PATTERNS OF ENDOTHECIAL WALL THICKENINGS IN ARACEAE: SUBFAMILIES POTHOIDEAE AND MONSTEROIDEAE

A survey of the patterns of endothecial wall thickenings in 106 representative species from 20 genera in the Pothoideae and Monsteroideae was made using cleared anthers, sections and macerations. The wide variety of wall thickenings that is present is based on an annular-helical pattern. Variations in thickenings are related to differences in cell shape, cell orientation, intergradation between helical and annular patterns, pitch of helices, presence of branched thickenings, and various types of discontinuities in thickenings. Notable exceptions to the annular-helical pattern include Culcasia, which lacks a differentiated endothecial layer with thickenings, and Acorus, which has a peculiar stellate pattern that is unique in the family. No single pattern consistently characterizes either subfamily, although continuous helices are common in the Monsteroideae, and rare in the endothecium of Pothoideae (except Anadendrum). Monsteroideae frequently exhibit a series of slanted separate thickenings on anticlinal walls, which is absent from Pothoideae except in Heteropsis. The slanted pattern is considered a variation on a rectangular helix, involving discontinuities of thickenings on the periclinal walls. Some monsteroid genera show considerably more interspecific variation (Rhaphidophora) than others (Monstera). Endothecial thickenings constitute an anatomical character that is useful in the systematic study of Araceae; present results support other anatomical studies in identifying Culcasia and Acorus as highly divergent genera in the Pothoideae.     

CLADISTIC ANALYSIS OF PATTERNS OF ENDOTHECIAL THICKENINGS IN THE POALES/RESTIONALES

The three-dimensional structure of the endothecial thickenings in the anthers was investigated in 87 species from 70 genera, chosen to provide representative coverage of the families Cyperaceae, Restionaceae, Anarthriaceae, Ecdeiocoleaceae, Centrolepidaceae, Joinvilleaceae, Flagellariaceae, Poaceae, Xyridaceae, and Eriocaulaceae. There is complex variation in the patterns of endothecial thickening: the Eriocaulaceae, Flagellariaceae, and most Poaceae have thickenings with a complete baseplate; the Cyperaceae and most of the Restionaceae are characterized by helical thickenings; some genera in the Bambusoideae have annular thickenings; and U-shaped thickenings occur in the Xyridaceae and Eriocaulaceae and in some Poaceae and Restionaceae. Joinvillea and Ecdeiocolea have unique thickening types. Endothecial characters were subjected to cladistic analysis. Including endothecial characters in an existing cladogram of the group indicates that there is no single, well-corroborated cladogram available for the Poales/Restionales.     

Arabidopsis MYB26/MALE STERILE35 regulates secondary thickening in the endothecium and is essential for anther dehiscence

The Arabidopsis thaliana MYB26/MALE STERILE35 (MS35) gene is critical for the development of secondary thickening in the anther endothecium and subsequent dehiscence. MYB26 is localized to the nucleus and regulates endothecial development and secondary thickening in a cell-specific manner in the anther. MYB26 expression is seen in anthers and also in the style and nectaries, although there is no effect on female fertility in the ms35 mutant. MYB26 expression in anthers occurs early during endothecial development, with maximal expression during pollen mitosis I and bicellular stages, indicating a regulatory role in specifying early endothecial cell development. Overexpression of MYB26 results in ectopic secondary thickening in both Arabidopsis and tobacco (Nicotiana tabacum) plants, predominantly within the epidermal tissues. MYB26 regulates a number of genes linked to secondary thickening, including IRREGULAR XYLEM1 (IRX1), IRX3, IRX8, and IRX12. Changes in expression were also detected in two NAC domain genes, NAC SECONDARY WALL-PROMOTING FACTOR1 (NST1) and NST2, which have been linked to secondary thickening in the anther endothecium. These data indicate that MYB26 regulates NST1 and NST2 expression and in turn controls the process of secondary thickening. Therefore, MYB26 appears to function in a regulatory role involved in determining endothecial cell development within the anther and acts upstream of the lignin biosynthesis pathway.     

