Two cryptospore-bearing land plants from the Lower Devonian (Lochkovian) of the Welsh Borderland

Two examples of fragmentary, coalified plant fossils with cellular preservation and in situ spores are described using scanning and transmission electron microscopy (SEM and TEM), from a Lower Devonian (Lochkovian) locality in the Welsh Borderland. A sporophyte which produced dyads of ?Cymbohilates of. horridus is unique in that stomata are numerous on the supporting axis, and are comparable with those described from contemporary vascular plant remains. The in situ dyads possess a bilayered exospore wall, with an outer exosporal envelope present over the distal faces. A fusiform sporangium, with externally smooth epidermis, contains specimens of the tetrad Velatitetras sp. Each tetrad is encompassed by a laevigate, folded, exosporal envelope of uniform thickness, which contains a layer of regular voids. Spores within the tetrads are ultrastructurally bilayered, with a complex, digitate outer margin presumably representing spore wall ornamentation. Neither the tetrads nor dyads reported in this paper are lamellate in ultrastructure. The combination of stomata, branching and dyads in the same sporophyte holds significance for the understanding of cryptospore affinities.     

Sporangia containing Scylaspora from the Lower Devonian of the Welsh Borderland

ABSTRACT. Bulk maceration of Early Devonian (Lochkovian) deposits from the Welsh Borderland has yielded eight specimens of a new type of sporangium characterized by its elongate shape and distinctive spores. The specimens have been examined using scanning electron, transmission electron and light microscopy. The elongate sporangia occur isolated and are fragmented to varying degrees. They contain trilete spores that possess a proximal surface with shallow murornate ornament and a distal surface that is laevigate. The spores belong to the dispersed spore genus Scylaspora , and this is the first report of such spores in situ . Ultrastructural studies demonstrate that the spore walls are bilayered with a lamellate inner layer overlain by an essentially homogeneous outer layer. There is little or no associated extra-exosporal material. To date this is the earliest reported example of lamellate wall ultrastructure in trilete spores. Models of spore wall development are suggested in the light of evidence provided by spore wall ultrastructure. Detailed comparisons of the characters preserved in the fossils (morphological, anatomical and ultrastructural), with those in other fossil and extant plants, currently shed little light on the phylogenetic relationships of these specimens, primarily due to the paucity of comparable data. It is proposed that the plant is probably of vascular status, but in the absence of evidence for vascular tissue, it must be classified as rhyniophytoid.     

Ultrastructure of laevigate hilate spores in sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland

Spore masses and isolated sporangia, containing laevigate hilate cryptospores attributable to the dispersed taxon Laevolancis divellomedia sensu lato, have been recovered on bulk maceration of Upper Silurian (Pridoli) and Lower Devonian (Lochkovian) deposits from the Welsh Borderland. Detailed morphological, anatomical and ultrastructural analysis, using light microscope, scanning electron microscope and transmission electron microscope techniques, reveals subtle differences between the specimens and they can be grouped into five distinct types. The different groups are distinguished principally by using sporangia-spore mass characteristics, presence or absence of extra-exosporal material and nature of spore-wall ultrastructure. Of the groups, one has a uniformly homogeneous exospore and the other four groups have a bilayered exospore. In the former the spores lack extra-exosporal material and occur in a discoidal sporangium. Of the bilayered groups, two have exospores of homogeneous composition but with the two layers differing in electron density. They occur in discoidal sporangia and spore masses and are distinguished on the presence or absence of extra-exosporal material and differences in the widths of the two layers. Finally, two bilayered groups possess a lamellate inner layer, but vary in presumed sporangial shape. Elongate sporangia have spores with concentric continuous lamellae, lacking further ultrastructure. In contrast, spores from a discoidal spore mass have white-line-centred, presumably tripartite, lamellae which are laterally discontinuous, overlapping and irregularly spaced. These findings, which suggest that morphologically similar spores were produced by a number of plant taxa, have important implications regarding the assessment of early land-plant diversity. The affinities of hilate cryptospore-producing plants are unknown and problematic, particularly as no extant non-angiosperm plants produce dyads, other than through meiotic irregularity, and spore-sporangial characters have no exact counterpart in coeval plants. Studies of specimens with in situ hilate cryptospores suggest that they derive from rhyniophytoids, i.e. plants that resemble the simplest of vascular plants but lack evidence of vascular tissue, although hilate cryptospore-containing examples show no axial branching. It might be argued, based on evidence from spore wall ultrastructure, that some of the plants have more in common with lycopsids and filicopsids than bryophytes, a surprising finding bearing in mind the stratigraphic distribution of hilate cryptospores-dyads and inferences that the producers were bryophyte-like. Detailed studies of wall structure in the hilate cryptospores permit consideration of spore wall development. It is suggested that extra-exosporal material derives from a tapetum and is thus produced by the diploid sporophyte. The white-line-centred lamellae in a single specimen provide the earliest evidence for the presence of such structures in early land plant spores and provide further evidence that sporopollenin deposition on such structures is the most primitive mode of sporopollenin deposition among land plants.     

