Embryology of Swertia tetraptera Maxim. (Gentianaceae) and its systematic implication

The embryology of Swertia tetraptera is reported for the first time. The anther is tetrasporangiate and its wall, of which the development conforms to the dicotyledonous type, is composed of epidermis, endothecium, middle layer and tapetum at the mature stage. The tapetum has dual origin and is similar to the glandular type. Cytokinesis is of the simultaneous type and microspore tetrads are mostly tetrahedral. Pollen grains are mostly 3-celled when shed. The ovary is bicarpellate and unilocular with four series of ovules. The fused margins of two carpels intensively protrude into the locule. The ovule is unitegmic, tenuinucellate and orthotropous. The development of embryo sac is of the Polygonum type. The development of endosperm conforms to the nuclear type and the embryogeny corresponds with the Physalis I variation of Solanad type. Three antipodal cells persist at the mature embryo sac. The embryo is at the globular stage when seeds are released from the capsule. Compared with the other studied species in Swertia, S. tetraptera shows differences from them in three distinctive embryological traits: enlargement and protrusion of the fused margins of two carpels into the ovary locule; orthotropous ovules and a developed hypostase. These three characters were previously only found in the genus Halenia of the tribe Gentianinae in the Gentianaceae,and thus suggest that S. tetraptera should be related to Halenia. The results partly correspond with the molecular evidence (Yuan & Kupfer, 1995). The monophyletic Halenia clade in Yuan & Kupfer's molecular cladogram includes Halenia, S. tetraptera and Frasera (section Frasera in Swertia s. l.); nevertheless, up to now, no embryological study on Frasera has been reported. The embryological results imply that the taxonomic status of S. tetraptera needs to be revaluated and the genus Swertia s. l. may not be a mono-phyletic group. Decontaminated thianthrene disproportion. 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四数獐牙菜的胚胎学及其系统学意义

四数獐牙菜花药四室,药壁发育为双子叶型;绒毡层腺质型。小孢子母细胞减数分裂为同时型, 四分体的排列方式为四面体形;成熟花粉为3-细胞。子房2心皮,2心皮连接处强烈内凸,4列胚珠。薄 珠心,单珠被,直生胚珠。胚囊发育为蓼型。胚乳发育为核型。胚胎发育为茄型酸浆I变型。反足细胞 在胚囊成熟时期宿存。果实成熟时,种子只发育至球型胚阶段。承珠盘存在。与已研究的獐牙菜种类 的胚胎特征相比较,该种存在三个显著的特点：直生胚珠,心皮连接处强烈内凸,承珠盘存在。这三个特 征目前仅在龙胆族中的花锚属中发现。经过性状分析,它们均是演征,此三个共有演征表明四数獐牙菜 与花锚属是一单系群,它们的亲缘关系最近。这与Yun &; Kupfer(1995)的分子证据部分吻合,因为分 子系统树上的花锚单系支包括花锚、四数獐牙菜和北美的密花属(广义獐牙菜属中的密花组),而后者的 胚胎学特征未有任何报道。胚胎学研究暗示四数獐牙菜的分类学等级有待重新评价,广义的獐牙菜属 有可能不是一单系发生群。     

Embryology of Zippelia begoniaefolia (Piperaceae) and its systematic relationships