ENDOTHECIUM IN IRIDACEAE AND ITS SYSTEMATIC IMPLICATIONS

An examination of the endothecial thickenings in 44 species of Iridaceae, selected from the four subfamilies and all major tribes, provides useful information about generic and tribal relationships in the family. U-shaped thickenings occur in Nivenioideae and Iridoideae—Sisyrinchieae, the latter the least specialized tribe of its subfamily. The occurrence of helical thickenings in all members examined of Iridiodeae tribes Irideae, Mariceae, and Tigridieae (a putatively monophyletic group) and Ixioideae is consistent with the recognition of these two lines as distinct taxa based on anatomical, morphological, phytochemical, and in the case of Ixioideae, palynological criteria. Baseplate thickenings are restricted to Patersonia. However, the shrubby Cape genera—Nivenia, Klattia, and Witsenia—have U-shaped thickenings which show a tendency for the bars on the inner periclinal cell walls to anastomose, suggesting a trend towards the baseplate condition in Patersonia. This accords with a suggested relationship between these genera, based on anatomical and flavonoid similarities. The pattern of variation in endothecial thickenings in Iridaceae is consistent with the phylogeny proposed by Goldblatt (1990). The distribution of thickening types within the family does not make it possible to polarize this character, but the most parsimonious interpretation assumes that U-shapes are basic. However, in at least some other monocotyledonous families the pattern suggests that U-shaped thickenings are derived from helices.     

ENDOTHECIAL THICKENINGS IN ANTHERS OF PORATE MONOCOTYLEDONS

A survey of the presence of endothecial thickenings in 20 genera from 11 families of porate monocotyledons tested the long-standing hypothesis that porate anther dehiscence is correlated with the absence of thickenings. Although this hypothesis is widely repeated in texts, it has been supported by a limited, biased data base from studies of Ericaceae and Melastomataceae, two large families of dicotyledons. The results of our broad study of monocotyledons show that endothecial thickenings are present in most porate monocotyledons. Thickenings are absent in only two of the 11 porate families surveyed, Mayacaceae and Commelinaceae, both in the Commelinales. Thickenings are also absent in Xyris (Xyridaceae), a nonporate genus. These findings indicate that in monocotyledons porate anther dehiscence and the absence of thickenings are not as closely related as previously supposed.     

Characterization and genetic mapping of a mutation (ms35) which prevents anther dehiscence in Arabidopsis thaliana by affecting secondary wall thickening in the endothecium

The male sterile mutant, ms35 , of Arabidopsis thaliana was produced by X-irradiation of seeds. The mutant produces fertile pollen, but is male sterile because the anthers do not dehisce. Anther development in ms35 plants occurs as in wild-type Arabidopsis until shortly after microspores are released from meiotic tetrads. Thereafter, in the wild type, bands of lignified, cellulosic secondary wall thickenings are laid down around the cells of the anther endothecium. In contrast, wall thickenings are not formed in the endothecium of the ms35 mutant. Development of other lignified tissues, for example the vascular tissue of the stamen, occurs normally in ms35 plants. In mutant anthers, as pollen maturation is completed, the stomium is cleaved but the anther wall does not retract to release pollen. The block in anther dehiscence in ms35 plants is specifically correlated with the absence of endothecial wall thickenings. The ms35 mutation represents the first genetic evidence in support of the proposed role of the endothecium in anther dehiscence. The ms35 gene was mapped to the top arm of chromosome 3 ( hy2 -(4.17±2.31 cM)- ms35 -(32.14±5.45 cM)- gl1 ).     

Additional Notes on the Begonia Sect. Petermannia (Begoniaceae) from China

Begonia pellionioides YM. Shui & WH. Chen, a new species of the genus Begonia L. (sect. Petermannia (Klotzsch) A. DC., Begoniaceae), is described and illustrated. The new species is endemic to limestone areas in Southeastern Yunnan Province, China, and is most similar to Begonia hainanensis Chun & F. Chun but distinguished by its more elongated leaves with cuneate base and acuminate to caudate apex, tepals adaxially with spiny hairs and sharply triangular capsule wings with truncate superior margin. Besides, the unknown flower morphology of previously published Begonia sublongipes YM. Shui in Begonia sect. Petermannia is described.     

国产秋海棠属(秋海棠科)花粉形态及其分类学意义(英文)

秋海棠属是世界有花植物第六大属,是被子植物分类困难的类群之一,亟需增加形态性状的比较研究,以便于今后对该属开展分类学修订。该研究选取国产秋海棠属中较为常见的侧膜组、秋海棠组、单座组和二室组共21种,应用扫描电镜观察花粉微形态,探讨花粉形态对秋海棠属植物的分类学意义。结果表明:秋海棠属植物的花粉多为单粒花粉,辐射对称,等极,三孔沟,超长球形到长球形。选取9个稳定的花粉特征进行无序和不加权的性状编码,应用非线性多维标度分析对花粉特征矩阵进行聚类分析,结果支持这些组都不是单系类群需要重新修订,其中花粉边缘形状、极面观轮廓、萌发沟和花粉的外壁纹饰具有一定分类学意义,特别是塞缘特征具有重要的分类意义。根据塞缘特征可以将研究类群区分为2个类群:(A)无塞缘或塞缘光滑;(B)塞缘颗粒状。类群B中具规则颗粒状和精细颗粒状塞缘的种类聚在一起,而具粗糙颗粒状塞缘的种类位于类群A和B的中间,很可能是2个类群的过渡性状,这需要增加取样做进一步的研究。     

