SIPHUNCULAR STRUCTURE IN SOME FOSSIL COLEOIDS AND RECENT SPIRULA

Abstract: In Jurassic Phragmoteuthis huxleyi Donovan (Order Phragmoteuthida) the siphuncular wall shows unique structural and morphological features. The septal neck is short, about one‐eighth of chamber length, but the connecting ring is extremely long, extending through 5–6 chambers. The permeable siphuncular wall in each chamber is, therefore, unusually thick and consists of 5–6 consecutive connecting rings. Each connecting ring is calcified and has a highly porous structure in being composed of bundles of spicular crystallites, orientated more or less at right angles to the siphuncular wall, and separated by smaller or larger interspaces. A restudy of the belemnoid Megateuthis gigantea (Schlotheim) and the aulacoceratid Mojsisovicsteuthis? shows that the connecting rings in these taxa are also calcified. Each ring has a length of two chambers and consists of several calcified lamellae that are traversed by minute pores. The permeable siphuncular wall in each chamber therefore consists of two consecutive connecting rings separated by a porous prismatic layer. In Recent Spirula the connecting ring is composed of two layers: an outer spherulitic‐prismatic layer and an inner glycoprotein layer, of which the latter is not preserved in dry shells. The connecting ring structure is here similar to that in Recent Nautilus. Our study shows that at least three different structural types of siphuncular wall occur in coleoids. The phragmoteuthid connecting ring has a primitive structure, unknown in other cephalopods. This indicates that this taxon has no closer relationship with other coleoid taxa. The belemnitid‐aulacoceratid connecting ring is calcified and traversed by numerous pore canals. It shows a certain structural similarity to that in fossil actinoceratid and orthoceratid nautiloids. The spirulid connecting ring is structurally similar to that in Recent Nautilus and fossil nautilitid and tarphyceratid nautiloids. Thus the connecting ring structure indicates that coleoids include several, phylogenetically clearly separated lineages.     

Characterization of actinoceratoid cephalopods by their siphuncular structure

Partially phosphatized connecting rings show well-preserved structural details in the actinoceratoid Adamsoceras holmi (Ormoceratidae) from the Lower Ordovician (Kundan) of Estonia. Each connecting ring is composed of an outer, thin, spherulitic-prismatic layer, and an inner, thick, calcareous, lamellar layer, the latter being traversed by numerous large pores. The calcareous lamellar inner layer occurs in the connecting rings of three other actinoceratoids: Eushan-tungoceras pseudoimbricatum, Rayonnoceras solidiforme , and Huroniella sp. In Huroniella sp. and R. solidiforme , this layer is traversed by similar large pores as in Adamsoceras holmi , but in E. pseudoimbricatum the pores were probably too narrow to be recognized. The described structural type of the connecting rings, together with well-developed annular endosiphuncular deposits, are here considered as characteristic for actinoceratoids. The inflated connecting ring in actinoceratoids had a great surface area, which increased the number of pores across it. The permeability and emptying rates of the cameral liquid through the connecting ring could therefore have been as high or higher than that in Nautilus . In being composed of numerous calcified lamellae, the mechanical strength of the connecting ring against hydrostatic pressure could have also been the same or higher than that in Nautilus.     

A new view on Nematothallus: coralline red algae from the Silurian of Gotland

Abstract:  The thalloid carbonaceous fossil Nematothallus Lang, 1937, has been widely interpreted as an early Palaeozoic land‐plant, despite the absence of a convincing modern analogue. Exceptionally well‐preserved nematophyte cuticle from the Late Silurian Burgsvik Sandstone Formation, Gotland provides additional insight into the organism’s anatomy, phylogenetic affiliations and ecology. Because this material exhibits additional characters not present in the type material we assign it to Nematothallopsis gotlandii gen. et sp. nov. The organism was constituted of a close‐packed layer of palisade‐like filaments covered by a cuticle that bears a characteristic pseudocellular pattern on its inner surface. Apertures in this cuticle are often encircled by a ring of multicellular filaments, which are sometimes associated with spheroidal, spore‐like entities. In the light of the conspicuous similarity of the palisade layers to the pseudoparenchymatous tissue of coralline red algae, and of the filament‐fringed apertures to their reproductive conceptacles, we reconstruct the Nematothallopsis organism as an extinct rhodophyte and re‐evaluate the putative terrestrial habit of cuticular nematophytes in general.     