Microsporogenesis and embryology of the monotypic Zippelia (Z. begoniaefolia) Blume (Piperaceae) is described for the first time to assess its systematic relationships. The formation of the anther wall is of Basic Type such that the anther wall, consisting of an endothecium with fibrous thickenings, two middle layers, and a glandular septum with 2‐nucleate cells, is derived from a primary parietal layer. Simultaneous cytokinesis follows meiosis of the microspore mother cell thence forming a tetrahedral tetrad of microspores. The single basal ovule is orthotropous, crassinucellate and bitegmic but only the inner integument forms the micropyle. The sporogenous cell of the nucellus functions directly as a megaspore mother cell. A coenocyte with four nuclei forms after meiosis of the megaspore mother cell. The formation of the embryo sac is tetrasporic ab initio and is of, or similar to, the Drusa Type of embryo sac in which the nuclei of the coenocyte undergo two successive mitoses and forms a 16‐celled or 16‐nucleate embryo sac that is ovoid in shape. The embryo sac has an egg apparatus consisting of an egg cell and two synergids (but one of the latter is less discernable). Two polar cells occur just beneath the egg apparatus and 11 antipodal cells or nuclei are arranged along the lower part of the inner wall of the embryo sac. They are linked by threads of cytoplasm. The two polar cells are separated or fused before fertilization. A large primary endosperm nucleus with many nucleoli, which resulted from the fertilized polar cells and with the participation of antipodal cells, divides into a free nuclei stage. The free nuclei are arranged along the lower part of the inner wall of the embryo sac or rarely assemble at the central part. The development of endosperm is thus of the Nuclear Type. The zygote remains undivided and fails to develop even when the seed is nearly mature. Frequently, the zygote and the endosperm abort later and leave an empty chamber in the top part of the seed. Most of the seed content is starchy perisperm. Only the inner integument forms the seed coat and the pericarp develops glochidiate hairs (anchor‐like hairs) when the endosperm begins to develop. By comparison with the other piperaceous taxa using embryological and botanical features, Zippelia is referred to as a basal taxon and a more isolated evolutionary line or a blind branch in the Piperaceae. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society, 2002, 140 , 49–64.     

Embryology ofCrocus thomasii (Iridaceae)

The embryology ofCrocus thomasii is described. Male meiosis is of simultaneous type, and gives rise to starchy microspores which develop into lipoid pollen grains; these are two-celled and show a spinulate acolpate, abaculate exine lacking apertures. The tapetum is glandular and its cells become bi- or sometimes multinucleate. The ovule is anatropous and bitegmic; the inner integument forms the micropyle. Megasporogenesis is heteropolar with starch accumulation in the functional chalazal megaspore. Embryo sac development conforms to thePolygonum type. The endosperm development is nuclear. The embryo develops according to the Caryophyllad type. In the ripe seed it is differentiated and enveloped by a starchy cellular endosperm. The embryological characters observed strongly favour a close relation betweenC. thomasii andC. sativus.     

Floral development of Iris decora Wall. (Iridaceae)

The outer tepal and stamen primordia arise as secondary primordia on the outer tepal-stamenprimordia, which are formed on the floral apex. The inner tepal primordia are formed directly on the floral apex. All the floral appendages are initiated in the second tunica layer and are homologous with regard to their origin and early development. A short perianth tube is formed as a result of intercalary growth in the common bases of the tepal primordia. The intercalary growth in the fused bases of the floral appendages elevates the peripheral zone. The floral apex thus appears as a shallow cup. Further intercalary growth results in the formation of an inferior ovary. The ovules are initiated as outgrowths on placental ridges from the lateral ovary wall, the trilocular appearance being the result of secondary cohesion of the parietal placentae.     

玉蝉花繁殖生态学研究

通过野外调查、人工授粉和室内种子萌发实验等,对天目山玉蝉花（Iris ensata）种群繁殖生态特征进行了研究。结果显示：玉蝉花自然种群呈聚集型分布,种群更新主要依赖贡献率为0.756的无性繁殖。人工同瓣和异瓣自花授粉座果率分别为10%和20%,结实数分别为0和4±1（n=2）粒/果;同株和异株异花授粉座果率分别为80%和100%,结实数分别为59±7（n=8）和64±9（n=10）粒/果,自交主要来自同株异花授粉,而同株花异熟、雌雄异位和异向等可有效降低自交。自然套袋不结实,不存在无融合生殖。花粉/胚珠比1277±270（n=10）,辅证玉蝉花为兼性异花授粉、异花授粉植物。有效授粉昆虫为蜜蜂（Apismellifica）,访花频率为0.019次.花-1.h-1,单只蜜蜂回访同朵花的比率为3.5%。自然种群立地种子数估算值为368粒/m2,种子宿存,具非深度生理休眠,萌发需光,低温层积60d萌发率达94.9±0.7%。玉蝉花种群繁殖适应对策为k对策,具备自我维持机制,但现正受人为干扰。     

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 ofKyllinga monocephala (Cyperaceae) and its systematic position

Aspects of the life history ofKyllinga monocephala are described. Anther wall development corresponds to the Monocot type. The endothecium shows spiral thickenings. The tapetum is glandular and has uninucleate cells. Ubisch granules are present. Mature pollen grains (pseudomonads) are 3-celled at maturity. Ovules are bitegmic, crassinucellate and develop a funicular obturator. The embryo development conforms to theJuncus-variation of the Onagrad type. Endosperm, seed coat and pericarp are described.     

