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 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.     

Sporogenesis and gametogenesis in Sladenia and their systematic implication

Sporogenesis and gametogenesis in the monotypic genus Sladenia is reported for the first time. The anther wall is of the monocotyledonous type of formation and consists of four-layers, namely a glandular tapetum, one middle layer, an endothecium, and an epidermis. Fibrous thickenings are developed in the epidermis at or near the stage of anther maturity rather than in the endothecium. Successive cytokinesis in microsporogenesis results in tetrahedral or rarely isobilateral tetrads of microspores. Pollen grains are two-celled when shed. The ovule is bitegmic, tenuinucellate, and anatropous. The nucellus degenerates during meiosis of the megaspore mother cell. The micropyle is formed by only the inner integument. Development of the female gametophyte conforms to the Adoxa type. Sladenia has been treated either as a member of Actinidiaceae, Dilleniaceae or Theaceae s.l. , or as belonging to an independent family, Sladeniaceae. Recent molecular studies have suggested that it belongs to Ternstroemiaceae within the Ericales s.l . Several noteworthy embryological characters are distinct from those of the other putatively related families and support the recognition of a separate family, Sladeniaceae. Additional embryological evidence confirms this conclusion.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 305–314.     

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.     

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     

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。     

山茶科核果茶属和石笔木属的胚胎学研究

观察了短叶核果茶,石笔木,粗毛石笔木和屏石笔木的大小孢子和雌雄配子体的发生和发育过程,４个种的胚胎学特征高度相似,均为基本型药壁发育,腺质线毡层,同时型小孢子母细胞胞质分裂,四面体形小孢子四分体,二细胞成熟花粉,倒生胚珠,双珠被,薄珠心,单孢原,蓼胚囊,卵细胞与助细胞区分明显,均具有珠被绒毡层和承珠盘,以蓼型胚囊区别于邻近的山茶属。     

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     

北柴胡大小孢子发生和雌雄配子体发育的研究

用石蜡切片法对北柴胡的大、小孢子发生和雌、雄配子体发育过程进行观察研究.结果显示,北柴胡的胚珠为倒生型,单珠被,薄珠心.孢原细胞不经分裂直接发育成大孢子母细胞;四分体线性,多数情况下合点端的大孢子为功能大孢子,少数情况下亚合点端的大孢子也可发育为功能大孢子,蓼型胚囊;八核胚囊时期,珠心基部和两侧的一些珠心细胞保持自己的细胞质和形状,留存较久,成为珠心座细胞,珠被内表皮细胞发育为珠被绒毡层;花药壁发育类型为基本型,绒毡层为腺质绒毡层.小孢子母细胞减数分裂的胞质分裂为同时型,产生正四面体型四分体.成熟花粉为3细胞型.     

白穗花有性器官与胚胎发育形态的研究

利用石蜡切片技术对白穗花(Speirantha gardenii (Hook.) Baill.)胚胎发育过程进行了显微观察。研究结果表明: 白穗 花具四室花药, 腺质绒毡层; 小孢子母细胞减数分裂为连续型, 四分体主要为左右对称型, 偶有直线型、T型和交互对生型; 成熟花粉为二细胞型; 子房三室, 中轴胎座; 倒生胚珠, 双珠被, 厚珠心型, 珠孔由内珠被形成; 胚囊的发育为蓼型; 胚的发育类型为柳叶菜型, 核型胚乳。根据已有的胚胎学资料, 比较白穗花和铃兰族(Convallarieae)其它植物的胚胎学特征, 结果显示: (1) 白穗花属(Speirantha Baker)与蜘蛛抱蛋属(Aspidistra Ker-Gawler)的胚胎学特征更为接近, 二者可能起源于共同的祖先; (2) 夏须草属(Theropogon Maxim.)是铃兰族的异质类群。     

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.     

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.     

条纹龙胆的胚胎学研究

首次报道了条纹龙胆（Ｇｅｎｔｉａｎａ ｔｓｒｉａｔａ Ｍａｘｉｍ．）的胚胎学特征,研究结果用以讨论龙胆属狭蕊组（Ｇｅｎｔｉａｎａ Ｓｅｃｔ．Ｓｔｅｎｏｇｙｎｅ）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;绒毡层细胞仅来源于初生壁细胞,故绒毡层起源属单型起源,细胞单核,原位退化,属腺质型绒毡层,药隔处的绒毡层细胞经多次平周分裂形成２层至多层的绒毡层细胞,其余部位的绒毡层细胞仍为１     