The role of mid-palaeozoic mesofossils in the detection of early bryophytes

Recently discovered Silurian and Devonian coalified mesofossils provide an additional source of data on early embryophytes. Those reviewed in this paper are considered of some relevance to understanding the early history of bryophytes while highlighting the difficulties of recognizing bryophytes in often very fragmentary fossils. The first group comprises sporophytes in which terminal sporangia contain permanent dyads and tetrads. Such spores (cryptospores) are similar to those found dispersed in older Ordovician and Silurian strata, when they are considered evidence for a land vegetation of embryophytes at a bryophyte grade. The phylogenetic significance of plants, where the axes associated with both dyad- and tetrad-containing sporangia are branching, a character state not found in extant bryophytes, is discussed. The second group comprises axial fossils, many with occasional stomata, in which central conducting strands include G-type tracheids and a number of novel types of elongate elements not readily compared with those of any tracheophyte. They include smooth-walled, evenly thickened elongate elements as well as those with numerous branching +/- anastomosing projections into the lumen. Some of the latter bear an additional microporate layer, but the homogenized lateral walls between adjacent cells are never perforate. Such cells, which occur in various combinations in central strands, are compared with the leptoids and hydroids of mosses, hydroids of liverworts and presumed water-conducting cells in coeval Lower Devonian plants such as Aglaophyton. It is concluded that lack of information on the chemistry of their walls hampers sensible assessment of their functions and the affinities of the plants. Finally, a minute fossil, comprising an elongate sporangium in which a central cylindrical cavity containing spores and possible elaters terminates in a complex poral dehiscence apparatus, is used to exemplify problems of identifying early bryophytes. It is concluded that further progress necessitates the discovery of pre-Upper Silurian fossils with well-preserved anatomy, as well as a re-evaluation of criteria used to assess existing and new Devonian fossils for bryophyte affinity.     

New insights into early land ecosystems: a glimpse of a lilliputian world

Small, relatively uncompressed, very fragmentary plant remains (mesofossils) are described from a Silurian (P ídolí) and a Lower Devonian (Lochkovian) locality in the Welsh Borderland. Excellent cellular preservation provides characters leading to the demonstration of diversity in plants of simple gross morphology and allows deliberations on functional anatomy (e.g. of stomata), and reproductive biology (including development and dehiscence of sporangia). A survey of in-situ spores is presented, and preliminary comparisons made with dispersed spore assemblages especially in relation to reconstruction of vegetation on local and regional scales. The earliest body fossils of unequivocal terrestrial arthropods isolated from the same locality as the P ídolí plants suggest that the decomposer/microherbivore/predator soil and litter communities found in the Lower and Middle Devonian extend back at least into the Silurian. Evidence for plant—animal interaction in the Lower Devonian comes from spore-dominated coprolites believed to have been produced by litter-feeding myriapods.     

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。     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age (micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.     

Lower Devonian coalified sporangia from Shropshire: Salopella Edwards & Richardson and Tortilicaulis Edwards

A diverse assemblage of coalified sporangia from Lochkovian/Gedinnian, fluvial rocks imicrornatus-newportensis Spore Biozone) contains fusiform forms assignable to Salopella Edwards & Richardson and Tortilicaulis Edwards. Both show bifurcation within some sporangia. In specimens assigned to Salopella cf. marcensis , longitudinal dehiscence produces two equal valves revealing spores resembling Aneurospora , although trilete marks may be difficult to distinguish. The spores are in two forms with distal ornament of either coarse or fine coni, each occurring in separate sporangia. A further sporangium of Salopella shape contains distally apiculate, hilate cryptospores. Elongate sporangia exhibiting spiralling of superficial cells and sometimes gross twisting, are placed in a new species of Tortilicaulii, T. offaeus. Anatomical data include details of irregularly thickened epidermal cells, sporangial wall and in situ spores. The latter are trilete, equatorially thickened and highly distinctive, because the entire exospore surface is covered with grana, a feature not common in dispersed spores of this age. The nomenclatural and curatorial problems associated with these small, coalified fossils, whose anatomical examination by scanning electron microscope necessitates elimination of gross morphology, are discussed.     