Author and type of the name Iris cengialti (Iridaceae)

Abstract

A. Kerner is correctly the author of the name Iris cengialti previously ascribed to Ambrosi, and consequently the type of the species is here selected, as lectotype, among Kerner's original material kept in WU.     

Leaf anatomy and relationships of Dietes (Iridaceae)

The leaf anatomy of Dietes is described, including all 6 species. Characters support the morphological evidence that Dietes is a distinct genus and related to other Iridoideae. Certain features, notably the epidermal structure and sheath vasculature, distinguish it from other Iridaceous genera, although the sheath vasculature indicates a relationship with some New World Iridoideae. The leaf margin type is similar to that of other Old World Iridoideae. Many leaf anatomical characters in Dietes are associated with xeromorphy. Leaf anatomy supports the division of Dietes into 2 subgenera.     

Embryology and relationships ofSuriana maritima L. (Surianaceae)

An intensive study of the embryology ofSuriana maritima L. is made to provide the information for discussing relationships and systematic position of the genus which is uncertain yet. Notable embryological features ofSuriana include: ovular archesporium multi-celled; ovule unitegmic, campylotropous, and crassinucellate with a nucellar cap; obturator not formed; ovule and seed pachychalazal; chalazal haustorium of endosperm formed; mature seed exalbuminous. Comparisons with possibly related families suggest thatSuriana is distinct from any of the families discussed but best resembles Rutaceae (and Simaroubaceae). Until a study onStylobasium and several other genera, which might be closer toSuriana, comes up in future, we tentatively support the placement ofSuriana as a monotypic family Surianaceae in Rutalessens. Takhtajan.     

Embryology of Plagiopteron suaveolens Griffith (Plagiopteraceae) and its systematic implications

A study of the embryology of Plagiopteron suaveolens Griffith is presented in order to evaluate the affinities of the genus. Anther wall formation is of the dicotyledonous type. The anther tapetum is glandular and its cells become two to four nucleate. Cytokinesis in the microspore mother cell is simultaneous and the resultant tetrad is tetrahedral. The mature pollen grains are two-celled. The anther has 4-locules at anthesis. The ovule is anatropous, bitegmic and weakly crassinucellate with only one cell-layered parietal tissue. The micropyle is zig-zag in section and formed by both inner and outer integuments which are three and three to four cell layers thick respectively. The megaspore archesporium is one-celled and the embryo sac development is of the Polygonum type. An endothelium is formed with early disintegration of nucellar tissue. The antipodals multiply to 15 to 20 cells. Embryological evidence supports the placement of Plagiopteron in its own family, Plagiopteraceae, with closest relationship with Celastraceae and close relationship with Elaeocarpaceae, Rhizophoraceae and some families of the Geraniales.     

Isolation and characterization of polymorphic microsatellites in Iris ensata (Iridaceae)

• Premise of the study: Microsatellite primers were developed in Iris ensata (Iridaceae) to provide polymorphic markers for further studies into population genetics. • Methods and Results: Thirteen polymorphic microsatellite loci were isolated from I. ensata. These loci were successfully amplified in two natural populations of I. ensata from eastern China (Longwangshan, Zhejiang Province) and northeastern China (Jinchuan, Jilin Province). There was no significant linkage disequilibrium found for any pair of loci. These loci contained between two and 12 alleles per locus across all 48 individuals of I. ensata. The number of alleles per locus varied from two to 10 at the population level and the observed and expected heterozygosities ranged from 0.167 to 0.958 and from 0.284 to 0.853, respectively. • Conclusions: These loci showed high levels of polymorphism and could be used to study the population genetic structure, genetic relationships, and phylogeography of I. ensata.     

