A TETRAHEDRAL MEGASPORE ARRANGEMENT IN A SEED FERN OVULE OF PENNSYLVANIAN AGE

A tetrahedral arrangement of one functional and three aborted megaspores has been found in the fossil seed fern ovule Conostoma anglo-germanicum. This is the first conclusive report of a Pennsylvanian age ovule with such a primitive megaspore arrangement, suggesting that this feature was more widespread and persistent among Paleozoic pteridosperms than previously assumed.     

耳叶补血草大、小孢子发生与雌、雄配子体的发育

本文较系统地报道了耳叶补血草(Limonium otolepis(Schrenk.)Kuntze.)的大、小孢子发生和雌、雄配子体形成过程.主要结果:(1)小孢子母细胞减数分裂过程中的胞质分裂为同时型,小孢子四分体多为正四面体形,少数左右对称形;(2)成熟的花粉为三细胞型,具3个萌发孔;(3)花药壁由5层细胞组成,最外层为表皮,其内分别为药室内壁、中层、绒毡层,绒毡层为变形型,花药壁的发育属于基本型;(4)耳叶补血草的雌蕊由5心皮组成,柱头、花柱分离,子房合生,1室,基生胎座,胚珠1个,拳卷型,双珠被,厚珠心,珠孔由内珠被形成;(5)孢原细胞发生于珠心表皮下,经1次平周分裂,形成造孢细胞,由造孢细胞直接发育成大孢子母细胞,大孢子母细胞减数分裂形成4个大孢子呈直线排列,合点端大孢子具功能,属于典型的蓼型胚囊发育.     

Developmental Mechanisms in Heterospory: Features of Post-meiotic Regression in Selaginella

Post-meiotic regression in the sporangia of Selaginella sulcataresults in tetrads comprising small and large spores. From germinationtests it has been found that the small spores are abortive andthe large ones fertile, a circumstance which suggests an innateconstitutional difference between them. Densitomctric measurementsshow that the cytoplasm in the prophase megasporocyte is polarized.At cytokinesis this cytoplasm is simultaneously partitionedinto four cells, equal in size, but presumably not in composition.Initially, all four megaspores in the tetrad grow at the samerate, but later, development becomes differential with one,or more usually two spores increasing rapidly in size and theothers much more slowly. The onset of this phase is signalledby a difference in protoplasmic staining and it is the megasporeswith a densely-staining protoplast which develop to fertility. A comparable process of regression occurs also in the microsporangium;the spores develop differentially after a period of equal growth.Examination of young microspore tetrads shows that daughterspores differ in the density of free cytoplasmic ribosomes.It is suggested that this is a consequence of cither disproportionateinheritance of ribosomes from the parental cell resulting frompolarization or, more probably, since the deficiency is notre-dressed in later development, an inherited difference inthe ability to effect ribosome synthesis after prophase elimination.Endoplasmic reticulum-ribosome complexes, in other organismsbelieved to be implicated in cell degeneration, o0ccur in themicrospore cytoplasm.     

INVESTIGATIONS OF NORTH AMERICAN CYCADEOIDS: EARLY OVULE ONTOGENY

Ovules on cones of the genus Cycadeoidea from the Cretaceous were found preserved at the pregametophytic stage of nucellar development. At this stage, the ovules are distinct from those of extant gymnosperms in that there is a greater degree of differentiation in both the integuments and the nucellus in Cycadeoidea. Linear clusters of three or four cells, which consistently occur in the middle of the nucellus, are interpreted to be the remains of linear tetrads of megaspores. This example of exceptional preservation allows, for the first time, a comparison of early ontogenetic events in a fossil seed with those of extant seeds.     

