蕨类植物紫萁绒毡层细胞凋亡的细胞学特征

绒毡层凋亡过程是小孢子发生中的重要事件,以往的研究主要集中在被子植物,蕨类植物尚未见此方面的报道。该研究首次采用透射电镜和免疫荧光技术对蕨类植物紫萁(Osmunda japonica Thunb.)绒毡层细胞凋亡的细胞学过程进行了观察,以明确紫萁绒毡层细胞的发育类型和凋亡特征,为蕨类植物绒毡层细胞凋亡的深入研究以及孢子发育研究提供依据。结果显示：(1)紫萁的绒毡层属于复合型,即外层绒毡层为分泌型,该层细胞发育过程中液泡化,营养物质被吸收;内层绒毡层为原生质团型,经历了细胞凋亡的过程。(2)绒毡层内层细胞在凋亡过程中细胞壁和细胞膜降解,细胞质浓缩且空泡化;细胞核内陷、变形,染色质浓缩凝聚,形成多数小核仁,DAPI荧光由强变弱;线粒体、质体、内质网、高尔基体等细胞器逐渐退化,液泡中多包含纤维状物、絮状物、黑色嗜锇颗粒和小囊泡等;出现多泡体、多膜体和细胞质凋亡小体,上述特征与种子植物绒毡层凋亡特征基本一致。(3)与种子植物相比,紫萁绒毡层的细胞凋亡开始得早,在整个凋亡过程中没有核凋亡小体的产生;除了产生孢粉素外,绒毡层细胞内产生了大量的丝状物质、絮状物质和电子染色暗的颗粒物,这些物质可能用于...     

白头翁绒毡层和周绒毡层膜的发育和组织化学研究

白头翁绒毡层为分泌型,发育过程分为：发生－分化,生长－合成和分泌－解体3个阶段。第1阶段,绒毡层细胞的形态结构和组织化学特征同造孢组织和花粉母细胞的大体一致,所不同的是大量积累淀粉和形成少量原乌氏体,第2阶段为重要的合成时期;细胞体积膨大,具多倍体双核或畸形核,蛋白质大量合成,淀粉水解;原乌氏体和孢粉素荧光物质大量累积。第3阶段为分泌最活跃时期,裸细胞在妨行分泌功能上起着重要作用。原乌氏体,荧光物质,碳水化合物,或许包括分解胼胝质壁和纤维素壁酶等成分均在裸细胞阶段排出,周绒毡层膜,乌氏体和花粉外壁均属于孢粉素性质的结构,根据周绒毡层膜的外切向位置和结构,初步结论：它可能是绒毡层细胞质膜向着药壁中层一面的残存部分,而膜内表面上的孢粉素纹理则是后来添加的。乌氏体的基本形态为短颈烧瓶状,单个或2－5个组成复合结构,本文对绒毡层,绒毡层膜和乌氏体的功能,及孢粉素的转运和聚合等问题进行了讨论。     

白头翁绒毡层和周绒毡层膜的发育和组织化学研究

白头翁绒毡层为分泌型,发育过程分为：发生－分化,生长－合成和分泌－解体3个阶段。第1阶段,绒毡层细胞的形态结构和组织化学特征同造孢组织和花粉母细胞的大体一致,所不同的是大量积累淀粉和形成少量原乌氏体,第2阶段为重要的合成时期;细胞体积膨大,具多倍体双核或畸形核,蛋白质大量合成,淀粉水解;原乌氏体和孢粉素荧光物质大量累积。第3阶段为分泌最活跃时期,裸细胞在妨行分泌功能上起着重要作用。原乌氏体,荧光物质,碳水化合物,或许包括分解胼胝质壁和纤维素壁酶等成分均在裸细胞阶段排出,周绒毡层膜,乌氏体和花粉外壁均属于孢粉素性质的结构,根据周绒毡层膜的外切向位置和结构,初步结论：它可能是绒毡层细胞质膜向着药壁中层一面的残存部分,而膜内表面上的孢粉素纹理则是后来添加的。乌氏体的基本形态为短颈烧瓶状,单个或2－5个组成复合结构,本文对绒毡层,绒毡层膜和乌氏体的功能,及孢粉素的转运和聚合等问题进行了讨论。     

