Fine structure of the nematocytes of Polypodium hydriforme Ussov (Cnidaria)

The fine structure of the stinging cells (nematocytes) and stinging capsules (nematocysts) is described for Polypodium hydriforme. a freshwater coclenterate with a prominent endoparasitie stage in its life cycle. All the nematocysts belong to the type of lesser glutinants (atrichous isorhiza) and fall into three size classes. The internal structure of the capsules is similar in the three classes. A novel type of organization of the cnidocil apparatus of the nematocysts is described. The cnidocil lacks a root fibre and its kinctosome sits directly on the operculum of the nematocyst, so that the entire cnidocil apparatus has a radial rather than bilateral symmetry. It is compared with that of other types of nematocytes and its similarity with the mechanoreccptors of the coelentcratcs is noted. The possible place of the Polypodium nematocytes in the evolution of the collar receptors of the Metazoa is discussed.     

Peculiarities of the embryonic development of Polypodium hydriforme Ussov (Coelenterata), a parasite of acipenserid oocytes

"Unicellular" stages (107 specimens) and multicellular stages (64 specimens) of embryogenesis of Polypodium, found in 14 sterlet (Acipenser ruthenus L.) females, have been studied with light microscopy, cytophotometry, and autoradiography following incubation with 3H-uridine. All stages of the embryonic development occur inside host oocytes. The "unicellular" stage includes a binucleate cell with unequally sized nuclei; separation inside it of a small cell around the smaller nucleus, i.e. transformation of the single cell into a complex of 2 cells, the larger one enveloping the smaller; formation of a cavity inside the nucleus of the large (outer) cell, and migration of the small cell into it, and "cell-in-a-cell" stage, the small (generative) cell being inside the cavity formed by the nucleus of the large (trophic) cell. The latter gives rise to a hypertrophied but still unicellular envelope around the embryo, the trophamnion. The multicellular stages start with segmentation of the generative cell into blastomeres. These form a morula lying inside the cavity of the trophamnion. Gastrulation occurs by morular delamination. The inversion of the germ layers, typical of parasitic Polypodium stages, apparently arises during gastrulation. Both the generative cell ("egg") and the blastomeres are haploid, at least until the morula stage. The eggs of Polypodium are the smallest ones among coelenterates; they lack yolk and develop without fertilization. Diploidy seems to be restored during segmentation. The trophamnion cell grows, its nucleus becomes highly polypoid, and its cytoplasm accumulates mucoprotein inclusions. Both the blastomere nuclei and the trophamnion nucleus have large nucleoli and actively synthesize RNA. The stages of embryogenesis of Polypodium closely correspond to stages of the host oogenesis. The embryonic development of Polypodium lasts several years and is the slowest among coelenterates. However, it has some features typical of the class Hydrozoa.     

Morphology,ultrastructure, and development of the parasitic larva and its surrounding trophamnion of Polypodium hydriforme Ussov (Coelenterata)

Summary The larval stage of Polypodium hydriforme is planuliform and parasitic inside the growing oocytes of acipenserid fishes. The larva has inverted germ layers and a special envelope, the trophamnion, surrounding it within the host oocyte. The trophamnion is a giant unicellular provisory structure derived from the second polar body and performing both protective and digestive functions, clearly a result of adaptation to parasitism. The trophamnion displays microvilli on its inner surface, and irregular protrusions anchoring it to the yolk on its outer surface. Its cytoplasm contains long nuclear fragments, ribosomes, mitochondria, microtubules, microfilaments, prominent Golgi bodies, primary lysosomes, and secondary lysosomes with partially digested inclusions.The cells of the larva proper are poorly differentiated. No muscular, glandular, neural, interstitial, or nematocyst-forming cells have been found. The entodermal (outer layer) cells bear flagella and contain rough endoplasmic reticulum; the ectodermal (inner layer) cells lack cilia and contain an apical layer of acid mucopolysaccharid granules. The cells of both layers contain mitochondria, microtubules, and Golgi bodies; their nuclei display large nucleoli with nucleolonema-like structure, decondensed chromatin, and some perichromatin granules. At their apical rims, the ectodermal cells form septate junctions; laterally, the cells of both layers form simple contacts and occasional interdigitations. The lateral surfaces of entodermal cells are strengthened by microtubules.     

Correction: Phylogenetic placement of the enigmatic parasite, Polypodium hydriforme, within the Phylum Cnidaria

Correction to Evans, N.M., Lindner, A., Raikova, E.V., Collins, A.G. and Cartwright, P. Phylogenetic placement of the enigmatic parasite, Polypodium hydriforme, within the phylum Cnidaria. BMC Evol Biol, 2008, 8:139.     