REASSESSMENT OF THE LITTLE‐KNOWN CRUSTOSE RED ALGAL GENUS POLYSTRATA (GIGARTINALES), BASED ON MORPHOLOGY AND SSU rDNA SEQUENCES1

The taxonomic distinctiveness of the crustose red algal genus Polystrata Heydrich (Peyssonneliaceae) is confirmed on the basis of morphological and molecular data. The vegetative and reproductive morphology of the type species Polystrata dura Heydrich is newly described. Polystrata thalli are thick multi‐layered crusts, each crust of which is composed of a mesothallus, a superior perithallus, and an inferior perithallus. P. dura is characterized by a poorly developed inferior perithallus consisting of single‐celled perithallial filaments and each layer of multi‐layered crusts being closely adherent to the parental layer. This Polystrata species is identical to Peyssonnelia species, the type genus of the Peyssonneliaceae in the morphology of sexual reproductive organs: a carpogonial branch and an auxiliary cell branch are formed laterally on respective nemathecial filaments; the gonimoblasts are developed from connecting filaments and auxiliary cells; the spermatangia are produced in male and female nemathecia; and the spermatangial filament produces a series of one to four paired spermatangia that form a whorl surrounding each central cell (the Peyssonnelia dubyi‐type development). Polystrata fosliei (Weber‐van Bosse) Denizot is clearly distinguished from P. dura by an inferior perithallus as well‐developed as the superior perithallus, and each layer of multi‐layered crusts being loosely adherent to the parental layer. In our small subunit rDNA trees of the Peyssonneliaceae, these Polystrata species formed a clade with low to medium supports, although the phylogenetic position of Polystrata was unresolved in this family. Therefore, the thallus structure of Polystrata may be regarded as an important taxonomic character at the genus rank.     

Embryonic development of the axial column in the little skate,Leucoraja erinacea

The morphological patterns and molecular mechanisms of vertebral column development are well understood in bony fishes (osteichthyans). However, vertebral column morphology in elasmobranch chondrichthyans (e.g., sharks and skates) differs from that of osteichthyans, and its development has not been extensively studied. Here, we characterize vertebral development in an elasmobranch fish, the little skate, Leucoraja erinacea , using microCT, paraffin histology, and whole‐mount skeletal preparations. Vertebral development begins with the condensation of mesenchyme, first around the notochord, and subsequently around the neural tube and caudal artery and vein. Mesenchyme surrounding the notochord differentiates into a continuous sheath of spindle‐shaped cells, which forms the precursor to the mineralized areolar calcification of the centrum. Mesenchyme around the neural tube and caudal artery/vein becomes united by a population of mesenchymal cells that condenses lateral to the sheath of spindle‐shaped cells, with this mesenchymal complex eventually differentiating into the hyaline cartilage of the future neural arches, hemal arches, and outer centrum. The initially continuous layers of areolar tissue and outer hyaline cartilage eventually subdivide into discrete centra and arches, with the notochord constricted in the center of each vertebra by a late‐forming “inner layer” of hyaline cartilage, and by a ring of areolar calcification located medial to the outer vertebral cartilage. The vertebrae of elasmobranchs are distinct among vertebrates, both in terms of their composition (i.e., with centra consisting of up to three tissues layers—an inner cartilage layer, a calcified areolar ring, and an outer layer of hyaline cartilage), and their mode of development (i.e., the subdivision of arch and outer centrum cartilage from an initially continuous layer of hyaline cartilage). Given the evident variation in patterns of vertebral construction, broad taxon sampling, and comparative developmental analyses are required to understand the diversity of mechanisms at work in the developing axial skeleton of vertebrates. J. Morphol. 278:300–320, 2017. © 2017 Wiley Periodicals, Inc.     