Embryology and relationships of Akania (Akaniaceae)

We present the first extensive study of embryology of Akania , the only genus of Akaniaceae (one of the 15 glucosinolate-producing families). Akania has a distinctive combination of embryological features, which includes bitegmic and crassinucellate ovules; thick, multiplicative and vascularized outer integument; a Polygonum-type embryo sac; albuminous seed; 'exotestal' seed coat with a palisade of columellar, thick-walled cells and a thick and sclerotic mesotesta with the inner tissue aerenchymatous. A comparison of its embryological characteristics with those of some other groups indicates that Akania closely resembles Bretschneideraceae (another glucosinolate-producing family) and Sapindaceae, but it retains some archaic embryological features and is best treated as a separate family in or near Sapindales, as in most modern general classifications. Based on evidence from embryological and other data, Akaniaceae appear to be the sister group of Bretschneideraceae, possibly near to Sapindales. With the data at hand, we cannot fully assess the recent suggestion that Akaniaceae-Bretschneideraceae are a basal group in Capparales.     

Embryology and relationships of Lauraceae (Laurales)

Embryology of Lauraceae, hitherto poorly known, was investigated on the basis of 35 species from 23 genera to contribute to a better understanding of familial and generic relationships. Data from the genera investigated and from the literature show that the genera of Lauraceae are very similar embryologically, but that differences do exist in a few characters among the genera. Based on comparisons with other families of Laurales, Lauraceae consistendy had a pachychalazal ovule or seed with a ramified raphal vascular bundle at chalaza (an apomorphy) in common with Hernandiaceae. However, since several core lauralean families such as Amborellaceae, Monimiaceae, and Gomortegaceae are little known embryologically, these must be studied for critical comparison. Wimin Lauraceae, Cassytha is clearly distinct from the rest of the family in having an ab initio cellular type endosperm (a plesiomorphy, also reported in Umbellularia ) instead of a nuclear type endosperm (an apomorphy) as in the rest of the family, in lacking the nucellar cap and in having the micropyle formed by bom the inner and outer integument, facts supporting the traditional taxonomic placement of Cassytha in its own subfamily and the remainder of the family in the other subfamily. Widiin the rest of the family, the amoeboid tapetum (an apomorphy) distinguishes 15 genera ( Actinodaphne, Cinnamomum, Laurus , etc.) from the six genera with the glandular tapetum. In addition, a mature embryo sac protruding from the nucellus (an apomorphy) distinguishes five genera ( Beibchmiedia, Caryodaphnopsis, Cryptocarya, Endiandra, Potamria) and one species of Ocotea (O. rubra ) from the rest of the family. These results can properly be incorporated in a future suprageneric classification. The distinctness of Ocotea rubra wimin the genus is also discussed.     

Embryology and relationships ofGyrocarpus andHernandia (Hernandiaceae)

Two representative genera of Hernandiaceae,Gyrocarpus andHernandia, were investigated embryologically to contribute to a better understanding of their respective evolutionary position. Comparisons with other lauralean families using Chloranthaceae or Annonaceae (as a representative of Magnoliales) as an outgroup of Laurales (if present, plus other related taxa) support a lauraceous affinity for the two genera because of the presence of ramified raphal vascular bundles at the chalaza (a synapomorphy), but do not provide evidence for the separation of Hernandiaceae from Lauraceae.Hernandia rather shares with Lauraceae two apomorphies (i.e., the seed pachychalazy and the ruminate seed) which may be homoplasies judged from results of cladistic and molecular studies published elsewhere.Hernandia is greatly divergent from an ancestral line common withGyrocarpus and is even diversified within the genus. Based on evidence from embryology as well as from other sources, it seems best to accept two separate subfamilies in Hernandiaceae as usually have been accepted: one is a derived subfamily Hernandioideae, and the other a less specialized Gyrocarpoideae.     

Isolation and characterization of 10 microsatellite loci from Iris hexagona (Iridaceae)

Ten microsatellite loci isolated from Iris hexagona are described. All these loci are polymorphic, with three to 13 alleles across 24 individuals from a single natural population. Heterozygosity ranged from 0.125 to 0.870. Three loci depart significantly from Hardy–Weinberg equilibrium in our test population. The test population shows significant heterozygote deficiency in these and two other loci. Three loci exhibit significant linkage disequilibrium. These loci will be utilized to investigate patterns of genetic variation in the species throughout the Florida peninsula.     