中国秋海棠属 (秋海棠科) 植物的DNA条形码评价

秋海棠属植物种类繁多,形态变异多样,导致种类的系统放置混乱,近缘种类鉴定困难。利用DNA条形码实现物种快速准确的鉴定技术具有不受形态特征约束的优势,为秋海棠属植物的分类鉴定提供了新的方法。本研究选择4个DNA条形码候选片段（rbcL,matK,trnH psbA,ITS）对中国秋海棠属26种136个个体进行了分析。结果显示：叶绿体基因rbcL,matK和trnH psbA种内和种间变异小,对秋海棠属植物的鉴别能力有限;ITS/ITS2种内和种间变异大,在本研究中物种正确鉴定率达到100%/96%,可考虑作为秋海棠属DNA条形码鉴定的候选片段。研究结果支持中国植物条形码研究组建议将核基因ITS/ITS2纳入种子植物DNA条形码核心片段中的观点。     

Two New Records of Begonia L. (Begoniaceae) from China

Two species of Begonia L. (Begoniaceae), viz. Begonia demissa Craib and Begonia subperfoliata Parish ex Kurz are reported as new records in China, and the two species assigned to Begonia sect. Parvibegonia A. DC. and Begonia sect. Diploclinium (Wight) A. DC. respectively. The type locality of Begonia demissa is in Thailand and of Begonia subperfoliata in Myanmar. The distribution of Begonia demissa is now extended to Mengyuan, Mengla County of Xishuangbanna Dai Autonomous Prefecture and that of Begonia subperfoliata to Liushun Community, Pu’er City, both in Yunnan province, China.     

Phylogenetic position and biogeography of Hillebrandia sandwicensis (Begoniaceae): a rare Hawaiian relict

The Begoniaceae consist of two genera, Begonia, with approximately 1400 species that are widely distributed in the tropics, and Hillebrandia, with one species that is endemic to the Hawaiian Islands and the only member of the family native to those islands. To help explain the history of Hillebrandia on the Hawaiian Archipelago, phylogenetic relationships of the Begoniaceae and the Cucurbitales were inferred using sequence data from 18S, rbcL, and ITS, and the minimal age of both Begonia and the Begoniaceae were indirectly estimated. The analyses strongly support the placement of Hillebrandia as the sister group to the rest of the Begoniaceae and indicate that the Hillebrandia lineage is at least 51-65 million years old, an age that predates the current Hawaiian Islands by about 20 million years. Evidence that Hillebrandia sandwicensis has survived on the Hawaiian Archipelago by island hopping from older, now denuded islands to younger, more mountainous islands is presented. Various scenarios for the origin of ancestor to Hillebrandia are considered. The geographic origin of source populations unfortunately remains obscure; however, we suggest a boreotropic or a Malesian-Pacific origin is most likely. Hillebrandia represents the first example in the well-studied Hawaiian flora of a relict genus.     

广西秋海棠属(秋海棠科)4新种

描述了广西秋海棠属(秋海棠科) 的侧膜胎座组和秋海棠组4新种。     

Four new species of Begonia L. (Begoniaceae) from Guangxi,China

Four new species of the genus Begonia L. (Begoniaceae) from Guangxi, China are described and illustrated.     

A novel F-box protein represses endothecial secondary wall thickening for anther dehiscence in Arabidopsis thaliana

In plants, the regulation of protein turnover by the ubiquitin proteasome system (UPS) is a key posttranslational mechanism underlying diverse cellular processes. However, the participation of the UPS in cellular processes involved in anther dehiscence, especially endothecial secondary wall (ESW) thickening, has not been characterized. Here, we report that a novel F-box protein in arabidopsis, designated SAF1 (Secondary wall thickening-Associated F-box 1), negatively regulates ESW thickening in the anther. SAF1 is predominantly expressed in flower tissues and interacts with Arabidopsis-Skp1-like 19 (ASK19). SAF1-overexpressed (Ox) lines showed reduced fertility due to a lack or loss of ESW thickening in the anther and inhibition of the expression of relevant genes, such as IRREGURAR XYLEMs (IRXs) in flowers. These findings suggest that the novel Skp/Cul/F-box (SCF) complex consisting of SAF1 as an F-box protein and ASK19 as a Skp functions in secondary wall thickening of the anther endothecium.     

