中国特有属藤山柳属(猕猴桃科)的花药和胚珠发育及其分类学意义

首次报道了刚毛藤山柳（Clematoclethra scandens subsp.scandens）的花药和早期胚珠发育的胚胎学特征：花药4室;发育完整的花药壁6层,为基本型花药壁,由外至内分别为表皮、药室内壁、两层中层和两层绒毡层;腺质型绒毡层为2核或3核,纤维性加厚发生于药室内壁;小孢子母细胞减数分裂为同时型胞质分裂,小孢子四分体多为四面体型;成熟花粉粒为2-细胞型,3孔沟或稀4孔沟。早期胚珠为近倒生,单珠被,薄珠心,珠孔由单珠被构成。比较胚胎学研究认为藤山柳属和猕猴桃科的其它两个属都具有单珠被胚珠及其他相似的花药发育特征,这些特征和形态分类系统范畴的第伦桃科和山茶科有明显区别。因此,本文支持花粉形态学、细胞学和分子系统学的观点,认为藤山柳属与水东哥属和猕猴桃属构成一个较好的单系,一并放在猕猴桃科。     

Anther wall development,placentation, sporogenesis and gametogenesis in Smilax davidiana A.DC: A contribution to the embryology of Smilax

Embryological characters can be used to address taxonomic relationships and complement molecular phylogenetics and are of special value at the genus level. However, embryological information is fragmentary in Smilax and completely unknown in Smilax davidiana, a Chinese species. Anther wall development, placentation, sporogenesis and gametogenesis of S. davidiana are characterized here. The anther is bisporangiate, anther wall formation is of the Dicotyledonous type, both epidermis and endothecium develop fibrous thickenings, and the tapetum is secretory and of dual origin. Cytokinesis in the microsporocyte meiosis is successive, the microspore tetrad is tetragonal, and mature pollen is two-celled. The ovary is mostly trilocular with an axile placentation (a small fraction of the ovaries are unilocular with parietal placentation), the ovule is anatropous, bitegmic and crassinucellate, with embryo sac development of the Polygonum type. This study documents for the first time the embryological characters of S. davidiana in detail and contributes much to the embryology of Smilax.     

Embryology of theaceae — Anther and ovule development ofAdinandra,Cleyera andEurya

Anther and ovule development of the theaceous Ternstroemioideae is reported for the first time on the basis of eight specles of three generaAdinandra, Cleyera andEurya. Anthers of these three genera are similar and can be characterized by the following traits: tapetum of glandular type, anther dehiscing latrorse-introrse, both connective and anther epidermis heavily tanniniferous, and connective and even anther wall layers having abundant druses. Their ovules are also very similar in being bitegmic and tenuinucellate, and in having a micropyle formed by the inner integument only, three cell-layered integuments, a raphe bundle terminating at chalaza, usually amphitropous or less often campylotropous ovule, embryo sac formation of Polygonum type, ephemeral antipodal cells, and tanniniferous ovule epidermis. Such ovules are readily distinguishable from those of Camellioideae and all other families. It is suggested that the three genera studied are closely related, and that the degree of embryological specialization is highest inAdinandra and lowest inEurya. On the basis of the significant embryological discrepacies, the Ternstroemioideae seem to have diverged rather distantly from the other core-subfamily Camellioideae of the Theaceae.     

Embryological observations on Tipuana tipu (Benth.) O. Kuntze (Dalbergieae,Papilionaceae)

Abstract

The anther and ovule development of Tipuana tipu is described. Some embryological aspects previously not reported for the family have been pointed out, i.e. the Basic type of anther wall development; the constant occurrence of several megasporo-cytes undergoing meiosis, the concurrent development of more than one megaspore and finally the coexistence of three or four mature gametophytes in the same ovule.     

Embryology of the Theaceae - anther and ovule development of Camellia, Franklinia, and Schima

The anther and ovule development of Camellia, Franklinia, and Schima (Theaceae, Camellioideae) were observed. The three genera share the following embryological traits: anther wall formation of basic type, tapetum of glandular type, walls of endothecial cells with secondary thickening, and production of pseudopollen grains in connective, which are dispersed into pollen sacs at anthesis, ovule bitegmic-tenuinucellate, micropyle formed by inner integument alone, hypostase present, and both integuments generally five-to-seven cell layered. One autapomorphy of the Camellioideae found in the present study is the production of pseudopollen. The three genera surveyed differ with respect to the number of middle layers in the anther, the presence or absence of stomata on connective epidermis, morphology of pseudopollen, type of embryo sac formation, form of ovule, ovular vasculature, and the proliferation of ovular epidermis, etc. Among the three genera, Franklinia and Schima are presumed to be closer embryologically, and Schima possesses more numerous specialized features.     