Phylogeography of Iris missouriensis (Iridaceae) based on nuclear and chloroplast markers

We investigated the phylogeography of Iris missouriensis (Iridaceae), which is widely distributed in western North America. We utilized transposon display and DNA sequencing to quantify nuclear and chloroplast genetic structure. Our objectives were (i) to characterize the geographic structure of genetic variation throughout the species range, (ii) to test whether both margins of the range show reduced genetic diversity as predicted by north-south expansion and contraction associated with climate change, and (iii) to determine whether the subspecies Iris missouriensis ssp. longipetala is genetically distinct. We found that genetic diversity was significantly lower in the northern part of the range but was not significantly different between the central and southern regions, indicating greater stability of the southern margin vs. the northern. Among-population differentiation was high (PhiPT=0.52). The largest divisions in each marker set were concordant and separated the southern Rocky Mountains and Basin and Range provinces from the remainder of the range. The boundaries of this phylogeographic break do not coincide with gaps in present-day distributions or phylogeographic breaks identified in other species, and may indicate a measure of reproductive isolation. Consistent with current treatments, we did not find support for the taxonomic placement of I. missourienis ssp. longipetala as a distinct species. Although transposon display has been used to investigate relationships among crop accessions and their wild relatives, to our knowledge, this is the first use of these markers for population-level phylogeography of a nonmodel species and further demonstrates their utility in species recalcitrant to amplified fragment length polymorphism protocols.     

Embryology and probable relationships of Eriospermum (Eriospermaceae)

Three South African species of the African genus Eriospermum Jacq ., which makes up Eriospermaceae Endl., have been studied in connection with the project "Families of Vascular Plants", partly to establish the phylogenetic relationships of the family. The embryology is unusual in several features reflecting the advanced character of the genus in this respect. The following features are the most important: the tapetum is secretory; microsporogenesis is successive; the ovules are anatropous and crassinu-cellate; the primary archesporial cell cuts off a parietal cell; the embryo sac formation is of the Polygonum type; endosperm formation is nuclear, but the endosperm is soon consumed, nucellus cells bordering on the sides of the chalazal half of the embryo sac divide to form a perispermal sheath around the embryo; embryo formation follows the Nicotiana variation of the Solanad type; the embryo of the mature seed is large and cylindrical-obconical, reaches above the perisperm. The seed coat is formed by both the outer and the inner integuments, both 2-layered; the epidermal cells of the test a grow out into long trichomes. The embryology and seed shape of Eriospermum is compared to those in Cyanastrum, Walleria , and genera of Tecophilaeaceae and some other families as far as details in these are known.     

Embryology and systematic position ofLiparis (Orchidaceae)

The embryology ofLiparis paradoxa andL. rostrata has been studied. The young anther wall consists of an epidermis, endothecium, three middle layers and secretory tapetum with uninucleate cells. In the mature anther, two or three sub-epidermal layers develop fibrous thickenings. The anther wall development corresponds to the massive type. Simultaneous cytokinesis results in decussate, isobilateral, linear, T-shaped and tetrahedral pollen tetrads. The pollinia are compact and at anthesis the pollen grains are 2-celled. Ovules are anatropous, bitegmic and tenuinucellate. Both the integuments are dermal in origin and 2-layered. Development of female gametophyte is of the monosporic, 8-nucleate type. Double fertilization occurs. The primary endosperm nucleus degenerates. The mature embryo is organized from the derivatives of tiers 1, 1, m, and n. Its development conforms to a variation of the Onagrad type. The seed coat is formed entirely by the outer layer of the outer integument. There are three sterile and three fertile valves in the ovary. In the prefertilization stages valves consist of parenchymatous cells. After fertilization, the sterile valves become sclerenchymatous whereas the fertile valves remain parenchymatous. The embryological characters support the disputed systematic position ofLiparis within subtribeLiparidinae ofEpidendreae.