Comparative microsporogenesis and anther development of selected species from Magnoliaceae

Stamen development and microsporogenesis of four species from Magnoliaceae was investigated in order to provide additional data from this family. Stamen bases were found to be wide and short, without morphological differentiation in Magnolia moto, M. paenetalauma and Woonyoungia septentrionalis. In contrast, stamens are distinctly differentiated into anther and filament regions in Michelia crassipes. The orientation of dehiscence is introrse, introrse‐latrorse and latrorse in M. moto, M. paenetalauma and M. crassipes, respectively. The vascular bundles range from three to five (M. moto, M. paenetalauma) to one (M. crassipes). The amount of the connective tissue has been reduced from three to two times of the sporogenous tissue in M. moto and M. paenetalauma. The two parts are nearly equal in M. crassipess. In W. septentrionalis, the orientation of dehiscence, the vascular bundles and the size of the connective tissue vary in different parts of the floral receptacle. The endothecium and endothecial‐like cells form a ring that encloses the entire anther. The middle layer cells originate from both the outer and inner secondary parietal layers, and start to degenerate gradually at the microspore interphase stage or meiosis stage. The tapetum is of the secretory type, derived from the inner secondary parietal cells. The mature anther wall is composed of one epidermal, one endothecial, three to four middle layer(s) and one glandular tapetum. Only one epidermis, one endothecium, and the remnants of the middle layer and tapetum are left before anther dehiscence. Microspore tetrads appear as isobilateral, tetrahedral, decussate and T‐shaped, produced by a modified simultaneous microsporogenesis, which have evolved from the common ancestor of all Magnoliaceae. Our results support an ancestral state with stamens with non‐marginal sporangia and the amount of sterile tissue exceeding the amount of sporogenous tissue, and evolutionary trends toward equalization of the amount of fertile and sterile tissue on the stamen.     

The jaw apparatuses of Cretaceous Phylloceratina (Ammonoidea)

The jaw apparatuses of two species of Late Cretaceous Phylloceratina (Ammonoidea), Hypophylloceras subramosum and Phyllopachyceras ezoensis, are described on the basis of well‐preserved in situ material from Hokkaido, Japan. Gross morphological and X‐ray CT observations reveal that the upper and lower jaws of the two species are essentially similar in their overall structure. Their upper jaws consist of a shorter outer lamella and a pair of larger, wing‐like inner lamellae that become narrower and join together in the anterior portion, as in those of other ammonoids. The upper jaws of the two phylloceratid species are, however, distinguishable from those of other known ammonoids by the presence of a thick, arrowhead‐shaped calcified rostral tip. The lower jaws of the two species consist of a short, reduced inner lamella and a large, gently convex outer lamella covered with a thin calcareous layer, the features of which are common with the rhynchaptychus‐type lower jaws of the Cretaceous Lytoceratina. In the presence of a sharply pointed, thick calcareous tip on upper and lower jaws, the jaw apparatuses of the Phylloceratina resemble those of modern and fossil nautilids, suggesting that they were developed to serve a scavenging predatory feeding habit in deeper marine environments. This and other studies demonstrate that at least some Mesozoic rhyncholites and conchorhynchs are attributable to the Phylloceratina and Lytoceratina.     

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.     

INTERMEDIATE FEATURES OF CYANELLE DIVISION OF CYANOPHORA PARADOXA (GLAUCOCYSTOPHYTA) BETWEEN CYANOBACTERIAL AND PLASTID DIVISION1

Cyanelles of glaucocystophytes may be the most primitive of the known plastids based on their peptidoglycan content and the sequence phylogeny of cyanelle DNA. In this study, EM observations have been made to characterize the cyanelle division of Cyanophora paradoxa Korshikov and to gain insights into the evolution of plastid division. Constriction of cyanelles involves ingrowth of the septum at the cleavage site with the inner envelope membrane invaginating at the leading edge and the outer envelope membrane invaginating behind the septum. This means the inner and outer envelope membranes do not constrict simultaneously as they do in plastid division in other plants. The septum and the cyanelle envelope became stained after a silver‐methenamine staining was applied for in situ detection of polysaccharides. Septum formation was inhibited by β‐lactams and vancomycin, which are potent inhibitors of bacterial peptidoglycan biosynthesis. These results suggest the presence of peptidoglycan at the septum and the cyanelle envelope. In dividing cyanelles, a single electron‐dense ring (cyanelle ring) was observed on the stromal face of the inner envelope membrane at the isthmus, but no ring‐like structures were detected on the outer envelope membrane. Thus a single, stromal cyanelle ring such as this is quite unique and also distinct from FtsZ rings, which are not detectable by TEM. These features suggest that the cyanelle division of glaucocystophytes represents an intermediate stage between cyanobacterial and plastid division. If monophyly of all plastids is true, the cyanelle ring and the homologous inner plastid dividing ring might have evolved earlier than the outer plastid dividing ring.     