Embryology of Megacodon stylophorus and Veratrilla baillonii (Gentianaceae): descriptions and systematic implications

The embryological characters of Megacodon stylophorus (C. B. Clark) H. Smith and Veratrilla baillonii Franch. are described for the first time and the systematic relationships of Megacodon and Veratrilla are discussed. Both species share the following embryological features. The anthers are tetrasporangiate. The formation of anther walls is of the Dicotyledonous Type. The tapetum is of glandular type with uninucleate cells which have a dual origin. The tapetal cells on the connective side show radial elongation or periclinal division and intrude into the anther locule to form 'placentoids'. Cytokinesis in the microsporocyte meiosis is of the simultaneous type and the microspore tetrads are nearly always tetrahedral, rarely decussate. The ovary is bicarpellate and unilocular. The ovule is unitegmic and tenuinucellar. The formation of the embryo sac is of the Polygonum Type. Before fertilization, the two polar nuclei fuse into a secondary nucleus. Fertilization is porogamous. The development of the endosperm is of the Nuclear Type. However, the two species show variation in the following features: the number of cell layers which form the anther locule wall; construction of the wall of the mature anther; cell number in mature pollen grains; ovule number in cross sections of placentae; degree of ovule curvature; character of the hypostase and seed shape. In a comparison with the other taxa in the tribe Gentianeae using embryological features, Megacodon is referred to as an independent genus and should be treated as a member of the subtribe Swertiinae; Veratrilla is better separated from Swertia s . l . as an independent genus. Veratrilla is more derived than Swertia s . s. and shows a close relationship with S. tetraptera . © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 317–331.     

Embryology of Abeliophyllum (Oleaceae) and its phylogenetic relationships

Abeliophyllum, a monotypic endemic genus of Oleaceae, resembles Forsythia in various morphological characters, but its phylogenetic position is disputed and no embryological study of the genus has been carried out. We investigated more than 40 embryological characters of Abeliophyllum, compared them with previous information on Oleaceae, and discusses its phylogenetic relationships. Abeliophyllum is similar to other genera of Oleaceae in many embryological features, having some distinct features such as the mode of anther wall formation, formation of a nucellar cap, and formation of obturator and hypostase. The basic type of anther wall development and formation of a nucellar cap have not previously been reported in Oleaceae. In addition, differentiation of the obturator and formation of hypostase are not reported in the previously investigated genera of the family. Compared with close relatives, the seed coat structure of Abeliophyllum resembles Forsythia more than Fontanesia and supports existing molecular data which place Abeliophyllum as the sister group of Forsythia.     

Embryology of Gomortegaceae (Laurales): characteristics and character evolution

The embryological characteristics of Gomortegaceae, which are poorly understood, were investigated on the basis of Gomortega nitida, the only species of the family, to understand better the evolution of this group within Laurales. Comparisons with other Laurales and Magnoliales (a sister group of Laurales) show that Gomortega has many embryological features in common with the other lauralean families. Notably, Gomortega shares a testa without or with at best only a poorly developed mesotesta as a synapomorphy with all other Laurales. The genus further shares anthers dehisced by valves as a synapomorphy with the other Laurales (except for Calycanthaceae and Monimiaceae), and a non-multiplicative testa and bisporangiate anther as synapomorphies with Atherospermataceae and Siparunaceae (although the non-multiplicative testa occurs as a homoplasy in Monimiaceae, and the bisporangiate anther in Monimiaceae pro parte, Lauraceae pro parte and Hernandiaceae, respectively). Gomortega shows simultaneous cytokinesis to form pollen grains, a one-celled ovule archesporium and non-specialized chalaza, all or part of which may be synapomorphies shared with Atherospermataceae. Gomortega appears to have no embryological autapomorphies, but further comparison with Atherospermataceae is required.Kweon Heo and Yukitoshi Kimoto contributed equally to this work.     