Amoebocytes in mesoglea of Polypodium hydriforme

Mesogleal amoebocytes of free-living Polypodium hydriforme have been studied with transmission electron microscope. The amoebocytes have numerous processes and contain cytoplasmic vacuoles with fibrous material of different density. The phenomenon of cell death (apoptosis) of mesogleal amoebocytes is described. Chromatin of dying cells becomes condensed forming picnotic "caps" in the nucleus. No mitotic cells were encountered among mesogleal amoebocytes. The origin and functions of mesogleal amoebocytes of P. hydriforme are discussed.     

Polypodium hydriforme in the eggs of the sterlet from the northern Dvina river

The paper provides data on the invasion of Acipenser ruthenus (sterlet) in the North Dvina River by Polypodium hydriforme (Cnidaria). Prevalence and intensity of invasion proved to be similar for two fishing sites. Prevalence of invasion exceeded 88%, thus being exceptionally high for P. hydriforme. Intensity of invasion was from 1 to 436 eggs per female. A verage percentage of infected eggs was about 1%.     

Structural characteristics of nematocyst stinging threads from the parasitic cnidarian Polypodium hydriforme

The structure of discharged nematocyst stinging threads present in free-living individuals of Polypodium hydriforme was studied by scanning and transmission electron microscopy. Not all cnidae of P. hydriforme proved to be atrichous isorhizas (as previously was accepted), but only one of the five nematocyst categories studied. A unique feature of P. hydriforme nematocysts was revealed: their stinging threads possess two strands of spines, rather than 5 or 3, as in Narcomedusae and other Cnidarians, respectively. This fact supplements the evidence in favour of P. hydriforme being a rather isolated branche in the phylogenetic tree of Cnidaria.     

Cytomorphological characteristics of Polypodium hydriforme and problems of myxozoan and cnidarian phylogeny

The present review analyses cytomorphological characters of the parasitic cnidarian Polypodium hydriforme, discriminating between those of bilateral (triploblastic) animals, common characters shared with the Myxozoa, and the unique characters of this species. Phylogenetic position of the group of parasitic cnidarians and of the class Polypodiozoa is discussed. A conclusion is made that the cytomorphological characters as well as 18S rDNA analysis of P. hydriforme and Myxozoa justify establishment of a new taxonomic group (a clade) of parasitic cnidarians (Endocnidozoa) uniting Polypodiozoa and Myxozoa (Zrzavy, Hypsa, 2003). The unique characters of P. hydriforme suggest that the phylum Cnidaria is more diverse than commonly supposed, and that P. hydriforme is not an aberrant cnidarian species but a relic organism, which might originally belong to the cnidarian class Polypodiozoa, which underwent reduction in the course of adaptation to parasitism.     

Ultrastructure of the calcium-sequestering gastrodermal cell in the hydroid Hydractinia symbiolongicarpus (Cnidaria, Hydrozoa)

Large, free-floating crystals of calcium carbonate occur in vacuoles of gastrodermal cells of the hydroid Hydractinia symbiolongicarpus. Here, morphological details about the process by which these cells accumulate and sequester calcium are provided by a cytochemical method designed to demonstrate calcium at the ultrastructural level. Electron-dense material presumably indicative of the presence of calcium was EGTA-sensitive and was shown by parallel electron energy loss spectroscopy (EELS) and energy spectroscopic imaging (ESI) to contain calcium. Calcium occurred in only one cell type, the endodermally derived gastrodermal cell. In these cells, the electron-dense material appeared first as a fine precipitate in the cytosol and nucleus and later as larger deposits and aggregates in the vacuole. During the life cycle, gastrodermal cells of the uninduced planula and the planula during metamorphic induction sequestered calcium. In primary polyps and polyps from established colonies, gastrodermal cells sequestered calcium, but the endodermal secretory cells did not. Our observations support the hypothesis that gastrodermal cells function as a physiological sink for calcium that enters the organism in conjunction with calcium-requiring processes such as motility, secretion, and metamorphosis.     

The parasitic coelenterate, Polypodium hydriforme USSOV, from the eggs of the American acipenseriform Polyodon spathula.

No significant differences in macro- and micromorphology were found between the parasitic stolon and free-living polyps of Polypodium sp. obtained from infected eggs of the North American acipenseriform fish Polyodon spathula and corresponding developmental stages of Polypodium hydriforme Ussov, parasitic in the Volga sterlet (Acipenser ruthenus). Therefore, both the American and the European forms of Polypodium belong to the species P. hydriforme Ussov.     