白穗花有性器官与胚胎发育形态的研究

利用石蜡切片技术对白穗花（Speirantha gardenii（Hook.）Baill.）胚胎发育过程进行了显微观察。研究结果表明：白穗花具四室花药,腺质绒毡层;小孢子母细胞减数分裂为连续型,四分体主要为左右对称型,偶有直线型、T型和交互对生型;成熟花粉为二细胞型;子房三室,中轴胎座;倒生胚珠,双珠被,厚珠心型,珠孔由内珠被形成;胚囊的发育为蓼型;胚的发育类型为柳叶菜型,核型胚乳。根据已有的胚胎学资料,比较白穗花和铃兰族（Convallarieae）其它植物的胚胎学特征,结果显示：（1）白穗花属（Speirantha Baker）与蜘蛛抱蛋属（Aspidistra Ker-Gawler）的胚胎学特征更为接近,二者可能起源于共同的祖先;（2）夏须草属（Theropogon Maxim.）是铃兰族的异质类群。     

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。     

西南獐牙菜的胚胎学及其系统学意义

报道了西南獐牙菜的大、小孢子发生及雌,雄配子体发育过程,并以此讨论了獐牙菜属宽丝组和多枝组的分类等级和系统演化关系,西南獐牙菜花药四室;药壁发育为双子叶型;绒毡层二型起源,腺质型;中层细胞2层;药室内壁纤维状加厚,药壁表皮宿存,小孢子母细胞减数分裂为同时型;小孢子四分体的排列为四面体形,成熟花粉为3-细胞型,孔子房为3心皮,1室,胚珠12列,故为超侧膜胎座,薄珠心,单珠被,倒弯生胚珠,大孢子母细胞减数分裂形成4个大孢子呈直线形排列,合点端的大孢子具功能,胚囊发育为蓼型,两极核在受精前融合为次生核,3个反足细胞次生增殖为5-8个,宿存,比较宽丝组和多枝组的胚胎党性一状表明宽丝组从多枝组中分出是合理的,在系统位置上宽丝组较多枝组进化。     

Embryology of Hortonioideae and Monimioideae (Monimiaceae,Laurales): characteristics of the ‘lower’ monimioids

We investigated the embryology of the ‘lower’ monimioids, i.e. Monimioideae (Monimia, Palmeria and Peumus) and Hortonioideae (Hortonia), which are poorly described embryologically. Our results show that, contrary to what has been reported in the literature, ‘lower’ monimioids show very little variation in their embryological characters. Comparisons with Mollinedioideae (a large derived subfamily in Monimiaceae) and other families in Laurales show that the ‘lower’ monimioids are relatively consistent in sharing predominantly isobilateral tetrads of microspores and megaspores, a non‐specialized chalaza, and a mesotestal–endotestal seed coat (with tracheoidal cells of the meso‐ and endotesta). It is likely that, while the shared successive cytokinesis during meiosis of microspore mother cells supports the Monimiaceae–Hernandiaceae–Lauraceae clade obtained by molecular evidence, no synapomorphies exist to support a sister‐group relationship of Monimiaceae with Hernandiaceae or Lauraceae. Instead, the lack of hypostase in ovules and/or young seeds, the lack of endosperm in mature seeds and the amoeboid tapetum in the anther are likely synapomorphies of Hernandiaceae and Lauraceae. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 228–241.     

家榆胚胎学研究

采用常规石蜡切片法,借助光学显微镜对家榆胚胎发育进行研究,研究结果表明：（1）家榆的花药具4个药室;小孢子母细胞减数分裂过程中的胞质分裂为同时型;小孢子四分体排列为四面体型和左右对称型;成熟花粉为2-细胞型;（2）花药壁发育类型为双子叶型,腺质绒毡层;（3）家榆子房两心皮1室,侧膜胎座,1枚倒生胚珠,双珠被,厚珠心;四分体呈直线型和T型排列;（4）家榆胚囊类型为单孢子蓼型胚囊;胚的发育类型为茄型;核型胚乳;无胚乳型种子。     

The embryology and systematic relationships of Prionium serratum (Juncaceae: Juncales)

Although Prionium is included in Juncaceae, rbcL sequence data indicate that Juncaceae is paraphyletic, with most genera closer to Cyperaceae than to Prionium. Cyperaceae and Juncales have embryological synapomorphies: thus embryology is used to test the monophyly of Juncaceae. The embryology of Prionium is described and its systematic position discussed. Material was prepared using standard methods of paraffin embedding. Additional embryological data were extracted from the literature. The anther in Prionium is tetrasporangiate, and the wall has an epidermis, an endothecium and middle layer, and an irregularly bilayered, glandular secretory tapetum. Microsporogenesis is probably simultaneous; pollen is ulcerate with a granular exine, in tetrahedral and cross tetrads, and trinucleate at release. The trilocular ovary contains many crassinucellate ovules probably having a Polygonum-type embryo sac. Endosperm is helobial and the embryo is of the Onagrad type, Juncus variation. The seed is testal-tegmic and germination is epigeal. The embryology of Prionium is most like that of Juncaceae, which shares several synapomorphies with Cyperaceae. Some of the characters in Cyperaceae may be interpreted as specialized forms of those found in Juncaceae. Embryology supports the monophyly of Cyperaceae and Juncales, but not Juncaceae; thus the position of Prionium remains unresolved.     

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.