麻疯树花药发育过程中Ca2+的分布特征

采用超薄切片技术,在透射电镜下观察麻疯树(Jatropha curcasL.)花药发育过程中Ca2 的分布特征。在孢原细胞时期的花药中几乎看不到Ca2 沉淀,但花药维管束周围的细胞中有较多的Ca2 沉淀;到小孢子母细胞时期,细胞质中Ca2 沉淀依然较少,绒毡层壁上Ca2 沉淀明显增多;四分体形成时,小孢子细胞质和绒毡层细胞质中出现了较多的Ca2 沉淀;在小孢子发育早期,细胞质中Ca2 沉淀增加不明显,花粉壁部位累积有很多的Ca2 沉淀,绒毡层中Ca2 沉淀数量达到最多;到小孢子发育晚期,小孢子大液泡的液泡膜上有大量的Ca2 沉淀,绒毡层中Ca2 沉淀明显减少;随着二胞花粉中的大液泡消失,细胞质中积累淀粉粒以后,花粉中看到的Ca2 沉淀极少,同时,在花药维管束周围的薄壁细胞中,又出现了较多的Ca2 沉淀,表明花粉对Ca2 的需求可能降低。麻疯树花药发育过程中钙的动态分布特征暗示着钙参与了调控花粉发育过程,Ca2 的运输途径是由药隔薄壁组织运输到绒毡层,再进一步转移到小孢子表面和细胞质中,整个花药发育过程中,Ca2 沉淀表现为少—增加—减少的变化趋势。     

含笑花药发育的超微结构研究

对含笑花药发育中的超微结构变化进行观察,结果显示:(1)花粉发育中有三次液泡变化过程——第一次是小孢子母细胞在形成时内部出现了液泡,这可能与胼胝质壁的形成有关;第二次是在小孢子母细胞减数分裂之前,细胞内壁纤维素降解区域形成液泡,它的功能可能是消化原有的纤维素细胞壁;第三次是在小孢子液泡化时期,形成的大液泡将细胞核挤到边缘,产生极性。(2)含笑花粉在小孢子早期形成花粉外壁外层,花粉外壁内层在小孢子晚期形成,而花粉内壁是在二胞花粉早期形成;花粉成熟时,表面上沉积了绒毡层细胞的降解物而形成了花粉覆盖物。研究认为,含笑花粉原外壁的形成可能与母细胞胼胝质壁有关,而由绒毡层细胞提供的孢粉素物质按一定结构建成了花粉覆盖物。     

莴苣花药发育过程中钙的分布特征

减数分裂前,莴苣花药中的钙颗粒很少。减数分裂后,花药绒毡层细胞中的钙颗粒明显增加。同时在花药药室基质中也出现许多细小的钙颗粒。刚从四分体中释放出的小孢子内钙颗粒很少。伴随着花粉外壁物质在小孢子表面的沉积,钙颗粒开始积累在花粉壁部位。随后。小孢子中开始出现钙颗粒。当小孢子开始形成液泡后,钙颗粒向其中聚集,伴随着小液泡融合成大液泡。体积较大的钙颗粒主要集中在液泡中,而细胞质基质中的钙颗粒很少。随着二胞花粉中的大液泡消失,花粉细胞质中的钙颗粒变得很少。在以后的发育中,只有花粉壁中积累较多的钙颗粒。在莴苣花药发育过程中,钙与绒毡层细胞的退化和小孢子液泡形成以及二胞花粉中大液泡的消失有关。而花粉外壁表面积累丰富的钙与以后花粉的萌发有关。     

莴苣花药发育过程中钙的分布特征

减数分裂前,莴苣花药中的钙颗粒很少。减数分裂后,花药绒毡层细胞中的钙颗粒明显增加, 同时在花药药室基质中也出现许多细小的钙颗粒。刚从四分体中释放出的小孢子内钙颗粒很少,伴随着花粉外壁物质在小孢子表面的沉积,钙颗粒开始积累在花粉壁部位。随后,小孢子中开始出现钙颗粒。当小孢子开始形成液泡后,钙颗粒向其中聚集,伴随着小液泡融合成大液泡,体积较大的钙颗粒主要集中在液泡中,而细胞质基质中的钙颗粒很少。随着二胞花粉中的大液泡消失,花粉细胞质中的钙颗粒变得很少。在以后的发育中,只有花粉壁中积累较多的钙颗粒。在莴苣花药发育过程中,钙与绒毡层细胞的退化和小孢子液泡形成以及二胞花粉中大波泡的消失有关。而花粉外壁表面积累丰富的钙与以后花粉的萌发有关。     

