genera, and some subtypes distinguish species. The pollen morphology within the complex is discussed in relation to the rest of Mutisioideae and other palynologically allied tribes of Cichorioideae. There is little correlation between pollen types and tribes; only the Stenopadus exine type is exclusive to the Stifftieae tribe. The remaining types are shared by the two tribes of the complex. Pollen morphology supports the hypothesis that this group of genera is close to the Gochnatia complex and the Cardueae tribe.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 327–340.     

On some peculiarities of sporoderm structure in members of the Cycadales and Ginkgoales

The pollen morphology and ultrastructure of Cycas micholitzii, C. simplicipinna, Cycandra profusa, Ceratozamia mexicana, and Ginkgo biloba are studied. Pollen germination is also studied in C. mexicana and G. biloba. Although dehydrated pollen grains appear monosulcate, the study of hydrated pollen shows that the aperture occupies nearly half of the pollen surface and represents a pore rather than a sulcus. In the Ginkgoales, the inaperturate ectexine is characterized by a thick solid tectum, infratectum of columella-like elements or large granules, and distinct foot layer. On the contrary, in the Cycadales, the ectexine consists of a thin tectum, alveolar infratectum, and poorly discernable foot layer. Members of the Ginkgoales have a distinct distal aperture, which is constituted by an intine, endexine, and thin ectexine. In the modern Cycadales, an ectexine is well developed throughout the pollen perimeter; in the supposed aperture region the ectexine is not reduced in thickness, although it is characterized by a thinner tectum and thinner walls of infratectal alveoli. In Cycandra profusa, no unequivocal aperture region has been found. Thickened regions were observed in the intine of both the Cycadales and Ginkgoales.     

The pollen morphology of Phyllanthus (Euphorbiaceae) section Choretropsis

The pollen grain morphology of the eight South American species of the genus Phyllanthus , subgenus Phyllanthus , section Choretropsis , endemic to Brazil, was investigated using light and scanning electron microscopy. The high homogeneity in pollen shape, as well as aperture number and configuration, allowed the definition of a basic pollen type for this section, viz. prolate spheroidal or subprolate, zonocolporate with 4–5 invaginated colpi, sexine reticulate. Two subtypes have been observed, based on pollen grain apertures and exine sculpture. The first subtype is characterized by straight and narrow colpi provided with costae, lalongate endoapertures and a reticulum of rectilineous muri as in P. choretroides and P. spartioides . The second subtype has wider colpi without costae, circular endoapertures and a reticulum with curvilineous muri as in P. flagelliformis , P. goianensis and P. scoparius , or curvilineous-rectilineous muri as in P. angustissimus , P. gladiatus and P. klotzschianus . It was demonstrated that the pollen shape and the number of colpi varied within and between some of the species. The results suggest that pollen grain morphology, in association with plant architecture and the presence of phylloclades, constitute additional evidence to delimit the species of the section Choretropsis and to further understand their evolutionary origin within the genus Phyllanthus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 243–250.     

Allergenic and antigenic proteins released in the apertural sporoderm during the activation process in grass pollen grains

A combination of transmission electron microscopy with immunocytochemical methods was used to localize antigenic and allergenic proteins during the maturation and activation processes of Poaceae pollen grains. The intine undergoes a series of modifications that play a decisive role in these processes. Allergenic and antigenic proteins were detected particularly in the intine of activated in vitro grass pollen grains. Labelling of antigenic proteins was more abundant and less specific than that of allergenic proteins. At the time of hydration, the operculum was lifted up, the intine was swollen and labelling of allergenic proteins appeared highly localized in the Zwischenkörper. No significant labelling was observed when the Zwischenkörper gelatinized. Immunolocalization of allergenic proteins in the activated Zwischenkörper indicated the presence of proteins related to activation of the pollen grains. This confirms that the intine function is involved in the processes of pollen tube formation and fertilization, and also suggests the possible mechanism activated in the pollen grains when allergenic proteins reach the mucosa of sensitive subjects.     

Taxonomic revision and phytogeographic studies in Euphorbia (Euphorbiaceae) in the Khorassan provinces of Iran

This study is focused on the genus Euphorbia L. in a part of northeast Iran, viz. the three Khorassan provinces. Since there are many taxa of Euphorbia in Iran which are used in different industries and have significant effects on human and non‐human life it is important to revise their taxonomy. With about 90 species, following Turkey with 91 species, Iran is the second richest country for Euphorbia in Asia. Of these, 30 species are distributed in the Khorassan provinces. This is the first comprehensive work on the genus in this region. According to ‘Flora Iranica’, there are 17 species of Euphorbia in northeast Iran, while according to our results, there are 30 species of Euphorbia in the Khorassan provinces alone. In addition to various new taxonomic and biogeographic results, a new species, viz. E. chamanbidensis, is described. Euphorbia chamanbidensis is closely related to E. aucheri, but seed micro‐morphological characters differentiate them. Two identification keys to the Euphorbia species of the studied area are provided, one based on macro‐morphological characters and another based on seed micro‐morphological characters. Phytogeographic analysis and distribution maps for all species are also presented.     

