Morphological Studies on Circaeaster agrestis Maxim. (1) Process of Embryolo gical Development

The present paper aims at discovering the characters of embryological development of Circaeaster agrestis, which makes up a monotypic genus, Circaeaster, to establish the phylogenetic relationships of the genus. The different opinions on its systematic position among botanists are briefly explained. The embryological studies show that the most important advanced characters of the genus are as follows. The ovule is amphitropous, unitegmic and tenuinucelar; the embryo sac formation is in accordance with the Polygonum type; endosperm formation is of the cellular type, the primary endosperm nucleus dividing to form two cells and the first wall vertical; embryo formation follows the variation of the Caryophyllad type; at the early stage of development of embryo, the integument has been already atrophied and at last disappeared, so that the seed coat is absent in the mature fruit. On the basis of the embryological and some morphological evidence, the authors consider that a close relationship between the genus and Ranunculaceae and its related families seems to be unlikely. The affinities of the genus Circaeaster are still uncertain.     

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。     

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。     

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。     

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.     

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。     

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.     

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.     

白毛新木姜子[Neolitsea aurata var. glauca]胚胎学研究

应用光学显微镜对白毛新木姜子的胚胎学特征进行了研究,首次在非寄生性樟科植物中发现了细胞型胚乳。对樟科8属进行了胚胎学特征的比较。花药四室,药室壁的发育属于“基本型”,周原质团型绒毡层。小孢子母细胞连续型分裂。等四面体型四分体。二细胞成熟花粉,无孔沟。雌孢原多个,一般仅一个能继续发育。蓼型胚囊。助细胞具丝状器。反足细胞宿存。大孢子母细胞和合子具极性。宿存的一个助细胞具有吸器功能。细胞型胚乳,胚胎发育属于柳叶菜型的三叶变型。种皮源于外珠被,内表皮细胞壁螺旋状加厚。胚胎学特征表明,新木姜子属与木姜子属有密切的亲缘关系。较多的双胚囊异常现象,支持樟科与Monimiaceae具有紧密关系的推测。胚胎学特征不支持将无根藤属独立为科的观点。     

白毛新木姜子[Neolitsea aurata var.glauca]胚胎学研究

应用光学显微镜对白毛新木姜子的胚胎学特征进行了研究,首次在非寄生性樟科植物中发现了细胞型胚乳。对樟科8属进行了胚胎学特征的比较。花药四室,药室壁的发育属于"基本型",周原质团型绒毡层。小孢子母细胞连续型分裂。等四面体型四分体。二细胞成熟花粉,无孔沟。雌孢原多个,一般仅一个能继续发育。蓼型胚囊。助细胞具丝状器。反足细胞宿存。大孢子母细胞和合子具极性。宿存的一个助细胞具有吸器功能。细胞型胚乳,胚胎发育属于柳叶菜型的三叶变型。种皮源于外珠被,内表皮细胞壁螺旋状加厚。胚胎学特征表明,新木姜子属与木姜子属有密切的亲缘关系。较多的双胚囊异常现象,支持樟科与Monimiaceae具有紧密关系的推测。胚胎学特征不支持将无根藤属独立为科的观点。     

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.     

Are there symplastic connections between the endosperm and embryo in some angiosperms?—a lesson from the Crassulaceae family

It is believed that there is symplastic isolation between the embryo (new sporophyte) and the endosperm (maternal-parental origin tissue, which nourishes the embryo) in angiosperms. However, in embryological literature there are rare examples in which plasmodesmata between the embryo suspensor and endosperm cells have been recorded (three species from Fabaceae). This study was undertaken in order to test the hypothesis that plasmodesmata between the embryo suspensor and the endosperm are not so rare but also occur in other angiosperm families; in order to check this, we used the Crassulaceae family because embryogenesis in Crassulaceae has been studied extensively at an ultrastructure level recently and also we tread members of this family as model for suspensor physiology and function studies. These plasmodesmata even occurred between the basal cell of the two-celled proembryo and endosperm cells. The plasmodesmata were simple at this stage of development. During the development of the embryo proper and the suspensor, the structure of plasmodesmata changes. They were branched and connected with electron-dense material. Our results suggest that in Crassulaceae with plasmodesmata between the endosperm and suspensor, symplastic connectivity at this cell-cell boundary is still reduced or blocked at a very early stage of embryo development (before the globular stage). The occurrence of plasmodesmata between the embryo suspensor and endosperm cells suggests possible symplastic transport between these different organs, at least at a very early stage of embryo development. However, whether this transport actually occurs needs to be proven experimentally. A broader analysis of plants from various families would show whether the occurrence of plasmodesmata between the embryo suspensor and the endosperm are typical embryological characteristics and if this is useful in discussions about angiosperm systematic and evolution.     