红豆草根瘤侵染细胞中液泡内含物的超微结构特征

用透射电镜研究了红豆草（Onobrychis viciifolia）根瘤侵染细胞中液泡内含物的超微结构特征。结果表明,早期发育侵染细胞的液泡中只含有少量的纤维状物质。随着细胞的发育,液泡不断变大,液泡中的纤维状物质和膜状物质越来越多。在中央液泡形成后,液泡中的纤维状物质逐渐减少,类细胞质、泡状和膜状物质明显增多,它们常由一层来自液泡膜的膜包围,其形状一般近似圆形或椭圆形。液泡内含物的大量出现可能与红豆草及其根瘤具有高度的抗旱件有关。     

白皮松绒毡层细胞超微结构的研究

白皮松绒毡层细胞的细胞器是十分丰富的,其中粗糙内质网、核糖体和造粉体在减数分裂过程中达到高峰。小孢子形成时,绒毡层细胞开始解体,内质网和线粒体是最后衰老的细胞器。在单核花粉形成时,处于绒毡层细胞外切向壁上的周绒毡层膜特别明显。但是,孢粉素体却主要分布在内切向壁和径向壁上。有趣的是,绒毡层细胞中的造粉体结构与壁层细胞不同。而且,脂体在二分体阶段基本消失,相反,此刻孢粉素体却在质膜外大量聚集。推测脂体的消长可能与原乌氏体和孢粉素体的形成有关。     

凤仙花花药发育中多糖和脂滴组织化学研究

以不同发育时期的凤仙花花药为实验材料,采用组织化学方法,对花药发育中的结构变化及多糖和脂滴物质分布进行观察。结果表明:(1)凤仙花的花药壁由6层细胞组成,包括1层表皮细胞,2层药室内壁细胞,2层中层细胞和1层绒毡层细胞。其中绒毡层细胞的形态不明显,很难与造孢细胞区分,且在小孢子母细胞时期退化。(2)在小孢子母细胞中出现了一些淀粉粒,但减数分裂后,早期小孢子中的淀粉粒消失,又出现了一些小的脂滴;随着花粉的发育,小孢子形成大液泡,晚期小孢子中的脂滴也消失;小孢子分裂形成二胞花粉后,营养细胞中的大液泡降解、消失,二胞花粉中又开始积累淀粉;接近开花时,成熟花粉中充满细胞质,其中包含了较多的淀粉粒和脂滴。(3)在凤仙花的花药发育中,绒毡层细胞很早退化,为小孢子母细胞和四分体小孢子提供了营养物质;其后的中层细胞退化则为后期花粉发育提供了营养物质。     

紫萁孢子囊发育中多糖和脂滴的细胞化学观察

采用光镜、透射电镜和细胞化学技术,对紫萁孢子囊发育过程中孢壁的超微结构和孢子囊内多糖和脂滴的分布及其动态变化进行研究,以探讨紫萁孢子囊发育过程中多糖和脂滴的代谢特征,为蕨类孢子发生的研究提供基础资料。结果表明:(1)紫萁孢子囊由1层囊壁细胞、2层绒毡层和产孢组织构成。(2)紫萁孢子壁由发达而分2层的外壁(外壁内层和外壁外层)和薄的不连续的周壁构成,由外壁形成棒状纹饰的轮廓;孢子外壁内层由多糖类物质构成,外壁外层和周壁均含有脂类物质。(3)在紫萁孢原细胞中观察到少量脂滴;随着紫萁孢壁的形成,囊壁细胞中淀粉粒的大小逐渐变小、数目先增加后减少,它们转运到内层绒毡层原生质团并转化为孢粉素前体物质,再穿过原生质团内膜表面进入囊腔,成为孢粉素团块或以小球形式填加到孢子表面形成孢壁。(4)紫萁孢子囊将多糖类营养物质转化为脂类,以脂滴的形式储藏在孢子中。     