Embryology of the dioecious Woonyoungia septentrionalis (Magnoliaceae)

The embryological characteristics and ovular integument development of the dioecious species Woonyoungia septentrionalis (Dandy) Law (Magnoliaceae), which are poorly understood, were investigated under laser scanning confocal microscope (LSCM) and light microscope (LM). The embryological characteristics conform to most of the previously studied species in Magnoliaceae. The anther has 4 microsporangia, and the anther wall develops according to the dicotyledonous type. Cytokinesis at meiosis of the microspore mother cells follows a modified simultaneous type, giving rise to isobilateral or decussate tetrads, and a cell plate is absent, but a membrane was observed. Mature pollen grains are 2‐cellular and have high germination rates. The ovule is anatropous, crassinucellate and bitegmic, and meiotic result in linear tetrads of megaspores, the one at the chalazal end functions directly as an embryo‐sac cell. The development of the embryo sac is of the Polygonum‐type and endosperm formation is of the nuclear type. The outer integument of the ovule differentiates into an outer fleshy and an inner stony layer while the inner integument is reduced to a tanniniferous layer. The normal embryological development, high germination rates of pollen and high seed set indicate that the primary reason for the decline of the species is not to be found in these developmental processes.     

Embryology and seed development of Syngonanthus caulescens (Poir.) Ruhland (Eriocaulaceae-Poales)

The embryology and the seed development of Syngonanthus caulescens are presented. This species possesses: a bithecous and tetrasporangiate anther, with a four-layered wall, a conspicuous endothecium of the baseplate type, a secretory tapetum formed by uninucleate cells, successive microsporogenesis resulting in isobilateral microspore tetrads, spiraperturate and binucleate pollen grains, an orthotropous, pendulous, bitegmic and tenuinucellate ovule, with a micropyle formed only by the inner integument, a megagametophyte of the Polygonum type, with formation of an antipodal cyst, free-nuclear and starchy endosperm, a broad and bell-shaped embryo, operculate and endotestal seeds, a seed coat derived from the inner layers of both integuments, and tanniniferous endotegmen. These embryological aspects are characteristic not only for Syngonanthus, but for the whole family, with few differences between genera. Furthermore, the pollen grain of the spiraperturate type and the cystic arrangement of the antipodals in the megagametophyte are peculiar and very distinctive features of Eriocaulaceae within the other Poales (commelinids).     

Microsporogenesis and megasporogenesis in Sinofranchetia (Lardizabalaceae)

Sinofranchetia Hemsl. (Lardizabalaceae) is a monotypic genus endemic to China. A recent combined analysis of molecular sequence data and morphology suggested that Sinofranchetia should be placed in tribe Sinofranchetieae. Embryology of taxa can be complementary to molecular phylogenetics and of special value at the genus level; however, embryological characters are completely unknown in Sinofranchetia. Here we characterize microsporogenesis and megasporogenesis in the male flowers and female flowers of Sinofranchetia, and provide embryological characters of the genus. In microspore development cytokinesis is simultaneous, the tetrads are tetrahedral and isolateral, and the mature pollens are two-celled and tricolpate (sometimes three-celled in the sterile anther of female flowers). Dicotyledonous-type and basic wall formations are found in Sinofranchetia, the tapetum is glandular. Megasporogenesis is successive, the female gametophyte of the Polygonum type. Anthers in female flowers degenerate at the tetrad stage, and some anthers produce two-celled or three-celled pollen. The newly revealed embryological characters are the basic type of anther wall formation, isolateral microspore tetrads, and both the epidermis and the parietal cell being involved in the formation of the crassinucellate ovules. The embryological peculiarities in Lardizabalaceae are discussed.     

Variations of floral traits among different life forms illustrate the evolution of pollination systems in Potamogeton species from China

Four leaf morpha groups (floating-leaved, submerged broad-leaved, submerged linear-leaved and filiform-leaved) and three inflorescences type groups (erect, facultative floating and floating) can be found in the genus Potamogeton. Life forms in the genus correlate with pollination types (anemophily, epihydrophily and hydroautogamy). Floral traits vary greatly among different life forms and pollination types. Both floating and submerged broad-leaved species produce erect inflorescences that are suited for anemophilous pollination. These flowers also have the highest pollen/ovule (P/O) ratio (25,013) and the smallest pollen grain volume (6046 μm³). On the contrary, submerged filiform-leaved species produce only floating inflorescences and their flowers produce the largest pollen grains (17,567 μm³) and have the lowest P/O ratio (6752), which is typical of the syndromes of hydroautogamy and epihydrophily. The relatively prolific seed production (about 70%) and wide dispersal in the floating and submerged broad-leaved species and the submerged filliform-leaved species indicate that these two groups are well adapted to the aquatic environment. This is in contrast with the less well-adapted complex and unstable pollination system found in linear-leaved species. We propose that the pollination systems and life forms in the Potamogeton, have undergone modification in response to the demands imposed by the aquatic environment during the evolutionary process in the genus.     

