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.     

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。     

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。     

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 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.     

Spore wall ultrastructure in Anogramma species (Pteridaceae) from Argentina

The sporoderm ultrastructure of Anogramma Link species, which grow in Argentina, was studied using scanning and transmission electron microscopy. The species A. chaerophylla (Desv.) Link and A. lorentzii (Hieron.) Diels were studied with TEM for the first time. The spores of both species have a sculptured, apparently two-layered exospore. The perispore is uniformly thickened on the whole surface: in A. chaerophylla it is three-layered, while, in A. lorentzii it is single-layered with a complex structure. Spherules are present on the perispore surface or incorporated into the structure of A. chaerophylla while, globules exist on and within the perispore in A. lorentzii. The sporoderm ultrastructure in these two species was compared with other cingulate genera within the Pteridaceae. The characteristics found in this work, with respect to spore wall structure and general morphology, suggest that these characters may differentiate species within genus.     

水蕨孢子壁的形成和发育

利用光镜、扫描电镜和透射电镜对水蕨科(Parkeriaceae)水蕨(Ceratopteris thalictroides (L.) Brongn.)孢子壁的形成和发育进行了研究。结果表明, 水蕨孢子呈辐射对称, 三裂缝, 表面具肋条状纹饰。孢子壁由内壁、外壁和周壁三部分构成。在四分体阶段外壁已基本形成, 其外壁显著, 表面光滑, 质地均匀, 由孢粉素形成, 外壁厚约3-5 μm, 脊高约5-7 μm。周壁由绒毡层残余物在外壁表面沉积形成, 较薄, 厚度只有0.1 μm, 表面具有杆状突起。研究结果对揭示孢子纹饰和孢子壁各层的形成过程、来源和稳定性有一定的意义, 并为蕨类植物孢粉学和系统学研究提供基础资料。     

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.     

水蕨孢子壁的形成和发育

利用光镜、扫描电镜和透射电镜对水蕨科（Parkeriaceae）水蕨（Ceratopteris thalictroides（L．）Brongn．）孢子壁的形成和发育进行了研究。结果表明,水蕨孢子呈辐射对称,三裂缝,表面具肋条状纹饰。孢子壁由内壁、外壁和周壁三部分构成。在四分体阶段外壁已基本形成,其外壁显著,表面光滑,质地均匀,由孢粉素形成,外壁厚约3—5μm,脊高约5—7μm。周壁由绒毡层残余物在外壁表面沉积形成,较薄,厚度只有0．1μm,表面具有杆状突起。研究结果对揭示孢子纹饰和孢子壁各层的形成过程、来源和稳定性有一定的意义,并为蕨类植物孢粉学和系统学研究提供基础资料。     

团扇蕨孢子发生和发育的显微观察

利用光学显微镜对膜蕨科（Hymenophyllaceae）团扇蕨(Gonocormus minutus(Blume) Bosch)孢子的发生和发育进行了观察。研究结果表明：团扇蕨孢子为多边圆形,三裂缝不明显,外壁表面光滑,周壁薄,紧贴外壁表面,由周壁形成乳头状或颗粒状纹饰。在外壁形成后期,孢子表面和囊腔中出现大量小球;在周壁形成时期,孢子表面和周围出现较多小球体;小球和小球体参与孢子壁的形成。团扇蕨绒毡层为混合型,内层为周原质团绒毡层;外层为腺质型绒毡层。本文为膜蕨科系统演化和发育生物学研究提供依据。     

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。     

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.     

Spore morphology and wall ultrastructure of Trachypteris species (Pteridaceae)

Trachypteris is a small genus consisting of four species native to America and Madagascar: Trachypteris induta, T.?pinnata, T.?gilliana, and T.?drakeana. The spores of the genus were studied using light microscopy and scanning and transmission electron microscopy. They are trilete, triangular to globose, with equatorial diameter of 29?C58???m and polar diameter of 26?C53???m. The exospore is 0.5?C1.1???m thick, plane, and two-layered. The perispore is 0.3?C6.8???m thick and two-layered. Depending on the species, two different ornamentations were observed: cristated with partially fused cristae, and ridged with partially fused ridges, forming an incomplete reticulum. Globules immersed in the perispore were occasionally observed. The systematic value of the Trachypteris spores is discussed, and their morphology and ultrastructure are compared with those of other related cheilanthoid ferns. The palynological characteristics presented here may be useful for phylogenetic studies within the Pteridaceae, and particularly within the cheilanthoid ferns.     