白菜核雄性不育花药超微结构的研究

对白菜核雄性不育两用系的可育与不育花药进行了超微结构的比较观察。结果显示不育花药的造孢细胞核仁靠边分布;包裹小孢子母细胞的胼胝质厚薄不均匀,不完整等早期异常现象。减数分裂后．四分体细胞中常有多个细胞核。从四分体释放出的小孢子外壁的孢粉素物质不均匀沉积,呈不连续的单层异常结构。最后小孢子通过细胞质收缩方式败育。在可育花药中．绒毡层细胞在小孢子发育后期已显示出退化迹象,同时在细胞中开始积累脂类物质。但在同时期的不育花药中．绒毡层细胞没有显示出退化的迹象,也不合成脂类物质。从时间上看,败育花药中小孢子母细胞及小孢子的异常在先,绒毡层细胞的异常在后。本研究揭示了白菜核雄性不育花药的超微结构特征．对我们以前的光学显微镜观察结果予以补充和修正。     

白菜核雄性不育花药超微结构的研究

对白菜核雄性不育两用系的可育与不育花药进行了超微结构的比较观察。结果显示不育花药的造孢细胞核仁靠边分布:包裹小孢子母细胞的胼胝质厚薄不均匀,不完整等早期异常现象。减数分裂后,四分体细胞中常有多个细胞核。从四分体释放出的小孢子外壁的孢粉素物质不均匀沉积．呈不连续的单层异常结构。最后小孢子通过细胞质收缩方式败育。在可育花药中,绒毡层细胞在小孢子发育后期已显示出退化迹象,同时在细胞中开始积累脂类物质。但在同时期的不育花药中, 绒毡层细胞没有显示出退化的迹象,也不合成脂类物质。从时间上看,败育花药中小孢子母细胞及小孢子的异常在先,绒毡层细胞的异常在后。本研究揭示了白菜核雄性不育花药的超微结构特征, 对我们以前的光学显微镜观察结果予以补充和修正。     

The rôle of the tapetum in the formation of sporopollenin-containing structures during microsporogenesis in Pinus banksiana

Summary In the microsporangium of Pinus the outer layer of the peritapetal membrane and the pro-orbicular cores are not only formed in a similar manner, but are composed of apparently identical materials. Precursors for this lipoidal material are produced by the tapetal protoplasts, as are the precursors of sporopollenin. Production the precursors is sequential and appears to involve different cytoplasmic structures.The sporopollenin synthesised by the tapetum condenses upon the pro-orbicular cores, the peritapetal membrane, the exine initials and, on fragmentation of the tapetum, parts of the disintegrating cytoplasm. The evident unpolarised nature of the tapetal protoplasts, and the sequential nature of the synthesis of the lipoid and the sporopollenin by them, may point to orbicule formation in gymnosperms being a necessary by-product of the development of the peritapetal membrane.     

The Changes in the Tapetum of Pinus banksiana Accompanying Formation and Maturation of the Pollen

As meiosis is completed, and following the synthesis of lipidduring meiotic prophase, the tapetum begins to form precursorsof sporopollenin. These accumulate in cisternae of the endoplasmicreticulum, resembling large dictyosome vesicles. They are releasedfrom the tapetal protoplasts intact, but rupture in the loculus.The liberated precursors polymerize either on lipid dropletsin the expanded tapetal walls, forming the orbicules, or onthe lipid layer surrounding the loculus, forming the secondcomponent of the peritapetal membrane. On rupture of the callosewall condensation also proceeds on the walls of the meiospores,already coated with a thin layer of sporopollenin synthesizedby the spore itself. The tapetal protoplasts expand considerablyduring synthesis of the precursors. Wide channels also formbetween the protoplasts, and the nuclei undergo irregular divisions. Ribosomes are conspicuous in the tapetal cytoplasm during thesporopollenin synthesis, but protein levels are low. It is proposedthat protein is exported to the loculus and untimately incorporatedinto the developing microspores. In the final phase of microsporogenesis the tapetum fragments,and parts move into the loculus.Protein levels in the tapetumare now high, possible indicating the massive synthesis of hydrolaseswhich accomplish the dissolution of the tissue. Removal of thelipid component of the peritapetal membrane precedes the desiccationof the anther. The surfaces of the mature pollen lack organizedor irregular deposits of tapetal debris.     