Anther structure and pollen ontogeny of Hosta Tratt. and its systematic significance

Anther structure and pollen ontogeny of six species of Hosta Tratt. was studied to provide additional data for its classification. Microscopic observation found that their male gametogenesis has the following structural characteristics: (1) the four microsporangia are linearly arranged on introrse side of anther connective tissue; (2) mature anther wall is composed of an epidermis, endothecium, middle layer(s), and tapetum; (3) microsporogenesis is successive; (4) mature pollen grains are three-celled at anthesis, with a vegetative cell filled with large amounts of starch grains; (5) pollen ontogeny is obviously asynchronous; (6) each tepal has about 9?C13 vascular traces in cross-section view. The above observations suggest that genus H. Tratt. should be classified as family Hostaceae, in agreement with Dahlgren and Clifford (1982), Takhtajan (1997), and Wu et?al. (2003).     

Taxonomic significance of trichomes micromorphology in cucurbits

Studies on trichomes micromorphology using Scanning Electron Microscope (SEM) were undertaken in 23 species with one variety under 13 genera of the family Cucurbitaceae (viz., Benincasa hispida (Thunb.) Cogn., Citrullus lanatus (Thunb.) Matsum. & Nakai, Cucumis melo var. agrestis Naudin, Cucumis sativus L., Diplocyclos palmatus (L.) C. Jeffrey, Edgaria dargeelingensis C.B. Clarke, Gynostemma burmanicum King ex Chakr., Gynostemma pentaphyllum (Thunb.) Makino, Gynostemma pubescens (Gagnep.) C.Y. Wu, Hemsleya dipterygia Kuang & A.M. Lu, Lagenaria siceraria (Molina) Standl., Luffa acutangula (L.) Roxb., Luffa cylindrica M. Roem., Luffa echinata Roxb., Melothria heterophylla (Lour.) Cogn., Melothria leucocarpa (Blume) Cogn., Melothria maderspatana (L.) Cogn., Sechium edule (Jacq.) Sw., Thladiantha cordifolia (Blume) Cogn., Trichosanthes cucumerina L., T. cucumerina var. anguina (L.) Haines, Trichosanthes dioica Roxb., Trichosanthes lepiniana (Naudin) Cogn. and T. tricuspidata Lour.). The trichomes in the family Cucurbitaceae vary from unicellular to multicellular, conical to elongated, smooth to ridges, with or without flattened disk at base and cyctolithic appendages, thin to thick walled, curved at apices to blunt. Trichomes micromorphology in the family Cucurbitaceae was found significant taxonomically.     

Taxonomic significance of leaves in family Aizoaceae

Aizoaceae is one of the most important and widespread succulent plant families in both tropical/subtropical regions and arid zones. In this study, 27 species were collected from various floristic regions in Egypt, Kingdom of Saudi Arabia and cactus farms (Kalupia – Egypt). The morphological characteristics of every taxon were recorded. The important morphological features included: the number of leaves per plant; leaf types; leaf position (cauline or radical; the latter indicates leaves arising from, or near, the roots); leaf arrangement; petiolate or sessile leaves; leaf sheath present or absent; leaf shape; leaf margin; leaf tip; presence of leaf ‘window area’; leaf texture; and presence of white or dark miniscule dots (white miniscule dots from calcium carbonate and dark miniscule dots from tannin sacs). The investigated anatomical features were as follows: shape of the transverse section; the type of epidermal cells; the presence of large epidermal cells (bladder cells); presence of papilla and simple hairs; presence of tannin sacs; shapes of calcium oxalate crystals; shape of the xylem vessels; and the presence of Kranz unit (the unit that constitutes the vascular bundle/s, parenchyma sheath, and surrounding mesophyll) or collenchyma sheath. All data were recorded in a data matrix (as either text or numerical data), which was used to construct the identification key and phylogeny tree using a multi-variate statistical package. The results of our analysis may open the possibility of using the morphological and anatomical features of leaves to distinguish between the subfamilies, genera, and species of Aizoaceae.     