Embryology of <Emphasis Type="Italic">Petrosavia</Emphasis> (Petrosaviaceae,Petrosaviales): evidence for the distinctness of the family from other monocots

The affinities of Petrosavia, a rare, leafless, mycoheterotrophic genus composed of two species indigenous to East to Southeast Asia, have long been uncertain. However, recent molecular analyses show that the genus is sister to Japonolirion osense. Japonolirion and Petrosavia comprise the Petrosaviaceae, which are now placed in its own order, Petrosaviales, distinct from other monocots based on molecular analyses. We conducted an embryological study of Petrosavia, comparing it to Japonolirion, as well as to basal monocots (Acorus and Araceae) and more derived monocots (Nartheciaceae, Velloziaceae, and Triuridaceae). Our results showed that Petrosavia is very similar in embryology to Japonolirion, with both genera sharing a glandular anther tapetum, simultaneous cytokinesis in microspore mother cells, anatropous and crassinucellate ovules, T-shaped tetrads of megaspores, ab initio Cellular-type endosperm, and a mature seed coat composed of the exotesta, endotesta, and endotegmen. The two genera of Petrosaviaceae are clearly distinct from Acorus, and all Araceae, Nartheciaceae, Velloziaceae, and Triuridaceae genera in various combinations of characters. Thus, both molecular and embryological evidence support the distinctness of the Petrosaviaceae from other monocots and its placement in its own order, Petrosaviales.     

Embryology of Swertia (Gentianaceae) relative to taxonomy

The embryological features of three species of Swertia ( s.l.) – S. erythrosticta , S. franchetiana , and S. tetraptera – were characterized, and the observations were used, together with previously gathered data on other species, to evaluate a recently proposed polyphyly, based on molecular data, of Swertia s.l. Comparisons of species within the genus showed that they have diversified embryologically, and there are significant between-species differences. Notable features that vary between species include the number of cell layers that form the anther locule wall, the construction of the wall of the mature anther, tapetum origin, the cell number in mature pollen grains, the structure of the fused margins of the two carpels, the ovule numbers in placental cross-sections, the shape of the mature embryo sac, the degree of ovule curvature, antipodal variation and the presence of a hypostase, and seed appendages. They share characters that are widely distributed in the tribe Gentianeae, such as a dicotyledonous type of anther wall formation, a glandular tapetum with uninucleate cells, simultaneous cytokinesis following the meiosis of the microsporocytes, tetrahedral microspore tetrads, superior, bicarpellary and unilocular ovaries, unitegmic and tenuinucellar ovules, Polygonum -type megagametophytes, progamous fertilization, nuclear endosperm, and Solanad-type embryogeny. The presence of variation in embryological characters amongst the species of Swertia s.l. strongly supports the view that Swertia s.l. is not a monophyletic group. Frasera is better separated from Swertia s.l. as an independent genus, and is only distantly related to Swertia s.s. judging from the numerous differences in embryology. Swertia tetraptera is very closely related to Halenia , as they show identical embryology.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 383–400.     

Embryology of <Emphasis Type="Italic">Hemerocallis</Emphasis> L. and its systematic significance

The embryology of the genus Hemerocallis L. was studied to re-evaluate its current systematic position proposed by recent phylogenies based on molecular data. Using the improved carbol fuchsin–aniline blue staining method and conventional paraffin sectioning technique, we followed the development of anther and pollen grain, ovule and female gametophyte, and embryo and endosperm up to seed maturity. Our results showed that the (1) anther wall development is of the Monocot type, with a one cell-thick middle layer and a secretory tapetum, (2) microsporocytes cytokinesis is of the intermediate type, (3) microspore tetrads are tetragonal or decussate, (4) pollen grains are two-celled, (5) ovary is superior and trilocular, with axile placentas bearing two collateral ovules per locule, (6) ovules are anatropous, tenuinucellate, and bitegmic, with micropyle formed by the inner integument, (7) megaspore tetrads are linear, and only the chalazal one is functional, (8) embryo sac development is typically of Polygonum type, (9) embryogenesis is of Graminad type, and (10) endosperm development is of nuclear type. Overall, our study thus confirms that the embryological features of Hemerocallis support its exclusion from Liliaceae in Liliales, its inclusion in Asparagales, and its affinities with Asphodelaceae.     