Studies of Fossil and Modern Spore Wall Biomacromolecules using13C Solid State NMR

A range of Carboniferous lycophyte megaspore exines have beeninvestigated using¹³C magic-angle spinning nuclear magneticresonance (MAS NMR) spectroscopy. Their composition differsconsiderably from sporopollenin obtained from an extant lycophyte.The differences observed result in part from varying degreesof diagenesis. Fossil fern spores, gymnosperm megaspore-membranes and pollenhave also been examined. These show a similar composition tothe fossil lycophyte megaspores. The constituent material ofall of these exines differs considerably from the sporopolleninobtained from comparable extant samples. Despite the changesin composition observed on fossilisation, differences in compositionbetween the major groups of plants may be preserved to someextent in the fossil material. Walls of the fossil prasinophyceanalgal cystTasmanites have been examined and these show a greatersimilarity to fossil cuticle and algaenans than to sporopollenins. The effect of oxidative maceration on fossil and modern sporopolleninshas also been investigated. The main influence of oxidativemaceration is the removal of unsaturated carbon environmentssuch as aromatics; this causes fossil spores to be more susceptibleto oxidative maceration than the modern exines. Heating of modernexine material models the alteration of exines by diagenesis.The changes that occur on heating an extant sample to 150–225°Cgive a chemical composition that is similar to those of thefossil sporopollenins. ¹³C solid state NMR; spores; pollen; fossil; Carboniferous lycopsids; ferns; pteridosperm; gymnosperm; oxidative maceration; heating; thermal maturation     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: A MIMOSOID INFLORESCENCE

Six examples of a spicate inflorescence from the Middle Eocene Claiborne Formation in western Tennessee have been investigated. Individual flowers are small, alternately arranged, nearly sessile, and perfect. The style protrudes 2 mm beyond the floral envelope and terminates in a slightly swollen, rounded stigma. Both the style and the stigma are hairy. Ten stamens are exserted and extend 5 mm beyond the floral envelope. Anthers are small, versatile, and dehisce longitudinally. Pollen grains are tricolporate, tectate, and are in permanent tetrahedral tetrads 32 μm in diameter. Comparison of the fossil inflorescences with those of extant families having multiple pollen grain configurations suggests that the fossil inflorescences are most closely allied to the subfamily Mimosoideae of the Leguminosae. These are the first structurally documented remains of the Mimosoideae from the Middle Eocene.     

Ordered Linear Tetrads Are Produced by the Sporulation of Newly Formed Zygotes of Saccharomyces cerevisiae

Diploid Saccharomyces cerevisiae strains normally sporulate to produce tetrahedral unordered asci containing four spores (tetrads). We report that when newly formed zygotes are subjected to the same sporulation conditions, they form predominantly linear ordered tetrads. We show that the two spores from each end of such a linear tetrad invariably contain nonsister centromeres. Spore viability, recombination and independence of centromere segregation appear unaffected.     

ULTRASTRUCTURAL STUDIES OF IN SITU DEVONIAN SPORES: BARINOPHYTON CITRULLIFORME

Each sporangium in the Upper Devonian taxon Barinophyton citrulliforme contains both microspores and megaspores. Microspores range up to 50 μm in diam and possess a homogeneous sporoderm characterized by an outer separable layer. The sporoderm of the megaspores (up to 900 μm) is constructed of sporopollenin units that are loosely arranged in the outer portion of the wall, and that give the megaspore wall a spongy organization. Ultrastructural evidence suggests that the small spores were not abortive megaspores, but that both spore types were functional. The spores of this plant, as well as other Devonian spores that show less dramatic size differences, are suggested as demonstrating a phase in the evolution of heterospory where sex determination was established in spores within the same sporangium prior to the evolution of micro- and megasporangia.     