Taxonomic significance of the inner structure of the ovary in the genusImpatiens (Balsaminaceae)

The inner structure of the ovary inImpatiens was examined from the viewpoint of taxonomy, based on the materials obtained during the 1979 Botanical Expedition to Thailand. The species studied areI. kerriae, I. larsenii, I. hongsonensis, I. psittacina, I. salaengensis, I. charanii, I. kanburiensis andI. macrosepala. It is worthy of special mention that they are all characterized by 4 lateral sepals, large round outer sepals, a nipple-like short spur of the lip and connate wing petals. Contrary to the prevailing information about the ovary ofImpatiens, these Thai species were found to have 4-locular and 4-carpellate ovaries. The arrangement and the number of ovules in a loculus show a tendency of reduction from biseriate to uniseriate and from several to single or none. The species of more advanced type have smaller numbers of ovules and often sterile loculi of the ovary, as represented byI. macrosepala. The types of ovule development were proved to correspond to the degree of specialization of floral characters. Supported by a Grant in Aid of Scientific Research of Ministry of Education, No. 404126 and No. 504352, and partly reported at the Meeting of the Japan Society of Plant Taxonomists in March 1981.     

Ovules and seeds in Acalyphoideae (Euphorbiaceae): structure and systematic implications

Acalyphoideae, the largest subfamily of Euphorbiaceae, are investigated with respect to ovule and seed structure on the basis of 172 species of 80 genera in all 20 tribes of Acalyphoideae sensu Webster. All species of Acalyphoideae examined have bitegmic ovules with a non-vascularized inner integument. However, noticeable differences exist among and sometimes within the genera in the thickness of the inner and outer integument, the presence or absence of vascular bundles in the outer integument, whether ovules are pachychalazal or not, the presence or absence of an aril, seed coat structure (in terms of the best-developed mechanical cell-layer), and the shape of cells constituting the exotegmen. For the latter two characters, two different types of seed coat (i.e., "exotegmic" and "exotestal") and three different types of exotegmic cell (i.e., palisadal, tracheoidal and ribbon-like) were distinguished. Comparisons showed that three tribes Clutieae, Chaetocarpeae and Pereae are distinct from the other Acalyphoideae as well as from the other Euphorbiaceae in having an exotestal seed coat with a tracheoidal exotegmen. The tribe Dicoelieae is also distinct from the other Acalyphoideae in having an exotegmic seed that is composed of ribbon-like cells of exotegmen (i.e., cells both longitudinally and radially elongated, sclerotic and pitted). The tribe Galearieae, which should be treated as a distinct family Pandaceae, is also distinct from the other Acalyphoideae in having an exotegmic seed with a tracheoidal exotegmen (i.e., cells longitudinally elongated, sclerotic and pitted). The remaining genera of Acalyphoideae always have an exotegmic seed with a palisadal exotegmen (i.e., cells radially elongated, sclerotic and pitted). The shared palisadal exotegmen supports the close affinity of Acalyphoideae (excluding five tribes) with Crotonoideae and Euphorbioideae. Within the remaining genera of Acalyphoideae, a significant diversity is found in ovule and seed morphology with respect to the thickness of the inner and outer integument, the size of chalaza, vascularization of an outer integument and an aril.     

Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications

Ovule and seed structure in Euphorbioideae, one of the five euphorbiaceous subfamilies, is surveyed to evaluate its systematic implications on the basis of 79 species representing four of five tribes. All Euphorbioideae, like two other "uniovulate" subfamilies Acalyphoideae and Crotonoideae, but unlike most of two "biovulate" subfamilies Oldfieldioideae and Phyllanthoideae, consistently have a persistent and palisadal exotegmen composed of radially elongate, sclerotic, and pitted cells. Within Euphorbioideae, the tribe Stomatocalyceae (also with the palisadal exotegmen) is unusual in having vascular bundles in outer integument and clearly distinct from the remaining Euphorbioideae and the other "uniovulate" subfamilies. With the exclusion of Stomatocalyceae, Euphorbioideae are not anatomically divided into major groups such as a pseudanthial and a non-pseudanthial clade, but instead have some remarkable diversity within a tribe, a subtribe, and even a genus in the three ovule and seed characters: (1) the thickness of the inner integument, (2) the thickness of the outer integument, and (3) the presence or absence of an aril. Groups of genera and species wrapped by different combinations of their characteristics, however, are not necessarily harmonized with tribal or subtribal classifications available. Anatomical similarities and dissimilarities presented in this paper, as well as relationships among taxa presented in the classifications available, will be critically evaluated in the light of results of ongoing molecular phylogenetic analyses. Electronic Publication