Permanent dyads in sporangia and spore masses from the Lower Devonian of the Welsh Borderland

Dyads, interpreted as cryptospore permanent dyads, are reported from sporangia and spore masses recovered from fluviatile deposits of Lochkovian (Early Devonian) age from the Welsh Borderland. The morphology, anatomy and ultrastructure of the specimens have been analysed using LM, SEM and TEM. Two specimens consist of a single sporangium terminating an axis: one is cup-shaped and attached to an unbranched axis and named Culullitheca richardsonii gen. et sp. nov., the other comprises an isotomously branching axis with one sporangium preserved and is named Fusitheca fanningiae gen. et sp. nov. Two specimens show dyads adhering to cuticular fragments presumably derived from sporangia. In one the cuticle is of irregular shape, in the other it is elliptical. The final specimen comprises an elongate spore mass. In all cases ultrastructural analysis reveals that the spore walls are essentially homogeneous. Our findings suggest that cryptospore permanent dyads were produced by rhyniophytoid plants of small stature. The affinities of these plants remain conjectural, as is the relationship of the cryptospore permanent dyads with other sporomorph morphotypes (cryptospores and trilete spores). However, the production of dyads by a plant with a bifurcating sporophytic axis is evidence against affinity with extant bryophytes. The mode of formation, adaptive significance, affinities, phylogenetic relationships and stratigraphical history (including demise) of cryptospore permanent dyads is discussed.     

A new group of Early Devonian plants with valvate sporangia containing sculptured permanent dyads

As part of a study to explore diversity and disparity in Early Devonian terrestrial vegetation, several hundreds of sporangia with in situ spores have been isolated from a Lochkovian locality in Shropshire. These include a small number (seven) of sporangia showing dehiscence into four valves and containing permanent sculptured dyads, belonging to the Cymbohilates horridus complex and C. cymosus, which are recorded in coeval dispersed spore assemblages. A further, previously described, mesofossil comprises an incomplete sporangium containing C. horridus that terminates a naked isotomously branching stem with stomata. The valvate sporangia are placed in a new genus, Partitatheca, containing four species, P. splendida (type), P. horrida, P. densa and P. cymosa, their names reflecting the names of the dispersed spore species and varieties. Complex ultrastructure in the walls of the dyads is similar to that in earlier dyads in the Dyadospora complex where it provides evidence for a hepatic affinity of the earliest embryophytes, but the new taxa present a combination of bryophyte and tracheophyte characters and are considered to represent a new embryophyte lineage. General discussion includes the development of dyads, more particularly their relevance to understanding the diversity in meiotic processes, and the disappearance of dyads from the dispersed spore record prior to the Middle Devonian. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 229–257.     

Implications of an early land plant spore assemblage for the late Silurian age of the Si Ka Formation,northern Vietnam

The first plant microfossil assemblage from the Si Ka Formation of the Song Cau Group, northern Vietnam is reported. It is composed of cryptospores in dyads and tetrads, trilete spores, tubular remains consisting of an association of smooth, banded, and externally thickened tubes, and cuticle-like fragments. The biostratigraphic assemblage of sporomorphs indicates a late Silurian (late Ludfordian) to Early Devonian (early Lochkovian) age. Further comparison with coeval reports using the characteristic features of the assemblage confines their age to the late Ludlow (late Ludfordian) to early Přídolí. This report presents the oldest spore assemblage from Vietnam and contributes to a broader understanding of its paleo-landscape during the late Silurian.     