三叶木通大小孢子发生和雌雄配子体发育研究

三叶木通（Akebia trifoliata）属于木通科（Lardizabalaceae）木通属（Akebia）,为雌雄异花,雌雄同株的木质藤本植物,具有药食两用的经济价值。本文运用石蜡切片技术观察了三叶木通的大小孢子发生和雌雄配子体发育过程中胚胎学特征,以期了解三叶木通有性生殖过程,分析该物种自然条件下结实率低的生殖原因,为其后续进行杂交育种和新品种培育提供理论基础。结果表明：三叶木通雄蕊6枚,每枚花药具4个小孢子囊,花药壁完全分化时由外到内依次为1层表皮、1层药室内壁、2～3层中层及1层绒毡层,且绒毡层为分泌型绒毡层。小孢子母细胞减数分裂的胞质分裂为同时型,小孢子四分体为四面体型排列,从四分体分离出来的小孢子经过进一步发育形成2-核成熟花粉细胞。小孢子囊中的小孢子四分体时期存在少量败育现象。雌蕊具1室子房,为侧膜胎座,胚珠横生,多枚,双珠被,厚珠心。大孢子母细胞减数分裂后形成的大孢子四分体呈线型排列,靠珠孔端的3个大孢子退化,合点端的1个大孢子发育成为功能大孢子,经3次连续有丝分裂并进一步发育为七细胞八核的蓼型胚囊。雌花中的雄蕊早期发育正常,但小孢子发育至单核靠边期不在进一步发育,...     

ACHLAMYDOCARPON VARIUS COMB. NOV.: MORPHOLOGY AND REPRODUCTIVE BIOLOGY

Structurally preserved arborescent lycopsid fructifications are described from Pennsylvanian age strata in eastern Kentucky and southern Illinois. Achlamydocarpon varius comb. nov. is the name proposed for these cones and previously reported isolated megasporophylls described as Lepidostrobophyllum varius. The specimens range up to 3.5 cm long and represent cone apices. Megasporophylls are spirally arranged and attached to the cone axis at an acute angle. Megasporangia are large with a wall two cell layers thick. Each sporangium contains one large, presumably functional megaspore, and three smaller, abortive megaspores. Functional and abortive spores possess a prominent apical tuft (massa) that covers up to one-third of the proximal surface of the spore. Sporoderm ultrastructure is detailed together with a comparison of morphologically similar sporae dispersae megaspores. The possible function of the megaspore massa is discussed as it relates to the reproductive biology of the cone.     

珍稀濒危植物巴东木莲胚胎学研究

对巴东木莲(Manglietia patungensis Hu)的花发育以及胚胎发育过程进行了系统研究。巴东木莲花顶生,花器官头年年底开始分化到第二年3月分化出花被、雌雄蕊群直至6月发育成熟。雌蕊成熟时胚珠倒生,双珠被,厚珠心,大孢子四分体线形排列,合点端发育成功能大孢子,珠孔端的3个退化,大孢子为单孢子发生型,胚囊发育方式属蓼型;雄蕊花药外侧壁玫瑰红色,内侧有4个白色花粉囊,绒毡层有1层多核细胞,小孢子四分体排列方式多为左右对称形和交叉形,四面体形,偶为T字形和线形,成熟花粉粒为二细胞型。在巴东木莲花发育和大、小孢子发生以及雌、雄配子体形成过程中未见异常现象,因此笔者认为该物种的花器官发育以及雌、雄配子体发育并不构成导致该物种濒危的因素。     

SPORE TETRAD STRUCTURES OF POSSIBLE HEPATIC AFFINITY FROM THE AUSTRALIAN LOWER CARBONIFEROUS

Palynological studies of Early Carboniferous (Viséan) sediments of the Bonaparte Gulf Basin, northwestern Australia, reveal the presence of intact tetrahedral spore tetrads (here described as Sagenotetradites gen. nov.) and, more usually, of their disjunct spore portions which had previously been interpreted as dispersed whole-miospore species. Two species of the new tetrad genus are recognized: S. bonapartensis (Playford) comb, nov., as type species; and S. spiritensis (Playford) comb. nov. The species are distinguished, as originally, on the basis of sculptural attributes of their distal exoexines. When intact, both species share a common internal “binding” element (“Cadiospora abrupta” Playford, 1971) composed of the fused proximal exoexinal faces of all four spores of a given tetrad. Morphological comparisons with modern hepatic spores suggest an alliance of the microfossils with the order Sphaerocarpales. The occurrence of Sagenotetradites in exclusively marine sediments suggests that its parent plants grew in close proximity to the marine depositional basin. Moreover, the morphological attributes of the tetrads would appear to have facilitated dissemination by water.     