GENERIC HOMES FOR BRITISH SILURIAN LINGULOID BRACHIOPODS

Abstract: Since the brachiopod Order Lingulida has been greatly revised in recent years, with particular emphasis on Cambrian and Ordovician genera, many well‐established Silurian linguloid species from the British Isles have been left without an appropriate genus in which to place them. This is rectified here by allocating the various species, mostly erected in the nineteenth or early twentieth centuries, to a variety of genera which have been mainly erected within the last forty years. Two new genera are erected, Mergliella, with type species Lingula bechei Davidson, 1866 , and Striatilingula, with type species Lingula? striata J. de C. Sowerby, 1839 .     

Silurian craniide brachiopods from Gotland

Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, 1939 , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.     

CONCEPTACLE STRUCTURE OF THE PARASITIC CORALLINE RED ALGA CHOREONEMA THURETII (CORALLINALES) AND ITS TAXONOMIC IMPLICATIONS1

The major diagnostic features for erecting the red algal subfamily Choreonematoideae (Corallinales) were a combination of 1) absence of both cell fusions and secondary pit connections, 2) conceptacle roof and wall comprised of a single cell layer, and 3) presence of tetrasporangial pore plugs within a uniporate conceptacle in the monotypic taxon Choreonema thuretii (Bornet) Schmitz. Because this alga is a parasite, the absence of secondary cell connections is most likely an adaptation to a reduced thallus. This study shows that all conceptacles are not composed of a file of cells but rather a single layer of epithallial cells that are underlain by a thick layer of calcified acellular material; both epithallial cells and the calcified layer are produced by peripheral sterile cells. Although the outermost tetrasporangial pore canal is uniporate, there is a calcified acellular multiporate plate recessed just below the rim. The plate is produced by interspersed sterile cells and is continuous with the calcified layer supporting the conceptacle. These unique structures are likely due to parasitism rather than to the ancestral state. Based on these results and a reexamination of published micrographs depicting lenticular cells in Austrolithon intumescens Harvey et Woelkerling, we propose that both subfamily Choreonematoideae and Austrolithoideae are closely allied with subfamily Melobesioideae. This distant relationship to its host (Corallinoideae) plus a combination of unique conceptacle and unusual type of parasitism indicates that C. thuretii is an alloparasite and that it is likely the most ancient red algal parasite studied to date.     

Revised correlation of Silurian Provincial Series of North America with global and regional chronostratigraphic units and δ13Ccarb chemostratigraphy

Cramer, B.D., Brett, C.E., Melchin, M.J., Männik, P., Kleffner, M.A., McLaughlin, P.I., Loydell, D.K., Munnecke, A., Jeppsson, L., Corradini, C., Brunton, F.R. & Saltzman, M.R. 2011: Revised correlation of Silurian Provincial Series of North America with global and regional chronostratigraphic units and δ¹³C_carb chemostratigraphy. Lethaia, Vol. 44, pp. 185–202. Recent revisions to the biostratigraphic and chronostratigraphic assignment of strata from the type area of the Niagaran Provincial Series (a regional chronostratigraphic unit) have demonstrated the need to revise the chronostratigraphic correlation of the Silurian System of North America. Recently, the working group to restudy the base of the Wenlock Series has developed an extremely high‐resolution global chronostratigraphy for the Telychian and Sheinwoodian stages by integrating graptolite and conodont biostratigraphy with carbonate carbon isotope (δ¹³C_carb) chemostratigraphy. This improved global chronostratigraphy has required such significant chronostratigraphic revisions to the North American succession that much of the Silurian System in North America is currently in a state of flux and needs further refinement. This report serves as an update of the progress on recalibrating the global chronostratigraphic correlation of North American Provincial Series and Stage boundaries in their type area. The revised North American classification is correlated with global series and stages as well as regional classifications used in the United Kingdom, the East Baltic, Australia, China, the Barrandian, and Altaj. Twenty‐four potential stage slices, based primarily on graptolite and conodont zones and correlated to the global series and stages, are illustrated alongside a new composite δ¹³C_carb curve for the Silurian. Conodont, graptolite, isotope, New York, Ontario, series, Silurian, stage.     

