Kingdonia,Embryology and Its Systematic Significance

The embryological studies show that Kingdonia is similar to Circaeaster while different from other members of the Ranunculales (sensu Takhtajan, 1980) in development of endosperm and embryo. We consider that there is close systematic relationship between Kingdonia and Circaeaster.     

独叶草属的胚胎学及其系统学意义

独叶草属的胚乳发育为五福花型,其早期分化属细胞型;胚胎发育为石竹型下的独叶草亚型;珠 被内层的部分细胞形成栅栏组织状,最后消失或近于消失;反足细胞宿存,部分不育大孢子宿存并和部 分珠心组织一起形成珠孔塞状结构。这些性状表明它与星叶草属间的相似性要大于与毛茛目其它成员 间的相似性。     

嵩草属植物的胚胎学

对嵩草属（ Ｋｏｂｒｅｓｉａ） 植物进行了初步的胚胎学研究。该属植物具假四合花粉（ｐｓｅｕｄｏｍｏｎａｄ） ; 药室内壁在二核花粉时期开始螺旋状加厚, 花药表皮在花粉成熟时形成乳突; 成熟花粉具三细胞。胚珠为倒生型, 具厚珠心和双层珠被, 珠孔由内珠被构成, 珠柄的近基部向珠孔增生形成珠孔塞。胚囊的发育为蓼型, 四分体线形排列, 合点端大孢子发育成八核胚囊。受精后, 胚乳核先于受精卵进行分裂, 胚乳的发育为核型。胚的发育为柳叶菜型灯芯草变型。通过比较, 嵩草属植物大小孢子的发育、胚珠的结构、胚囊的发育、胚乳的发育和胚的发育与莎草科中其它类群一致。所以, 根据胚胎学资料, 嵩草属及其近缘属应保留在莎草科中,不该另立为嵩草科。     

The Embryology of the Genus Kobresia(Cyperaceae)

对嵩草属(Kobresia)植物进行了初步的胚胎学研究。该属植物具假四合花粉(pseudomonad);药室内壁在二核花粉时期开始螺旋状加厚,花药表皮在花粉成熟时形成乳突;成熟花粉具三细胞。胚珠为倒生型,具厚珠心和双层珠被,珠孔由内珠被构成,珠柄的近基部向珠孔增生形成珠孔塞。胚囊的发育为蓼型,四分体线形排列,合点端大孢子发育成八核胚囊。受精后,胚乳核先于受精卵进行分裂,胚乳的发育为核型。胚的发育为柳叶菜型灯芯草变型。通过比较,嵩草属植物大小孢子的发育、胚珠的结构、胚囊的发育、胚乳的发育和胚的发育与莎草科中其它类群一致。所以,根据胚胎学资料,嵩草属及其近缘属应保留在莎草科中,不该另立为嵩草科。     

喉毛花的胚胎学研究

本文首次系统地记载了喉毛花属的胚胎发育过程,并以此为依据讨论了该属的分类等级和系统位置。喉毛花花药四室;药壁发育属双子叶型;绒毡层单型起源,细胞具单核,属腺质绒毡层;一层中层细胞;花药壁表皮层宿存,纤维状加厚和膨大;药室内壁减缩。小孢子母细胞减数分裂为同时型,四分体的排列为四面体型;成熟花粉为3-细胞型。子房为2心皮、l室,典型的侧膜胎座,胚珠8列,胚珠胎座靠近两心皮腹缝线。薄珠心,单珠被,倒生胚珠。大孢子母细胞减数分裂形成的4个大孢子呈直列式排列,其中合点端的大孢子具功能。胚囊发育为蓼型。极核在受精前融合为次生核。反足细胞宿存、分裂为8～12个,每个细胞均多核和异常膨大,反足细胞形成的吸器明显。异花传粉,珠孔受精。花粉管通过破坏一助细胞进入胚囊。受精作用属于有丝分裂前配子体融合类型。胚乳发育为核型,每核含2～3核仁。胚胎发育为茄型酸浆I变型,成熟种子胚只发育至球形胚阶段。反足细胞在合子分裂之后才开始退化,在胚的发育过程中反足细胞在胚乳层之外形成一层染色深、类似“外胚乳”的结构。比较喉毛花、龙胆属、假龙胆属以及肋柱花属的胚胎学特征表明喉毛花与假龙胆属的亲缘关系最近,在分类等级上作为一个独立的属较为合适,在系统位置上它比假龙胆属更为原始。     

