Spore morphology and ultrastructure of Cyathea (Cyatheaceae,Pteridophyta) species from southern South America

Spore sculpture and wall structure of eight Cyathea (Cyatheaceae) species from southern South America were studied using light microscopy (LM), scanning electron microscopy (SEM) and transmission electron microscopy (TEM) techniques. Two layers, i.e. an inner and an outer layer, were observed in the perispore. The inner layer has two strata: the inner stratum is attached to the exospore and composed of rodlets tangentially oriented to the spore surface and randomly intermixed; the outer stratum consists of a three-dimensional network of rodlets with either free or fused distal edges forming spinules. The outer layer is thin, darkly contrasted and covers the rodlets. In most cases, the exospore has two layers and a pitted surface. In Cyathea atrovirens, the exospore surface is smooth, while in C. delgadii and C. myriotricha it is verrucate. The homogeneity of perispore features within the genus Cyathea is evident, while exospore features are heterogeneous. The exospore has different kinds of surface-structures that are of potential interest for assessing evolutionary trends within the group.     

海金沙孢子壁结构和发育的研究

利用光镜、扫描电镜和透射电镜对海金沙科(Lygodiaceae)海金沙[Lygodium japonicum (Thunb.) Sw.]孢壁的形成和发育进行了研究.结果表明:海金沙孢子壁由内壁、外壁和周壁3部分构成.外壁由2层构成,即薄的内层和厚的外层,其中外层是在四分体分离前通过孢粉素的逐层沉积并浓缩凝聚而形成的均质层,其表面具不明显的疣状突起.周壁由绒毡层残余物在外壁表面逐层沉积形成,可分为周壁内层、周壁中层和周壁外层3部分;周壁中层具辐射状排列的长条形成分,周壁外层形成瘤状纹饰的轮廓.本研究为孢粉学和蕨类植物系统演化分析提供基础资料.     

瓦韦孢子壁的结构和发育的研究

利用光镜、扫描电镜和透射电镜对水龙骨科（Polypodiaceae）瓦韦(Lepisorus thunbergianus (Kaulf.) Ching)孢子壁的结构和发育进行了研究。研究结果表明瓦韦孢子两侧对称、单裂缝,表面具波纹状纹饰。孢壁从内到外由内壁、外壁和周壁三部分构成。外壁来源于绒毡层物质,由外壁内层和外壁外层构成,外壁外层表面的波纹状纹饰形成孢子表面的纹饰轮廓。周壁薄,紧贴外壁表面,由2层片状结构叠合而成。在外壁外层形成过程中,孢子表面和周围出现较多小球。本文探讨了孢壁各层的结构、来源和发育过程,为蕨类植物系统学和孢粉学研究积累资料。     

凤丫蕨孢子壁的结构和发育研究

利用光镜、扫描电镜和透射电镜对裸子蕨科（Hemionitidaceae）凤丫蕨（Coniogramme japonica（Thunb.） Diels）孢壁的结构和发育进行了研究。结果表明,凤丫蕨孢子外壁表面光滑,由2层构成,即薄的内层和厚的外层。周壁分为周壁内层和周壁外层两部分,周壁内层中上部具辐射状排列的小柱状成分,周壁外层由鳞片和小球体疏松交织成平面或立体网状,由两层周壁共同构成孢子表面皱状纹饰的轮廓。探讨了凤丫蕨孢子周壁的来源,为孢粉学和蕨类植物系统演化研究提供基础资料。     

朝鲜介蕨孢子周壁发育的研究

利用光镜、扫描电镜和透射电镜对朝鲜介蕨[Dryoathyrium coreanum(Christ)Tagawa=Lunathyrium coreanum(Christ)Ching]孢子周壁的发育规律进行了研究。结果表明,朝鲜介蕨孢子两侧对称,单裂缝,表面具粗大的脊状褶皱,褶皱形成网状或拟网状纹饰。孢壁包括内壁、外壁和周壁。孢子外壁表面光滑,在四分孢子时期就已发育成熟。四分孢子分离后,周壁开始形成,周壁来源于孢子囊的绒毡层,是由原质型绒毡层的残余物在外壁上沉积而成。成熟的周壁很厚,可分为外层和内层。周壁内有大的空腔,主要是由周壁外层向外隆起形成的,隆起进而形成了孢子的脊状褶皱和表面纹饰。     

乌蕨孢子壁的形成和发育

利用光镜、扫描电镜和透射电镜对鳞始蕨科（Lindsaeaceae）乌蕨（Stenoloma chusanum Ching）孢壁的形成和发育进行了研究。结果表明乌蕨孢子两侧对称、单裂缝,表面具疣状纹饰。孢壁由内壁、外壁和周壁三部分构成。外壁在四分体阶段已基本形成,其表面光滑,质地均匀,由孢粉素形成。周壁是由绒毡层残余物在外壁表面沉积形成,可分为周壁内层、周壁中层和周壁外层三部分。在周壁中层与外层之间有一层均匀的空间。最后,本文探讨了孢壁的形成和发育规律,研究结果对揭示孢子纹饰和孢壁各层的形成过程、来源和稳定性有重要的意义,并为孢粉学和系统学研究提供基础资料。     