Evolution of a complex coevolved trait: active pollination in a genus of fig wasps

Only three insect lineages have evolved complex active pollination behaviour and only fig wasps (Agaonidae) have also reverted from active to passive pollination. Previously, it was assumed that there was a single origin of active pollination in fig wasps, followed by one independent loss in each of five genera. We show here that there have been three to six changes in pollination behaviour within just one genus (Pleistodontes). The results suggest multiple gains of active pollination in fig wasps, but are sensitive to assumptions about the relative costs of gaining and losing this complex behaviour. In addition, previous comparative studies at higher taxonomic levels have reported correlated evolution between active pollination in wasps and low anther/ovule ratios in figs. We report that changes in pollination behaviour between congeneric species correlate perfectly with changes in anther/ovule ratios in the host figs, showing no phylogenetic inertia in coadaptation at the species level.     

Aquatic plants diversity in arid zones of Northwest China: patterns, threats and conservation

We investigated aquatic plant diversity by conducting the field investigation and collecting the published data in the arid regions of Northwest China. Two hundred and twenty four taxa of vascular aquatic plants representing 64 genera and 34 families occur in this area, 8.48% of which are endemic. Among these, 1 genus and 6 species were new state records and 1 family, 9 genera and 29 species were new area records. Typhaceae, Potamogetonaceae, Juncaginaceae and Haloragaceae were the most frequent families (considering relative frequency of occurrence), whereas Cyperaceae, Potamogetonaceae and Ranuncnlaceae are the most species-rich. The most frequent genera were Typha, Potamogeton, and Triglochin, and the most species-rich were Potamogeton, Eleocharis and Scirpus. The most frequent species are Triglochin palustre, Myriophyllum spicatum, Potamogeton pectinatus and Typha angustifolia. Aquatic plants diversity is distributed unevenly in the region. The maximum species occurs in Dzungarian basin while the least species in Hexi corridor. The aquatic flora in arid zone of China is not distinctive although some endemic species are found, most species are widely distributed. Local aquatic plants diversity can be influenced by many factors such as hydrological alteration, habitat loss, over-grazing, high human population pressure, global climate change, an inappropriate economic development policy. Among them, the largest threat to aquatic plants biodiversity may be habitat loss due to hydrological alteration. In order to conserve the aquatic plants biological resources and biodiversity in this region, some strategies and measures must be suggested including strengthening scientific research and biodiversity education in the local people, balancing economic development and ecological conservation, and enhancing governmental assistance and subsidy to the local residents.     

The embryology of <Emphasis Type="Italic">Xanthoceras</Emphasis> and its phylogenetic implications

Recent molecular analyses found the Chinese monotypic genus Xanthoceras to be either a sister genus to the rest of a broadly defined Sapindaceae sensu lato (s.l.) including Aceraceae and Hippocastanaceae or the only genus of the Xanthoceraceae. Here we investigate for the first time the embryology of Xanthoceras and compare the embryological data with related taxa to aid in understanding the phylogenetic relationships of this genus. The results indicate that Xanthoceras shares many embryological features with Sapindaceae sensu stricto (s.s.), Aceraceae, and Hippocastanaceae, including a glandular tapetum, simultaneous microsporogenesis, three-colporate pollen grains, a bitegmic and crassinucellate ovule, Polygonum type of embryo sac development, endosperm formation of the nuclear type, and exalbuminous mature seeds. The wide range of similarities in embryological features suggests that Xanthoceras is closely related to other members of Sapindaceae s.l. However, Xanthoceras differs from other Sapindaceae in several morphological characteristics, for instance, five to eight ovules in each locule in Xanthoceras versus one to two ovules in each locule in other Sapindaceae s.l., and five golden horn-like appendages present on both staminate and hermaphrodite flowers of Xanthoceras versus absent on the flowers of other Sapindaceae s.l. We propose that Xanthoceras merits recognition at the subfamily level as Xanthoceroideae, rather than at the family level as Xanthoceraceae. The present study also reveals that late-acting ovarian self-incompatibility (OSI) occurs in X. sorbifolium and that postzygotic OSI contributes to the self-sterility of the species.