峨眉双蝴蝶的胚胎学研究

首次报道了峨眉双蝴蝶Ｔｒｉｐｔｅｒｏｓｐｅｒｍｕｍ ｃｏｒｄａｔｕｍ（Ｍａｒｑ．）Ｈ．Ｓｍｉｔｈ的胚胎学特征,研究结果用以讨论双蝴蝶属的系统演化关系。主要研究结果如下：花药四室;药壁发育为双子叶型;绒毡层属单型起源,细胞具单核,属腺质型绒毡层,药隔处的绒毡层细胞形成类胎座,其余部位的绒毡层细胞仍为一层细胞;在花药成熟时,花药的药室内壁纤维状如厚且柱状伸长,表皮细胞减缩退化,纤维状加厚不明显。成熟花     

Embryological Studies on Sagittaria guayanensis H. B. K. Subsp. lappula (D. Don) Bojin (Alismataceae)

This paper deals with the embryological characteristics of Sagittaria guayanensis H. B.K. subsp. lappula (D. Don) Bojin. The anther wall development follows the Monocotyledonous type. The cytokinesis of microspore mother cell in meiosis is of the Successive type. The tetrads of microspores show an isobilateral arrangement, and the mature pollen grains are 3-celled. The ovule is bitegminous, pseudo-crassinucellate and anatropous. The megaspore mother cell originates directly from a single archesporial cell. The mature embryo sac consists of 7 cells including 8 nuclei and conforms to the Allium type. The two polar nuclei do not fuse into a secondary nucleus before fertilization. Instead, one sperm fuses with the micropylar end polar nucleus first , and the fertilized polar nucleus then migrates to the chalazal end, where it fuses with the second polar nucleus, forming the primary endosperm nucleus. The embryo development conforms to the Caryophyllad type. The mature embryo is U-shaped and forms the embryonic shoot apex accompanied by two leaves. The endosperm development corresponds to the Helobial type. The primary endosperm nucleus (invariably lying in the chalazal part of the embryo sac) divides and forms two chambers:large micropylar one and small chalazal one. The chalazal endosperm chamber remains binucleate, while, in the micropylar chamber free nuclear divisions occur and then cellnlarization takes place. During the embryo formation the endosperm gradually degrades and can not be found in the mature seed. The subgenus Lophotocarpus is different from the subgenus Sagittaria in some embryological aspects, especially in the structure of mature embryo sac and the double fertilization process.     

冠果草的胚胎学研究

冠果草花药壁的发育为单子口十型,绒毡层为周原质团型。小孢子母细胞减数分裂为连续型,四分体呈左右对称式排列,成熟花粉为三细胞型。双珠被,假厚珠心,倒生胚珠。胚囊发育为葱型,成熟胚囊的特点是两个极核分别位于中央细胞两端,不融合成次生核。受精过程中,一个精于与卵核融合形成合子,另一精子先与珠孔端极核融合,之后受精极核再移动到合点端与另一极核融合,形成初生胚乳核。胚的发育为石竹型。成熟胚呈马蹄形,具有2片真叶。胚乳发育为沼生目型。随着胚的发育,胚乳细胞逐渐解体,成熟种子中无胚乳。     

Early reproductive developmental anatomy in Decaisnea (Lardizabalaceae) and its systematic implications

Background and Aims

Decaisnea insignis, known as ‘dead man''s fingers’ (Lardizabalaceae), is widely distributed in China and the Himalayan foothill countries. This economically important plant, which is the only species in the genus, has not been the subject of any embryological studies aside from one brief, older paper that lacks micrographs. Data on Decaisnea are also important because its systematic position has been unstable since the genus was established in 1855. Therefore, the objectives of this study were: (a) to use modern microscopy to document early reproductive anatomical development in Decaisnea; and (b) to compare qualitatively these early embryological characters with allied taxa in a systematic context.

Methods

Decaisnea insignis floral buds and inflorescences were regularly collected from Shaanxi Province, China and prepared for light microscopy. The embryological characters studied were qualitatively compared with those of allied taxa via a thorough examination of the existing literature.

Key Results

Early reproductive anatomy in Decaisnea was documented and novel revelations made. It was discovered that the pollen is shed when three-celled (not two-celled, as previously reported), and that endosperm formation is nuclear (not cellular or helobial, as previously reported). These two newly revealed embryological characters are not found in any other members of Lardizabalaceae. Furthermore, neither are persistent antipodal cells, which we confirmed to be present in Decaisnea.

Conclusions

Decaisnea and other Lardizabalaceae characteristically have tetrasporangiate anthers, a secretory tapetum, simultaneous microsporocyte cytokinesis, primarily bitegmic, crassinucellate ovules, and a Polygonum type embryo sac. However, in the family, persistent antipodals, nuclear endosperm, and pollen shed at the three-celled stage are only found in Decaisnea. These embryological data prompted the suggestion that Decaisnea needs elevation above the level of genus.     