Cytological differentiation of the interrenal tissue of the Japanese quail,Coturnix coturnix

Summary As reported for several other avian species there are clearly distinguishable subcapsular (SCZ) and inner (IZ) zones of interrenal tissue in the Japanese quail. The SCZ contains large columnar cells (type I) with rounded nuclei, polymorphic mitochondria with shelf-like cristae, and relatively small numbers of lipid droplets. The IZ contains two and possibly three types of cells. Type II consists of large columnar cells with moderately dense cytoplasm containing large numbers of lipid droplets and many rounded mitochondria with tubular cristae. Smooth endoplasmic reticulum (SER) and Golgi apparatus are well developed; coated vesicles occur in the Golgi area and at the cell surface. Type-III cells occur in IZ and especially in its more peripheral areas. They are columnar cells with strikingly clear cytoplasm (in comparison with type II) containing mitochondria with plate-like cristae and tubular SER. Type-IV cells are sparsely distributed in IZ and occur rarely in SCZ. Type IV may be a degenerating phase of type III.After adenohypophysectomy or section of portal vessels type-I cells atrophy somewhat with a decrease in lipid droplets; type-II cells, also atrophy with conspicuous increase in size and number of lipid droplets, enlargement of mitochondria, and gradual disappearance of SER; type-III cells decrease in number whereas type-IV cells increase.After injection of ACTH, type-I cells enlarge and their mitochondria, SER and Golgi apparatus become more conspicuous; there is a decrease in lipid droplets in type-II cells and a development of SER, polysomes and Golgi apparatus; there is also a decrease in lipid droplets and a development of SER in type-III cells after injection of 2IU ACTH and an almost complete disappearance of lipid droplets after 4IU ACTH; type-IV cells increase in number.The investigation reported herein was supported by Scientific Research Grants from the Ministry of Education of Japan to Professor Mikami; and by grants from the Japan Society for the Promotion of Science, the National Science Foundation (USA), and the Graduate School Fund of the University of Washington to Professor Farner     

Pollen and Tapetum Development in Male Fertile Rosmarinus Officinalis L. (Lamiaceae)

The development of microspores/pollen grains and tapetum was studied in fertile Rosmarinus officinalis L. (Lamiaceae). Most parts of the cell walls of the secretory anther tapetum undergo modifications before and during meiosis: the inner tangential and radial cell walls, and often also the outer tangential and radial wall, acquire a fibrous appearance; these walls become later transformed into a thin poly-saccharidic film, which is finally dissolved after microspore mitosis. Electron opaque granules found within the fibrous/lamellated tapetal walls consist of sporopollenin-like material, but cannot be interpreted as Ubisch bodies. The middle lamella and the primary wall of the outer tangential and radial tapetal walls remain unmodified, but get covered by an electron opaque, sporopollenin-like layer. Pollenkitt is formed only by lipid droplets from the ground plasma and/or ER profiles, the plastids do not form pollenkitt precursor lipids. Tapetum maturation (“degeneration”) does not take place before late vacuolate stage.

The apertures are determined during meiosis by vesicles or membrane stacks on the surface of the plasma membrane. The procolumellae are conical, but at maturity the columellae are more cylindrical in shape. The columellar bases often fuse, but a genuine foot layer is lacking. The formation of the endexine starts with sporopollenin-accumulating white lines adjacent to the columellar bases. Later, the endexine grows more irregularly by the accumulation of sporopollenin globules. In mature pollen the intine is clearly bilayered.

Generative cells (GCs) and sperm cells contain a comparatively large amount of cytoplasm, and organelles like mitochondria, dictyosomes, ER, and multi-vesicular bodies, but no plastids; GCs and sperms are separated from the vegetative cell only by two plasma membranes.     

Steroidogenic characteristics of in vitro cultured epididymal epithelial cells of the rat

The paper presents the steroidogenic features of cultured epithelial cells of rat epididymis and their ability to synthesize steroid hormones. The cytoplasm of epididymal epithelial cells accumulated lipid droplets and contained active enzymes of steroidogenesis. Numerous mitochondria with lamellar cristae occurred near lipid droplets. Frequently, mitochondria formed a direct contact with lipid droplets and smooth endoplasmic reticulum. The hormone assay showed that the epididymal epithelial cells cultured without dihydrotestosterone synthesized and released the following steroids: dehydroepiandrosterone (DHEA), testosterone (T) and 17beta-estradiol (E). The levels of DHEA and T were very low. The concentration of E detected in media of cultured epididymal epithelial cells exceeded many times the concentration of E in control media. The cytoplasmic presence of organelles and enzymes that participated in the steroid synthesis indicated their similarity to steroidogenic cells. Epididymal epithelial cells were capable of moderate in vitro synthesis of androgens. It cannot be excluded that steroidogenesis in the cultured epididymal epithelial cells is maintained to sustain 17beta-estradiol synthesis pathways.     