Exceptional preservation in Lower Devonian coalified fossils from the Welsh Borderland: a new genus based on reniform sporangia lacking thickened borders

Reniform sporangia, comprising two equal valves and containing retusoid spores, recovered from Lower Old Red Sandstone strata of Devonian age ( micrornatus-newportensis Spore Biozone: lower Gedinnian lower Lochkovian) on North Brown Clee Hill in the Welsh Borderland are placed in Resilitheca salopensis gen. et sp. nov. Conventional compression fossils from Targrove, Ludlow of fertile axes showing isotomous branching with limited overtopping are considered conspecific because the terminal reniform sporangia contain the same spores. Spore ultrastructure is described using scanning and transmission electron microscopy. Sections show faint traces of lamellae. Particles associated with spores and sporangium wall are compared with the globules of pteridophytes and Ubisch bodies of angiosperms, and related to the development of the sporangium. The new plants are compared with Cooksonia caledonica Edwards known only from impressions, and with Renalia Gensel showing far more pronounced pseudomonopodial branching.     

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.     

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。     

ULTRASTRUCTURE OF DISPERSED AND IN SITU SPECIMENS OF THE DEVONIAN SPORE RHABDOSPORITES LANGII: EVIDENCE FOR THE EVOLUTIONARY RELATIONSHIPS OF PROGYMNOSPERMS

Abstract: The spore Rhabdosporites (Triletes) langii (Eisenack) Richardson, 1960 is abundant and well preserved in Middle Devonian (Eifelian) ‘Middle Old Red Sandstone’ deposits from the Orcadian Basin, Scotland. Here it occurs as dispersed individual spores and in situ in isolated sporangia. This paper reports on a detailed light microscope (LM), scanning electron microscope (SEM) and transmission electron microscope (TEM) analysis of both dispersed and in situ spores. The dispersed spores are pseudosaccate with a thick walled inner body enclosed within an outer layer that was originally attached only over the proximal face. The inner body has lamellate/laminate ultrastructure consisting of fine lamellae that are continuous around the spore and parallel stacked. Towards the outer part of the inner body these group to form thicker laminate structures that are also continuous and parallel stacked. The outer layer has spongy ultrastructure. In situ spores preserved in the isolated sporangia are identical to the dispersed forms in terms of morphology, gross structure and wall ultrastructure. The sporangium wall is two‐layered. A thick coalified outer layer is cellular and represents the main sporangium wall. This layer is readily lost if oxidation is applied during processing. A thin inner layer is interpreted as a peritapetal membrane. This layer survives oxidation as a tightly adherent membranous covering of the spore mass. Ultrastructurally it consists of three layers, with the innermost layer composed of material similar to that comprising the outer layer of the spores. Based on the new LM, SEM and TEM information, consideration is given to spore wall formation. The inner body of the spores is interpreted as developing by centripetal accumulation of lamellae at the plasma membrane. The outer layer is interpreted as forming by accretion of sporopollenin units derived from a tapetum. The inner layer of the sporangium wall is considered to represent a peritapetal membrane formed from the remnants of this tapetum. The spore R. langii derives from aneurophytalean progymnosperms. In light of the new evidence on spore/sporangium characters, and hypotheses of spore wall development based on interpretation of these, the evolutionary relationships of the progymnosperms are considered in terms of their origins and relationship to the seed plants. It is concluded that there is a smooth evolutionary transition between Apiculiretusispora‐type spores of certain basal euphyllophytes, Rhabdosporites‐type spores of aneurophytalean progymnosperms and Geminospora‐/Contagisporites‐type spores of heterosporous archaeopteridalean progymnosperms. Prepollen of basal seed plants (hydrasperman, medullosan and callistophytalean pteridosperms) are easily derived from the spores of either homosporous or heterosporous progymnosperms. The proposed evolutionary transition was sequential with increasing complexity of the spore/pollen wall probably reflecting increasing sophistication of reproductive strategy. The pollen wall of crown group seed plants appears to incorporate a completely new developmental mechanism: tectum and infratectum initiation within a glycocalyx‐like Microspore Surface Coat. It is unclear when this feature evolved, but it appears likely that it was not present in the most basal stem group seed plants.     