A comparative ultrastructural study of pit membranes with plasmodesmata associated thickenings in four angiosperm species

Recent micromorphological observations of angiosperm pit membranes have extended the number and range of taxa with pseudo-tori in tracheary elements. This study investigates at ultrastructural level (TEM) the development of pseudo-tori in the unrelated Malus yunnanensis, Ligustrum vulgare, Pittosporum tenuifolium, and Vaccinium myrtillus in order to determine whether these plasmodesmata associated thickenings have a similar developmental pattern across flowering plants. At early ontogenetic stages, the formation of a primary thickening was observed, resulting from swelling of the pit membrane in fibre-tracheids and vessel elements. Since plasmodesmata appear to be frequently, but not always, associated with these primary pit membrane thickenings, it remains unclear which ultrastructural characteristics control the formation of pseudo-tori. At a very late stage during xylem differentiation, a secondary thickening is deposited on the primary pit membrane thickening. Plasmodesmata are always associated with pseudo-tori at these final developmental stages. After autolysis, the secondary thickening becomes electron-dense and persistent, while the primary thickening turns transparent and partially or entirely dissolves. The developmental patterns observed in the species studied are similar and agree with former ontogenetic studies in Rosaceae, suggesting that pseudo-tori might be homologous features across angiosperms.     

Floral micromorphology and its systematic implications in the genus Sinosenecio (Senecioneae-Asteraceae)

Several floral microcharacters (configuration of stigmatic areas on the inner surface of the style branch, shape of the anther apical appendage, anther size, and configuration of the endothecial thickenings and of the filament collar) in 63 populations representing 35 species and one variety of Sinosenecio (Senecioneae-Asteraceae) were investigated. The floral micromorphological data obtained are highly consistent with evidence from molecular systematics and cytology, strongly suggesting a polyphyletic nature of the currently circumscribed Sinosenecio and the necessity of a taxonomic change at generic level.     

中国秋海棠属植物的叶表皮特征及其分类学意义

在光学显微镜下,对中国秋海棠属(Begonia)植物7组52种2变种的叶表皮进行观察。结果表明秋海棠属植物叶表皮形态在属内组间具有较大的相似性,表皮细胞为多边形或近多边形,垂周壁平直或弓形,大多数种类表皮细胞内具有晶体,气孔器仅分布于下表皮,且以不等型为主。叶表皮综合特征,例如表皮细胞形状,表皮毛类型,表皮细胞内晶体的类型和形态,气孔器形态以及与一些种类独有特征的组合,在种间,尤其在近缘种之间具有明显的差异。     

Pleistocene and pre-Pleistocene Begonia speciation in Africa

This paper presents a historical biogeographic analysis of African Begonia based on combined internal transcribed spacer (ITS) and trnL intron sequences. Age range estimates for Begonia in Africa ranged from only 1.5 Ma for some terminal nodes to 27 Ma for basal nodes when the ages of Réunion (2 Ma) andMayotte (5.4 Ma) were used to date the split between Begonia salaziensis and Begonia comorensis. Assuming a more recent origin age for Begonia salaziensis (2 Ma) provided age estimates in other parts of the phylogeny which agreed with patterns observed in other African organisms. A large proportion of the Begonia diversity seen today in Africa is of pre-Pleistocene origin. Species of Pleistocene origin are concentrated in species-rich groups such as sections Loasibegonia, Scutobegonia, and Tetraphila, which have their centre of diversity in western Central Africa. Phylogenetically isolated taxa such as Begonia longipetiolata, Begonia iucunda, and Begonia thomeana date to the late Miocene, a period of extended aridification on the African continent that had severe effects on African rain forest species. A general pattern is identified where phylogenetically isolated species occur outside the main identified rain forest refuges. Endemic species on the island of S?o Tomé such as Begonia baccata, Begonia molleri, and Begonia subalpestris appear to be palaeoendemics. Of these species, the most recent age estimate is for B. baccata, which is dated at ca. 3 Ma. Therefore, S?o Tomé appears to have functioned as an important (if previously unrecognised) pre-Pleistocene refuge. On the mainland, areas such as the Massif of Chaillu in Gabon, southern Congo (Brazzaville), and far western areas of Congo (Kinshasa) have played similar roles to S?o Tomé.