鹅毛竹大小孢子及雌雄配子体发育

利用扫描电镜、透射电镜、石蜡切片,对鹅毛竹的花芽分化、大、小孢子及雌、雄配子体的发育进行了详细观察.结果发现:鹅毛竹花药具4个药室,花药壁由表皮、药室内壁、中层、绒毡层4层结构组成,花药壁发育为单子叶型,绒毡层为腺质型,小孢子母细胞减数分裂中的胞质分裂为连续型,产生左右对称型小孢子.鹅毛竹成熟花粉大多2细胞型,都具1个萌发孔.鹅毛竹子房为单子房,子房1室,侧膜胎座,一个倒生胚珠,双珠被,薄珠心.大孢子母细胞由一个雌性孢原细胞直接发育而成,大孢子四分体呈线型,合点端一个大孢子分化为功能大孢子,由功能大孢子经过3次有丝分裂形成8核胚囊,发育类型为蓼型,位于核点端的3个细胞核进行多次分裂形成多个反足细胞.至此,成熟胚囊形成.并就鹅毛竹不结实的原因进行了探讨.     

Meiotic cytokinetic apparatus in the formation of the linear spore tetrads of Conocephalum japonicum (Bryophyta)

Most bryophytes produce tetrahedral spore tetrads. However, linear spore tetrads have been reported to occur in Conocephalum japonicum (Thunb.) Grolle. In this study, the distribution of microtubules (MTs) during meiosis in C. japonicum was examined to determine the division pattern resulting in a linear tetrad. Spore mother cells in the pre-meiotic stage were cylindrical with randomly distributed cytoplasmic MTs. In the prophase-metaphase transition, spindle MTs replaced cytoplasmic MTs and a barrel-shaped spindle with two flattened poles developed. Cortical MT arrays were not detectable throughout meiosis. Although a phragmoplast appeared between sister nuclei in telophase-I, it disappeared without expanding to the parental cell wall. Metaphase-II spindles oriented parallel to the long axis of the cell and in tandem to each other resulted in a linear arrangement of telophase nuclei. Radial arrays of MTs developed from the nuclear surfaces and three phragmoplasts appeared among the four nuclei to produce four spores. Two phragmoplasts separating the paired sister nuclei appeared prior to the appearance of a phragmoplast between non-sister nuclei. The MT cycle is basically the same as that reported in meiosis of C. conicum, which produces non-linear tetrads. A morphometric study indicated that the difference in the division pattern between C. conicum and C. japonicum is due to a difference in the shape of spore mother cells. The cylindrical shape of sporocytes of C. japonicum restricts the orientation of spindles and phragmoplasts so that the four resultant spores are arranged linearly. Received: 22 April 1998 / Accepted: 15 May 1998     

On Eosperma oxroadense gen. et sp. nov.: a New Lower Carboniferous Seed from East Lothian

A new petrified platyspermic seed is described from the CementstoneGroup (Tournaisian) of Scotland. It has a generally Lagenostomaleanappearance despite its unusual symmetry, and it is comparedwith some other members of that order. It possesses a well-preservedmegaspore membrane, one of the most notable features of whichis the polygonal markings on the outer surface. Comparison ismade with some megaspores from fossil and living seeds. Thepossible affinity of the seed is discussed.     

On the significance of tetrahedral tetrads of Precambrian algal cells

Tetrahedral tetrads of coccoid cells are known from two Precambrian localities; one example has been reported from the late Proterozoic Bitter Springs Formation and a second example, morphologically different from the first, is here described from the middle Proterozoic Amelia Dolomite. Among modern algae, sheath- or wall-enclosed tetrahedral tetrads are regularly produced only by eukaryotes, either during meiotic or mitotic reproduction. In evaluating the significance of Precambrian tetrads, consideration should be given to the following: whether the tetrad geometry is truly tetrahedral; if so, whether the tetrahedral configuration is original or artifactual; and if original, whether the tetrad was produced during meiotic or mitotic reproduction. We suggest several kinds of evidence which may assist in making these determinations. Considering the Amelia Dolomite specimen, we conclude that it most closely resembles a mitotically produced green algal autospore tetrad, although this interpretation cannot yet be confirmed because of the difficulty in ascertaining whether the tetrahedral configuration is original. Considering the Bitter Springs specimen, we conclude that the organism probably was eukaryotic but that its mode of origin (whether through meiosis or mitosis) cannot be unequivocally ascertained from data presently available.     