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。     

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。     

Spore morphology of pteridophytes from China Ⅱ. Sinopteridaceae

Spores of 61 species and 6 varieties in 9 genera of the Sinopteridaceae were examined under scanning electron microscope (SEM). Based on surface ornamentation and other features, the spores of the Sinopteridaceae are divided into three types. In type Ⅰ , the exospore is smooth and the surface ornamentation, which is reticulate, cristate, echinate or rugate, is formed by the perispore. All the other genera of this family, except for Onychium and Cryptogramma, have this pattern of spores. In type Ⅱ, the surface ornamentation is formed by both perispore and exospore. This pattern is found only in Cryptogramma. In type Ⅲ, the perispore is thin and the surface ornamentation is formed by the exospore. Onychium is characterized by this type of spores. Those genera with spores of type Ⅰ of the Sinopteridaceae seem to be closely related to each other and should be natural members of this family. The systematic position of Cryptogramma and Onychium, with spores of type Ⅱ and type Ⅲ respectively, however, should be reconsidered. Aleuritopteris might be the mostprimitive member of the Sinopteridaceae from the evidence of spore morph     

Spore Morphology of Pteridophytes from China Ⅵ .Pteridaceae

The spore morphology of 30 species 4 variety of 2 genera of Pteridaceae from China was investigated under scanning electron microscope (SEM) . Among them, 29 species 4 variety belong to the genus Pteris . The spores of Pteris are trilete , radiosymmetrical , subtriangular in polar view and hemispherical or subhemispherical in equatorial view . The polar axes are 26 - 54μm long , and equatorial axes are 34 - 97μm long . The perispore is thin and the surface ornamentation is formed by the exospore . The types of ornamentation are tuberculiform-rugulate , verruciform-rugulate or lophate , the spore of most species with the equatoial flange, some species with proximal ridge and distal ridge . The spore morphology of Pteris is stable, and the difference between species is distinct , but the features of spore and sporophyte are not related. The spore of Histiopteris is monolete and bilaterally symmetrical, elliptical in polar view and kidney-shaped in equatorial view . The polar axes are 22 - 23μm long , and equatorial axes are 29 - 36 μm long . The perispore is thin and the surface ornamentation is formed by the exospore . The surface is rugulate . The spore morphology of Histiopteris and Pteris is very differential , put genus Histiopteris in family Pteridaceae is not suitable, according to the feature of spore morphology .     

中国蕨类植物孢子形态的研究 Ⅵ . 凤尾蕨科

利用扫描电镜对国产凤尾蕨科( Pteridaceae ) 2 属30 种4 变种植物孢子进行了观察。其中凤尾蕨属( Pteris) 29 种4 变种, 孢子三裂缝, 辐射对称; 极面观为钝三角形, 赤道面观为半圆形或超半圆形; 孢子极轴长26～54μm, 赤道轴长34～97μm; 外壁厚, 形成孢子表面纹饰的轮廓; 周壁薄, 紧贴于外壁上; 多数种类孢子外壁具沿赤道加厚的赤道环, 孢子表面呈瘤块状、疣块状, 或隆起的脊连成网状或拟网状等纹饰, 有的种类具近极脊和远极脊。凤尾蕨属植物孢子的形态稳定, 种间差异明显, 但孢子形态特征与孢子体的形态特征是不相关的。栗蕨属( Histiopteris) 植物孢子为单裂缝, 两侧对称, 极面观椭圆形, 赤道面观豆形; 孢子极轴长22～23μm, 赤道轴长29～36μm; 周壁很薄, 由外壁形成孢子纹饰的轮廓; 表面呈粗的块状纹饰。从孢子形态上看, 栗蕨属与凤尾蕨属有很大差异, 将其放在一个科中不合适。     

中国蹄盖蕨属植物孢子形态的研究

利用扫描电子显微镜对我国产蹄盖蕨属44种植物的孢子进行了观察。结果表明,该属孢子形态为单裂缝,两侧对称,极面观为椭圆形,赤道面观为豆形。外壁表面光滑,由周壁形成表面纹装饰。根据周壁的结构和表面纹饰,可分为两种类型;一是周壁外层发达,形成粗大的脊状纹饰,有11种属此类型;二是周壁外层很薄或不完全发育,由周壁内层或中层形成表面纹饰,有33种属此类型纹饰。本文还就本属的孢子形态特征以及与本属的属下分类关系、本属与邻近属的关系等进行了讨论。