Ultrastructure of nematocyst discharge in catch tentacles of the sea anemone Haliplanella luciae (Cnidaria: Anthozoa)

The mature nematocyst lies just beneath the cnidodyte plasma membrane. A microtubule array surrounds the nematocyst capsule just beneath the capsule tip. We propose that the array helps to hold the capsule at the cnidocyte cell surface until discharge. The undischarged capsule tip is sealed by three apical flaps, joined together along complex radial seams. The seams are filled with subunits that appear to bind the flaps together. Upon discharge, the flaps separate along the radial seams to permit thread eversion. The everted thread is lined on both sides by subunits that are stained by antimonate, indicating that they bind calcium. We suggest that, together, the subunits hold the uneverted thread in its folded and coiled configuration. Thread eversion would follow subunit uncoupling. The capsule and thread interiors of partially discharged nematocysts are stained by antimonate. In contrast, the capsule and thread interiors of fully discharged nematocysts are not stained by antimonate. Thus, nematocyst calcium might be injected into the target tissue where it is presumed to act in conjunction with nematocyst venom to promote cell death.     

Ultrastructure of a novel eurytele nematocyst of Carybdea alata Reynaud (Cubozoa,Cnidaria)

The ultrastructural characteristics of nematocysts from the cubozoan Carybdea alata Reynaud, 1830 (Hawaiian box jellyfish) were examined using light, scanning and transmission electron microscopy. We reclassified the predominant nematocyst in C. alata tentacles as a heterotrichous microbasic eurytele, based on spine, tubule and capsule measurements. These nematocysts exhibited a prominent and singular stylet, herein referred to as the lancet. Discharged nematocysts from fixed tentacle preparations displayed the following structures: a smooth shaft base, lamellae, a hemicircumferential fissure demarking the proximal end of a stratified lancet, and a gradually tapering tubule densely covered with large triangularly shaped spines. The lancet remained partially adjoined to the shaft base in a hinge-like fashion in rapidly fixed, whole-tentacle preparations. In contrast, this structure was not observed in discharged nematocyst preparations which involved multiple transfer steps prior to fixation. Various approaches were designed to detect this structure in the absence of fixative. Detached lancets were located in proximity to discharged tubules in undisturbed coverslip preparations of fresh tentacles. In addition, examination of embedded nematocysts from fresh tentacles laid on polyacrylamide gels revealed still-attached lancets. To examine the function of this structure in prey capture, Artemia sp. laden tentacles were prepared for scanning electron microscopy. While carapace exteriors exhibited structures proximal to the lancet, i.e., the nematocyst capsule and shaft base, neither tubule nor lancet structures were visible. Taken together, the morphological data suggested a series of events involved in the discharge of a novel eurytele from C. alata.     

Occurrence and Ultrastructure of a Variant (rho) Form of Mycoplasma

The ultrastructure of a variant (rho) form of Mycoplasma is described. The rho-forms are characterized by dark-ground light microscopy as relatively rigid, unbranched, filamentous organisms with discoidal swellings, and by electron microscopy by the presence of an intracytoplasmic axial fiber extending throughout the length of the cell and associated with a terminal structure of characteristic appearance. In negatively stained preparations the fiber presents a pattern of transverse light and dark major bands, the dark band being divided by a central minor light band. The periodicity of the banding varies from 12.0 to 14.5 nm, and the width of the fiber varies from 40 to 120 nm. The fiber appears to be composed of fibrils aligned parallel to the long axis. The evidence indicates that the fiber contains protein and is devoid of nucleic acid. rho-Forms were commonly found in Mycoplasma strains derived from goats and occasionally in bovine strains. They may have a wider distribution, as the growth medium was shown to be important both for the expression of the rho-character and for the selection of the rho-variant. The functional significance, if any, of the fiber and the terminal structure is unknown.     