Electron Microscopy of the Megaspore Horstisporites Semireticulatus from Liassic Strata of Germany

Ultra-thin sections of paratypes of the megaspore Horstisporites semireticulatus Jung from Liassic strata of Germany have been investigated by the transmission electron microscope. By this it could be demonstrated that the fossil sporoderm consists of two distinct layers. The inner layer (exine) is very thin (about 0.5 μ) and reveals a lamellated structure. The outer layer (perine) is thicker by far (about 25 μ or more) and is composed of ramifying sporonin threads, which form a three-dimensional network. Proximally, in the region of the triradiate dehiscence commissure, both layers coalesce. Distally the exine separates from the perine, forming a cavity hitherto, erroneously, called mesosporoid. The structural similarity of the Horstisporites semireticulatus sporoderm and that of such megaspores of Selaginella which show a monozonal kind of perine formation e.g. Selaginella selaginoides, favours the idea that the fossil species in question belongs to the Selaginellaceae     

Silicon in sporoderms of micro- and megaspores of Isoetes echinospora Durieu registered by EDS and EELS

Protoplasma - The present study reveals silica in sporoderms of micro- and megaspores of the modern quillwort Isoetes echinospora Durieu and homologizes layers of the sporoderm in spores of this...     

Development of polarity in the ovules of the F2-progeny of theOenothera hybridalbicans · hhookeri

In the F₂-progeny of hybrids from crosses betweenOenothera biennis orsuaveolens andOe. hookeri with theRenner-complexesalbicans and^hhookeri, the development of callose pattern in meiocytes and megaspore tetrads is the same as in the F₁ and the parentOe. hookeri. During the development of the megaspore tetrads and the embryo sacs primary and secondary heteropolarity as well as homopolarity is observed. Estimates for the initial frequency of homo- and heteropolar tetrads at the end of the degeneration of megaspores in the tetrads immediately before the start of embryo sac development could be calculated. The F₂-plants can be arranged in three groups, distinguished by the frequency of the two polarity types. One of these groups behaves similar to the parentOe. hookeri, the two others have more homopolar tetrads. The segregation can be interpreted as recombination of genes, which influence the development of the polarity in the ovules. This is possible by crossing-over of genes between the twoRenner-complexes of the hybrid.     

Lageniastrum macrosporae (fossil Volvocales, Lageniastraceae nov. fam.), an endophyte in megaspores from the Carboniferous of the French Massif Central

Sublagenicula nuda lycophyte megaspores from the upper Visean of central France frequently host a colonial green alga as an endophyte. This association was first recorded more than 100 years ago by the French paleobotanist B. Renault, who introduced the name Lageniastrum macrosporae for the alga. However, the biological significance of the discovery was not fully assessed until recently. The L. macrosporae-S. nuda association represents the oldest compelling fossil evidence for algal endophytes in land plants, and the only example to date of an alga residing in the interior of spores of vascular cryptogams. Here we present a detailed re-evaluation and photographic documentation of the surviving original specimens from the Visean of Combres/Lay and Esnost. Moreover, a newly discovered specimen from the Stephanian of central France represents the first record of this association from the Upper Carboniferous. An emended diagnosis for L. macrosporae Renault, 1896 is provided, and a lectotype and paralectotype are designated. Although L. macrosporae displays a striking suite of morphological characters found in members of the extant chlorophyte family Volvocaceae (especially the genus Volvox), the peculiar biology of the fossil necessitates establishment of a new family, for which the name Lageniastraceae is proposed. Considerations of the adaptive advantages for the alga of occupying the interior of megaspores are offered. These include the possible effectiveness of the spores in protection during periods of desiccation and against plankton-feeding animals, and use of the spores as potential vectors for dispersal from one isolated body of water to another by spore-feeding animals.