Stages in development of<Emphasis Type="Italic"> Selaginella diffusa</Emphasis> megaspores

In mature megaspores of Selaginella diffusa (C. Presl) Spring the units of the exospore are ordered and become unordered toward the outer and inner surfaces. The exospore surface is coated with silica at maturity. The insertion of the future gap begins in early stages with formation of many minigaps within the inner part of the exospore distally. The mesospore, like the exospore, is resistant to the acetolysis reaction and can, thus, provisionally be considered to consist of sporopollenin. Unit structures within the outer part of the mesospore are unordered, but become ordered in the middle and inner parts. The inner surface of the mesospore appears verrucate. In maturing megaspores, the mesospore is mostly disintegrated and the inner exospore, which encapsulated the mesospore, remains as a somewhat isolated structure, and is again near the outer exospore. There are connecting strands across the gap between the inner surface of the outer exospore and the surface of the inner exospore. There are also spheres on the outer surface of the inner exospore. Electronic Publication     

Stages in development of Selaginella diffusa megaspores

In mature megaspores of Selaginella diffusa (C. Presl) Spring the units of the exospore are ordered and become unordered toward the outer and inner surfaces. The exospore surface is coated with silica at maturity. The insertion of the future gap begins in early stages with formation of many minigaps within the inner part of the exospore distally. The mesospore, like the exospore, is resistant to the acetolysis reaction and can, thus, provisionally be considered to consist of sporopollenin. Unit structures within the outer part of the mesospore are unordered, but become ordered in the middle and inner parts. The inner surface of the mesospore appears verrucate. In maturing megaspores, the mesospore is mostly disintegrated and the inner exospore, which encapsulated the mesospore, remains as a somewhat isolated structure, and is again near the outer exospore. There are connecting strands across the gap between the inner surface of the outer exospore and the surface of the inner exospore. There are also spheres on the outer surface of the inner exospore.     

乌蕨孢子壁的形成和发育

利用光镜、扫描电镜和透射电镜对鳞始蕨科(Lindsaeaceae) 乌蕨( Stenoloma chusanum Ching) 孢壁的形成和发育进行了研究。结果表明乌蕨孢子两侧对称、单裂缝, 表面具疣状纹饰。孢壁由内壁、外壁和周壁三部分构成。外壁在四分体阶段已基本形成, 其表面光滑, 质地均匀, 由孢粉素形成。周壁是由绒毡层残余物在外壁表面沉积形成, 可分为周壁内层、周壁中层和周壁外层三部分。在周壁中层与外层之间有一层均匀的空间。最后, 本文探讨了孢壁的形成和发育规律, 研究结果对揭示孢子纹饰和孢壁各层的形成过程、来源和稳定性有重要的意义, 并为孢粉学和系统学研究提供基础资料。     

蜈蚣草孢壁发育及其系统学意义

利用光镜、扫描电镜和透射电镜对凤尾蕨科(Pteridaceae)蜈蚣草(Pteris vittata L.)孢壁的形成和发育进行研究。结果表明:蜈蚣草孢子四面体型,极面观钝三角圆形,赤道面观半圆形或超半圆形,近极面具瘤状纹饰和近极脊,远极面具脊并连成网状,具赤道环;孢子具乌毛蕨型外壁,由外壁外层构成纹饰的轮廓;实心型周壁由2层构成,且内层薄、外层具小球体。结合孢子外壁和周壁的发育特征,认为凤尾蕨科与裸子蕨科和水蕨科的亲缘关系较近,支持将裸子蕨科和水蕨科置于凤尾蕨科。     

华中铁角蕨孢子纹饰形成的超微结构观察

利用透射电子显微镜对铁角蕨科(Aspleniaceae)华中铁角蕨(Asplenium sarelii Hook.)孢子及其纹饰的形成过程进行观察。结果表明:①华中铁角蕨孢子囊发育为薄囊蕨型;②孢子外壁表面光滑,远极面的外壁厚约0.8~1.1μm,近极面的外壁厚约1.4~1.8μm;③孢子周壁厚度约4~5μm,染色较外壁深,分为内层和外层;内层紧帖外壁表面,其上具柱状、瘤状或疣状突起;外层向外隆起形成脊状纹饰的轮廓,脊的下方具空腔,脊的顶端具翅;④铁角蕨型与鳞毛蕨型孢子外壁和周壁纹饰的形成过程具有相似性;⑤孢子的成熟度对于孢子形态的研究是至关重要的,只有完全成熟的孢子的表面纹饰才是稳定的。     