Ultrastructure of the neuromuscular system of the polyp of Aurelia aurita L., 1758 (Cnidaria,scyphozoa)

Fine structural study indicates that the neuromuscular system of stage I polyps of Aurelia aurita is exclusively ectodermal. The three major muscle fields are the radial muscles of the oral disc, the longitudinal muscles of the tentacles, and the muscle cords of the septae and the column; the muscle fields are in physical continuity at the peristomial pits and share a common innervation and type of myofibril. The myofibril is striated in the tentacle base, in the outer oral disc, and in the upper part of the muscle cord; it grades into a smooth muscle toward the tentacle tip, the mouth, and the lower part of the cord. There is a fourth field of longitudinal smooth muscle in the pharynx. The nervous system consists of an epithelial sensory cell in the tentacle and a single type of neuron found in the subepithelial layer of the tentacle, oral disc, and muscle cord. The lack of gap junctions suggests that there is no nonnervous conduction system. The subepithelial layer also contains three types of fibers and a type of soma which cannot be characterized as neuronal. The soma is identified as the “neurosecretory cell” described in Chrysaora. The absence of neuromuscular elements in the column and stolon distinguishes the Aurelia aurita collected from Washington, USA, from English polyps previously described.     

The Embryonic Origin of Neurosensory Cells and the Role of Nerve Cells in Metamorphosis in Phialidium gregarium (Cnidaria,Hydrozoa)

Summary

The embryonic origin of the nervous system in Phialidium gregarium was investigated. Entoderm-free planulae, surgically produced by bisection at mid-gastrulation, and normal planulae were examined by light and electron microscopy to determine their cellular composition. The cell types that occur in the epidermis of the normal planula were described. The entoderm-free planulae were found to be devoid of interstitial cells and their derivatives, the nematocytes and ganglion cells. Neurosensory cells were present, however, indicating that they are derivatives of the ectodermal epithelium.

The role of nerve elements in the initiation of metamorphosis was also examined. Normal and entoderm-free planulae treated for four hours with 0.4% colchicine at two, three, or four days of development fail to undergo cesium-induced metamorphosis. Since such treatment in other hydrozoans eliminates interstitial cells and their derivatives [1-3], it might be argued that ganglion cells are necessary to initiate metamorphosis. The observation that entoderm-free planulae, devoid of interstitial cell derivatives, are capable of responding to induction by bacteria or cesium, however, indicates that in Phialidium the colchicine effect is on other cell types. The results are compared with findings for other Cnidaria.     

Rediscovery of Polypodium calirhiza (Polypodiaceae) in Mexico

This study addresses reported discrepancies regarding the occurrence of Polypodium calirhiza in Mexico. The original paper describing this taxon cited collections from Mexico, but the species was omitted from the recent Pteridophytes of Mexico. Originally treated as a tetraploid cytotype of P. californicum, P. calirhiza now is hypothesized to have arisen through hybridization between P. glycyrrhiza and P. californicum. The tetraploid can be difficult to distinguish from either of its putative parents, but especially so from P. californicum. Our analyses show that a combination of spore length and abaxial rachis scale morphology consistently distinguishes P. calirhiza from P. californicum, and we confirm that both species occur in Mexico. Although occasionally found growing together in the United States, the two species are strongly allopatric in Mexico: P. californicum is restricted to coastal regions of the Baja California peninsula and neighboring Pacific islands, whereas P. calirhiza grows at high elevations in central and southern Mexico. The occurrence of P. calirhiza in Oaxaca, Mexico, marks the southernmost extent of the P. vulgare complex in the Western Hemisphere.     

Ultrastructure of neuro‐spirocyte synapses in the sea anemone Aiptasia pallida (Cnidaria,Anthozoa, Zoantharia)

Using transmission electron microscopy of serially sectioned tentacles from the sea anemone Aiptasia pallida, we located and characterized two types of neuro‐spirocyte synapses. Clear vesicles were observed at 10 synapses and dense‐cored vesicles at five synapses. The diameters of vesicles at each neuro‐spirocyte synapse were averaged; clear vesicles ranged from 49–89 nm in diameter, whereas the dense‐cored vesicles ranged from 97–120 nm in diameter. One sequential pair of synapses included a neuro‐spirocyte synapse with clear vesicles (81 nm) and a neuro‐neuronal synapse with dense‐cored vesicles (168 nm). A second synapse on the same cell had dense‐cored vesicles (103 nm). An Antho‐RFamide‐labeled ganglion cell and three different neurites were observed adjacent to spirocytes, but no neuro‐spirocyte synapses were present. Many of the spirocytes also were immunoreactive to Antho‐RFamide. The presence of sequential neuro‐neuro‐spirocyte synapses suggests that synaptic modulation may be involved in the neural control of spirocyst discharge. The occurrence of either dense‐cored or clear vesicles at neuro‐spirocyte synapses suggests that at least two types of neurotransmitter substances control the discharge of spirocysts in sea anemones. J. Morphol. 241:165–173, 1999. © 1999 Wiley‐Liss, Inc.