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.     

藏药椭圆叶花锚的胚胎学

本文首次报道了藏药椭圆叶花锚的胚胎学特征。花药四室,药壁发育为双子叶型;绒毡层异型起源,腺质型绒毡层。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型;成熟花粉为３－细胞。子房２心皮,而二心皮连接处强烈膨大、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。承珠盘发达。果实成熟时,种子只发育至球型胚阶段。比较了该种与龙胆族其它属、种的胚胎学特征,发现它们大部分特征是相似的,但在如下３个特征上存在区别：子房二心皮连接处强烈膨大、内凸;直生胚珠;具有发达的承珠盘。其胚胎学特征的系统学和分类学意义有待进一步比较与评价     

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.     

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.     

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.     

石香薷（唇形科）的胚胎学研究

石香薷(Mosla chinensis Buch.-Ham.ex Maxim.)花药壁发育属双子叶型。花药具4个小孢子囊;腺质绒毡层,细胞具2～4核,有3至数个核仁;初生造孢细胞直接行小孢子母细胞的功能,在小孢子囊中成单列。花粉母细胞减数分裂后胞质分裂为同时型;小孢子四分体呈四面体形,也有左右对称形,成熟花粉具2细胞。胚珠倒生,单珠被,薄珠心,大孢子四分体线形排列,功能性大孢子位于合点端,少数为合点端第二个细胞。胚囊发育属蓼型,珠孔区近卵圆形,比合点区稍短,合点区较狭窄。胚胎发生属柳叶菜型。细胞型胚乳,珠孔吸器为单孢3核,合点吸器为单孢2核。种子无胚乳,种皮由珠被发育。石香薷雌雄配子体的发育、胚胎发生及胚乳形成,与紫苏属的Perilla ocimoides几乎完全一致。不同点仅在于石香薷在2-细胞花粉时,药室内壁细胞切向伸长,壁尚未发生纤维状加厚(P.ocimoides药室内壁细胞径向伸长,胞壁纤维状加厚),珠孔吸器为单孢3核(P.ocimoides为单孢4核)。胚胎学显示石荠苎属与紫苏属有密切的亲缘关系。     

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.     

Development of ovule,embryo sac,and endosperm in Dipterostemon and Dichelostemma (Alliaceae) relative to taxonomy

Dipterostemon capitatus and all species of Dichelostemma have the following characters in common: the ovule is anatropous and bitegmic; the nucellar epidermis is penetrated by the embryo sac; the remaining chalazal part of the nucellus expands, partly due to periclinal divisions in its epidermis; the micropyle is formed by the inner integument only; parietal cells are present; the embryo sac develops according to the Polygonum type; the endosperm is nuclear; after fertilization, the cells of the inner integument enlarge greatly. One difference between the genera is that the inner integument is two cells thick in Dipterostemon, but five to seven cells thick in all Dichelostemma species. In Dipterostemon, embryogenesis is of the Asterad type, and the formation of walls in the endosperm occurs much as in wheat. The absence of variation in embryological characters among the species of Dichelostemma strongly supports the view that these species all belong in one genus, despite differences in gross morphology. Dichelostemma is very closely related to Brodiaea, since species of these two genera show identical embryology. Dipterostemon must be retained as a genus because a deviating inner integument adds to uniqueness in gross morphology. Dipterostemon, Dichelostemma, and Brodiaea, are embryologically quite different from Triteleia. Neither Dipterostemon nor Dichelostemma show any close affinity with Allium.     

Self-sterility in two Cytisus species (Leguminosae, Papilionoideae) due to early-acting inbreeding depression

In most angiosperms, the endosperm develops before the embryo, but with harmony between the two structures until final seed formation. In an embryological study, we show that inbreeding depression causes disharmony in development of the two structures in two Leguminosae shrubs, Cytisus multiflorus and C. striatus. Our main objective was to test the causes of self-sterility in the two species by comparing the embryological development of the self seeds with that of cross seeds. In developing selfed seeds of C. multiflorus, the embryo reaches at most the globular stage and never forms mature seeds, while in C. striatus a few mature selfed seeds are formed. In both species, the main cause of abortion of developing selfed seeds is diminished endosperm development (low values of the ratio of endosperm to embryo), which triggers collapse of the endosperm and embryo. The results indicate that self-sterility in C. striatus is postzygotic because of strong, early inbreeding depression, while in C. multiflorus there exists a mixed pre- and postzygotic mechanism; the prezygotic mechanism causes rejection of some self-pollen tubes in the style/ovary, and the early inbreeding depression triggers abortion of fertilized ovules that escaped that action.     

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.