Spore morphology and wall ultrastructure of Hymenophyllaceae Link (Pteridophyta) from north-west Argentina

The family Hymenophyllaceae is represented in the study area by six species in two genera, Hymenophyllum J. E. Smith and Trichomanes L. The study was based on herbarium material and spores were studied under light microscope (LM), scanning electron microscope (SEM) and transmission electron microscope (TEM). Both genera have trilete spores, 23 to 45 μm in equatorial diameter, with an ornamentation of echinulae and cones in Hymenophyllum and of verrucae, gemmae and granules in Trichomanes. Mature spores have a sporoderm composed of a perispore, an exospore and a fibrillar endospore; the exospore is 0.5 to 2.5 μm thick, compact and with an irregular margin. In some cases radial channels and other channels associated with the middle and inner parts of the laesurae were evident. A series of cavities filled with an opaque content line the inner margin of the exospore. The perispore is 20 to 400 nm thick and unevenly differentiated along the surface of a same spore. Under TEM, two main differentially contrasted portions could be distinguished: a dark massive portion with structural components could not be distinguished, and a light portion with several plates arranged in piles. The inner surface of the perispore exhibit short scales. Globules are immersed within the perispore at some depth from the perispore surface and others connected to it by structural threads. The spore characters observed including shape, ornamentation, laesurae length and wall structure are useful in distinguishing the two genera studied, but less useful in differentiation at the species level.     

ULTRASTRUCTURAL STUDY ON MICROSPORE WALL MORPHOGENESIS IN ISOETES JAPONICA (ISOETACEAE)

Spore wall morphogenesis of the microspore of Isoetes japonica was studied by transmission electron microscopy. The microspore wall consists of four layers: the perispore, outer exospore, inner exospore, and endospore. The perispore consists of electron-dense materials. The exospore is divided into outer and inner sections, with a large gap between the two. The outer exospore appears as an undulating plate consisting of tripartite lamellae with homogeneous sporopollenin. The inner exospore consists of an accumulation of tripartite lamellae on the microspore cell membrane. Immediately after meiosis, the tripartite lamellae of the outer exospore forms around the microspore. The lamellated inner exospore forms next, which adheres to the cell membrane of the microspore. The deposition of homogeneous sporopollenin material on the tripartite lamellae causes the plates of the outer exospore to thicken. Some homogeneous material may also be deposited on the inner exospore. Lastly, the electron-dense perispore is deposited on the outer exospore, and the electron-lucent endospore forms beneath the inner exospore. We conclude that the lamellae of the outer exospore, inner exospore, and endospore are formed and derived, in that order, from the gametophytic microspore cytoplasm. The homogeneous sporopollenin material of the outer exospore and perispore may be derived from the sporophytic tapetal cytoplasm.     

红盖鳞毛蕨孢子发育的超微结构和细胞化学研究

采用透射电镜和细胞化学技术对红盖鳞毛蕨(Dryopteris erythrosora(Eaton)O.Ktze.)的孢子发育过程进行了研究,根据超微结构和细胞化学特征可将其孢子发育过程分为3个阶段:(1)孢子母细胞及其减数分裂阶段:孢子母细胞壳在孢原细胞末期开始形成,位于孢子母细胞及其减数分裂形成的四分体外侧,PAS反应显示其为多糖性质,与胼胝质壁为同功结构;在减数分裂形成的四分孢子之间产生孢子外壳,从功能、形成位置和时间上看与胼胝质壁相似,但苏丹黑B反应显示其可能含有脂类物质,与孢子母细胞壳和胼胝质壁不同。(2)孢子外壁形成阶段:外壁为乌毛蕨型(Blechnoidal-type),由薄的多糖性质的外壁内层和表面平滑的孢粉素外壁外层构成;小球参与外壁外层的形成,组织化学分析显示小球的中央区域和外壁外层内侧部分由红色(多糖)变为黄色,小球的表面区域和外壁外层部分始终被染成黑色(脂类),可知小球与外壁同步发育。(3)孢子周壁形成阶段:周壁为凹陷型(Cavate-type),包括2层,内层薄,紧贴外壁,外层隆起形成孢子脊状褶皱纹饰的轮廓,以少见的向心方向发育;苏丹黑B和PAS反应观察周壁被染成橙色,推测其可能由多糖等成分构成;孢子囊壁细胞参与周壁的形成。本研究为揭示蕨类植物孢子发生的细胞学机制提供了新资料。     

Microsporidian Spore Wall: Ultrastructural Findings on Encephalitozoon hellem Exospore

A study of the spore wall of Encephalitozoon hellem was performed on thin sections, freeze-fracture, and deep-etched samples to obtain information on spore wall organization and composition. Our observations demonstrate that the spore wall is formed by an inner 30–35 nm electron-lucent endospore and an outer 25–30 nm electron-dense exospore. The exospore is a complex of three layers: an outer spiny layer, an electron-lucent intermediate lamina and an inner fibrous layer. Freeze-fracture and deep-etching techniques reveal that the intermediate lamina and the inner fibrous layer result from the different spatial disposition of the same 4-nm thick fibrils. In thin sections the endospore reveals a scattered electron-dense material that appears in the form of trabecular structures when analyzed in deep-etched samples. The presence of chitin in the exospore is discussed.     