TETRAXYLOPTERIS SCHMIDTII: ITS FERTILE PARTS AND ITS RELATIONSHIPS WITHIN THE ANEUROPHYTALES

The fertile branching system of Tetraxylopteris is composed of successive “nodes” bearing opposite and decussately arranged, upcurved sporangial complexes. By means of the transfer technique the morphology of the sporangial complex was revealed. It consists of a main stalk which dichotomizes twice producing four major branches. Each of the four branches is further subdivided three times, the subdivisions being arranged alternately and pinnately. The ultimate divisions bear the sporangia singly and terminally. The sporangial complexes decrease in size distally and are more tightly curved at the apex. The sporangia are oblong-oval with an acute apex. The spores are identical to the dispersed spore taxon Rhabdosporites langii, Richardson. They are spherical, trilete and pseudosaccate with a fine granular ornament on the pseudosaccus. They are 75–176 μ in diameter and show developmental stages from young tetrads to separated, fully mature spores depending on the age of the sporangium from which they were obtained. This is the first account of spores in sporangia of Tetraxylopteris. The diagnosis of the genus and species are emended to include the new information and the order Aneurophytales is redefined.     

Early vascular land plants: proof and conjecture

Megafossil evidence does not fill the 'evolutionary gap' between land plants and their hypothetical green algal ancestors. Rare Late Silurian vascular plant megafossils provide little information about the morphological, physiological, biochemical, and ecological steps that preceded their evolution. Dissociated trilete spores, spore tetrads, cuticle- and tracheid-like structures far exceed the abundance and diversity of Silurian vascular plant megafossils, and appear millions of years before them. In reference to whole-bodied organisms, these or analogous structures belong to land plants or emergent aquatics; they may represent plants evolutionarily intermediate between green algae and descendent vascular plants at the bryophyte or pre-bryophyte stages. Changes in the cellular biochemistry of pre-Devonian land plants in response to the selective pressures of terrestrial life may have led to the origin of lignin and cutin, neither of which has any counterpart among the algae, and to the evolutionary surge of the vascular plants in the Early Devonian represented by the plant megafossil record. Positive correlation between abundance and diversity of trilete spores and shallow-water, nearshore sites reinforces conclusions based on morphology that a terrestrial flora existed well prior to the appearance of vascular plant megafossils.     

Lower Devonian spore assemblages from the Arbuthnott Group at Canterland Den in the Midland Valley of Scotland

The Canterland Den locality in the Midland Valley of Scotland exposes sediments belonging to the Arbuthnott Group. They consist of typical Lower Old Red Sandstone terrestrial fluviatile deposits, which probably accumulated in an inland intermontane basin. Palynological investigation of these sediments has yielded diverse and well preserved palynomorph assemblages dominated by land-derived forms: spores, phytodebris (dispersed cuticles and tubular structures) and rare fragments of arthropod cuticle. Interestingly, rare acritarchs interpreted as deriving from non-marine algae are also present. The spore assemblages all belong to the lower subzone of the micrornatus–newportensis Spore Assemblage Biozone, indicating an early Lochkovian (Early Devonian) age. They are similar in general characteristics to previously described spore assemblages from the Arbuthnott Group, but exhibit minor differences in terms of taxon composition. They differ more significantly from coeval spore assemblages from the lowland floodplain deposits of the Anglo-Welsh basin, in terms of both relative abundance of morphotypes and taxon composition. An intriguing feature of the assemblage is the high abundance of undissociated spore tetrads. It is suggested that such tetrads are a genuine feature of Lochkovian spore assemblages, perhaps reflecting more flexible and unconventional reproductive strategies exhibited by early land plants.     

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。     

SPORE ULTRASTRUCTURE OF THE MYXOMYCETE PHYSARUM POLYCEPHALUM

Spores of the true slime mold Physarum polycephalum were examined at several stages of their development by means of scanning and transmission electron microscopy. The spores were globose, spine-covered structures produced within a sporangium enclosed in a tough, noncellular peridium. Cytologically, the spore represented a typical eukaryotic cell, having discrete organelles similar to spores of other myxomycetes. The presence of dictyosomes, helical filaments, and microbodies in these cells, as well as the further elucidation of the cell wall and the “polysaccharide-containing” areas, represent new contributions to the ultrastructure of the myxomycete spore. Of special interest were observations of metaphase nuclei just prior to spore cleavage, interphase nuclei in young spores, and nuclei in mature spores containing synaptonemal complexes. These observations indicate that in Physarum polycephalum mitosis occurs just prior to spore cleavage, and meiosis takes place after spore cleavage.