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。     

A developmental and evolutionary analysis of embryology in Platanus (platanaceae), abasal eudicot

The Platanaceae are an early derived eudicot lineage and therefore occupy a key position for understanding reproductive character diversification associated with the early evolutionary radiation of flowering plants. We conducted an embryological study of Platanus racemosa in order to provide critical data on defining angiosperm reproductive characters for this important group. Female gametophyte development is monosporic. Embryogenesis occurs in a series of stages including zygote elongation and division, development of a linear proembryo, formation of the embryo proper, histogenesis, organogenesis, and growth. Endosperm development is a complex process that includes four distinct phases: free nuclear proliferation, cellularization of the chalazal zone, centripetal cellularization of the micropylar zone, and cellular differentiation and growth. Only the outer endosperm layer persists at seed maturity. Our findings differ significantly from previously published reports for Platanus, in which endosperm development was described as ab initio cellular. A comparison of endosperm development in Platanus with several closely and distantly related free nuclear taxa reveals considerable developmental variability, consistent with a hypothesis of multiple origins of free nuclear endosperm in angiosperms. Our analysis indicates that much remains to be learned about embryology in basal angiosperms. Additional developmental and comparative studies will likely reveal critical insights into the early evolution of flowering plants.     

On the Systematic Position of Daphniphyllaceae

The present paper deals with the systematic position of Daphniphyllaceae. The genus Daphniphyllum was first described by Blume in 1826 as a member of Rhamnaceae. In 1858 Baillon removed it to the tribe Phyllantheae of Euphorbiaceae, while Müller (1869) raised this genus to the rank of family, Daphniphyllaceae. Although Müller’s treatment has been accepted by most botanists, including the present authors, its systematic position has been debated. The first aim in our studies on the cladistics of Hamamelidae is to answer the question which families should be included in this monophyletic group. By observing their pollen grains and stoma types of some representative species of Daphniphyllaceae, Hamamelidaceae and Buxaceae under light microscope (LM) and scanning electron microscope (SEM,) and analysing morphological, anatomical, palynological, embryological characters and chemical components in the three taxa and Euphorbiaceae, we find that Daphniphyllaceae is very similar to Hamamelidaceae, but greatly different from Euphorbiaceae, in inflorescence racemose or spicate, calyx nearly reduced, stamens numerous and sometimes synandry, connective usually exserted, disc absent, carpels 2; vessel with scalariform perforation plates and often not spiral-thickened, fiber bordered-pitted; stomata mostly paracytic; pollen 3-colpate; tapetum glandular, endosperm development cellular, obturator and caruncle absent; iridoid compounds present; sieve-element plastids S-type. The present authors have noticed the fact that Daphniphyllaceae is also similar to Magnoliaceae in the stamens numerous, anthers larger and filaments very short, connectives obviously exserted and with several bundles; anther wall thicker, endosperm development cellular, embryo small. It is considered that not only are Daphniphyllaceae and Hamamelidaceae phenetically close to each other but also much possibly derived from a common ancestor, the extinct group of Magnoliales. However, Daphniphyllaceae appears to be remote from Euphorbiaceae and Buxaceae in relationship and should be separated from Euphorbiales and Buxales. Meanwhile, since Daphniphyllaceae differs from the members of Hamamelidales in the incompletely septate ovary, drupaceous fruit, indistinct sexine sculpture of pollen grains, small embryo, and an unique alkaloid, daphniphylline, but lacking proanthacyanins, the establishment of an order, Daphniphyllales, for the family, is considered reasonable. According to our opinion, the order is related to Hamamelidales rather than to Euphorbiales as originally suggested by Huru-sawa (1954).     

利用激光扫描共聚焦显微技术观察同源四倍体双胚苗水稻的胚胎学特征

利用激光扫描共聚焦显微技术对同源四倍体双胚苗水稻的受精过程、胚胎发育过程和双胚来源进行观察鉴定。研究结果表明,同源四倍体水稻和二倍体水稻在发生受精作用的时间上不存在很大的差异。双胚苗水稻的受精作用和胚胎发育表现出明显的特异性,其双胚苗性状有其胚胎学根源,双胚可能有4种来源,即来源于由双套胚囊形成的双胚、由多卵卵器中的卵细胞和类卵细胞分别通过受精作用形成双胚、由特异性的反足细胞通过异常发育后形成额外的幼胚、由胚乳细胞通过异常发育后形成胚乳型幼胚。通过对试验材料的受精频率及其结实率进行比较后发现,双胚苗水稻具有更弱的有性生殖能力,其遗传可塑性更强,这为离子束介导技术的研究找到了比较好的受体材料。     

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。     

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 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.