Microspore morphology ofIsoetes species (lycophyta) from southern South America

Microspores of the 24 species ofIsoetes that grow in southern South America were analyzed under a light microscope and scanning electron microscope. The microspores are monolete, elliptic in polar view, 30–40 μm long, and 20–25 μm wide. A background with various characteristics is seen on each surface. A supra-laesural expansion is present. The perispore is ornamented and has a perforated background. In section, it has a lacunose structure. The exospore is smooth, and it has a compact structure in section. The studied species could be divided into three groups by their perispore ornamentation: equinate, rugulate, and tuberculate. Microspore size was positively correlated with increasing ploidy level, and larger microspores were associated with terrestrial habitats. A convergence in ornamentation was found between spores produced by the studied species and those that grow in regions outside of the area under study.     

Ultrastructural study of spore wall morphogenesis inOphioglossum thermale kom. var.nipponicum (Miyabe et Kudo) nishida

Spore wall morphogenesis ofOphioglossum thermale var.nipponicum was examined by transmission electron microscopy. The spore wall of this species consists of three layers: endospore, exospore, and perispore. The spore wall development begins at the tetrad stage. At first, the outer undulating lamellar layer of the exospore (Lo) is formed on the spore plasma membrane in advance of the inner accumulating lamellar layer (Li) of the exospore. Next, the homogeneous layer of the exospore (H) is deposited on the outer lamellar layer. Both lamellar layers may be derived from spore cytoplasm; and the homogeneous layer, from the tapetum. Then the endospore (EN) is formed. It may be derived from spore cytoplasm. The membranous perispore (PE), derived from the tapetum, covers the exospore surface as the final layer. Though the ornamentation of this species differs distinctly from that ofO. vulgatum, the results mentioned above are fundamentally in accordance with the data obtained fromO. vulgatum (Lugardon, 1971). Therefore, the pattern of spore wall morphogenesis appears to be very stable in the genusOphioglossum.     

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。     

Spore wall ultrastructure of Polypodiaceae from north-western Argentina

The spore wall ultrastructure of Campyloneurum, Microgramma, Pecluma, Phlebodium, Pleopeltis and Serpocaulum (Polypodiaceae) from north-western Argentina has been studied using transmission electron microscopy (TEM). The exospore is 0.4–3 μm thick, two-layered and variously ornamented in all taxa. The exospore surface is distinctive, but in general ultrastructure the exospore is similar in all species studied. The structural elements of the exospore consist of cavities in the inner part as well as channels with a radial orientation and channels at both sides of the laesura. Variation in the exospore surface was observed in spores at different stages of maturation. The perispore is darkly contrasted and 0.04–2 μm thick. Three different structure types were recognised, including fibrillar, multilamellar and lacunose. Scattered globules and spherules were always present on the perispore surface. The structural variability of the perispore was surveyed within complete sporangia. We concluded that the observed variability may be related to the stage in spore maturation and, consequently, to the stages in perispore differentiation. As the exospore ultrastructure is similar and interpreted as related to functional activity in the studied material, it cannot be used for systematic delimitations at this generic or specific level.     

Scanning electron microscope study of cat and dog enamel structure

Scanning electron microscopy revealed several similarities as well as significant differences in the enamel structure between cat and dog teeth. Three enamel layers were present in both species; a surface rodless (aprismatic) layer, an outer layer of parallel rods (only at some sites), and an inner layer with prominent Hunter-Schreger bands. In the inner layer of both carnivores, the diameter of individual rods varied significantly and frequently their course changed abruptly with respect to neighboring rods. In dog teeth the cross-sectional shape of inner enamel rods was pleomorphic, but hexagonal in outer enamel. In contrast, cat enamel rods were rounded in both inner and outer enamel layers. Hunter-Schreger bands of cats circumscribed the teeth in relatively straight segments, but these bands showed pronounced waviness in dog teeth. In cats and dogs the surface rodless layer was structurally continuous with subjacent interrod enamel and covered all tooth surfaces with the exception of the cervical areas. The data show that the structure of inner and outer enamel layers differ between these two carnivore species and that the enamel structure of the cat was most similar to that described in humans. One principal difference between carnivore and human teeth is that the growth lines of carnivores do not terminate at perikymata on the tooth surface.