Development and Histochemical Observations of Tapetum and Peritapetal Membrane in Anther of Pulsatilla chinensis

Tapetum of Pulsatilla chinensis is of secretory type. Its development proceeds rapidly in following sequence: (1) The stage of initiation-differentiation. At this stage cytological and histochemical features have been described in detail in this paper. (2) The stage of growth- synthesis: This stage appears to be the most important anabolic phase during the development of the tapetum. The salient features are that the tapetal cells become relatively enlarged and form two polyploid nuclei or aberrent polyploid nuclei resulting in synthetizing maximum proteins, fluorescing substances and maximum fluorescent Pro-Ubisch bodies in the tapetal cytoplasm. (3) The stage of secretion-disorganization: After the disintegration of the tapetal wall the enlarged naked cells appear at once. This is an important secretion period in which Pro-Ubisch bodies as well as all other fluorescing substances, carbohydrate or some enzymes are released into anther loculus. The naked cell layer becomes disorgnized until the beginning divition of the pollen grains into two ceils. As to peritapetal membrane of P. chinensis, mainly based on the membrane being on the outer side of the tapetum enclosing both the pollen, tapetal cytoplasm and Ubisch bodies, and the cellular configurations facing the pollen, Authors postulate that peritapetal membrane might be survival of the cytoplasmic membrane of tapetal cells. However, the peritapetal membrane of P. chinensis is similar to that of plasmodial, tapetum reported in certain Compositae and that of secretory tapetum reported in Pinus banksiana. Heslop-Harrison and Gupta et al. had conceded that the tapetal and peritapetal membrane belong to the general class of sporopollenin. On the contrary in P. chinensis the sporopollenin property of peritapetal membrane is only confined to its inner surface. But the thin mem- brane itself with the reticulate sporopollenin attched on its inner side appears negative staining reactions for sporopollenin though it has an ability to resist the acetolysis as well. In P. chinensis the Ubisch body is short necked flask shaped and their size is very similar. Ubisch body is either single or 2–5 in a group, resulting in compound bodies. When the Pro-Ubisch body is still within the tapetal cell it shows positive fluorescent reaction, while it eomletely unstains with Teluidine blue O. So Authors infer that the sporopollenin precur- sors may have permeated through Pro-Ubisch bodies. Finally, How sporopollenin precursor is synthesized in the tapetal cells, transported to pollen locula and polymerized into the sporopollenin on pollen, Ubisch body and peritapetal membrane? Future works along these problems may yield fruitful results.     

Sex pheromone gland of the female European corn borer moth,Ostrinia nubilalis (Lepidoptera,Pyralidae): ultrastructural and biochemical evidences

The sex pheromone gland of the female European corn borer moth, Ostrinia nubilalis was studied using light and electron microscopy. The pheromone gland is formed by hypertrophied epidermal cells at the mid-dorsal region of the intersegmental membrane between abdominal segments 8 and 9/10. Active glandular cells contain extensive apical membrane foldings, a single nucleus, many free ribosomes, numerous mitochondria, microtubules and lipid droplets. Smooth endoplasmic reticulum is scanty. In young moths, the glandular cells are smaller in size, the microvilli at the apical membrane are poorly developed and the cytoplasm contains fewer mitochondria, microtubules, and no lipid droplets. The surrounding unmodified epidermal cells are small cuboidal or squamous cells. These cells have ill-defined apical membrane foldings and do not contain lipid droplets in the cytoplasm and the overlying cuticle. Fatty acids analyses revealed the presence of the sex pheromone components, (E)-11-tetradecenyl acetate, and their immediate precursors, methyl (E)-11- and methyl (Z)-11-tetradecenoate, only in the dorsal portion of the cylindrical intersegmental membrane. Results of the present study show that the sex pheromone gland of O. nubilalis is restricted to the dorsal aspect of the intersegmental membrane between segments 8-9/10 and is not a ring-gland.