The extended structure of the periplasmic region of CdsD,a structural protein of the type III secretion system of Chlamydia trachomatis

The type III secretion system (T3SS) is required for the virulence of many gram‐negative bacterial human pathogens. It is composed of several structural proteins, forming the secretion needle and its basis, the basal body. In Chlamydia spp., the T3SS inner membrane ring (IM‐ring) of the basal body is formed by the periplasmic part of CdsD (outer ring) and CdsJ (inner ring). Here we describe the crystal structure of the C‐terminal, periplasmic part of CdsD, not including the last 60 residues. Two crystal forms were obtained, grown in three different crystallization conditions. In both crystal forms there is one molecule per asymmetric unit adopting a similar extended structure. The structures consist of three periplasmic domains (PDs) of similar αββαβ topology as seen also in the structures of the homologous PrgH (Salmonella typhimurium) and YscD (Yersinia enterocolitica). Only in the C2 crystal form, there is a C‐terminal additional helix after the PD3 domain. The relative orientation of the three subsequent CdsD PD domains with respect to each other is more extended than in PrgH but less extended than in YscD. Small‐angle X‐ray scattering data show that also in solution this CdsD construct adopts the same elongated shape. In both crystal forms the CdsD molecules are packed in a parallel fashion, using translational crystallographic symmetry. The most extensive crystal contacts are preserved in both crystal forms, suggesting a possible mode of assembly of the CdsD periplasmic part into a 24‐mer complex forming the outer ring of the IM‐ring of the T3SS.     

The histology and affinities of sinacanthid fishes: primitive gnathostomes from the Silurian of China

On the basis of well preserved specimens from the Lower Silurian of the Tarim Basin, Xinjiang Uygur Autonomous Region and Shiqian County, Guizhou Province, People's Republic of China we describe in detail the histological structure of sinacanthid spines, the only known remains of a group of fishes common in Silurian strata from China. The sinacanthids have previously been assigned either to the acanthodians or to the chondrichthyans. The spine structure is composed of an outer layer of atubular dentine and an inner layer of globular calcified cartilage, and the nature and distribution of these tissues indicates that the spines were formed as a result of interaction between the endoskeleton and dermoskeleton. The tissue distribution and style of growth described herein places the sinacanthids crownwards of the placoderms, and possibly within the total group Chondrichthyes. However, before they can be firmly placed within a phylogenetic scheme, further evidence is required both on the general anatomy of sinacanthids and on the nature of chondrichthyan apomorphies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 379–386.     

Embryology of the dioecious Woonyoungia septentrionalis (Magnoliaceae)

The embryological characteristics and ovular integument development of the dioecious species Woonyoungia septentrionalis (Dandy) Law (Magnoliaceae), which are poorly understood, were investigated under laser scanning confocal microscope (LSCM) and light microscope (LM). The embryological characteristics conform to most of the previously studied species in Magnoliaceae. The anther has 4 microsporangia, and the anther wall develops according to the dicotyledonous type. Cytokinesis at meiosis of the microspore mother cells follows a modified simultaneous type, giving rise to isobilateral or decussate tetrads, and a cell plate is absent, but a membrane was observed. Mature pollen grains are 2‐cellular and have high germination rates. The ovule is anatropous, crassinucellate and bitegmic, and meiotic result in linear tetrads of megaspores, the one at the chalazal end functions directly as an embryo‐sac cell. The development of the embryo sac is of the Polygonum‐type and endosperm formation is of the nuclear type. The outer integument of the ovule differentiates into an outer fleshy and an inner stony layer while the inner integument is reduced to a tanniniferous layer. The normal embryological development, high germination rates of pollen and high seed set indicate that the primary reason for the decline of the species is not to be found in these developmental processes.     

Ultrastructure of calcified cartilage in the endoskeletal tesserae of sharks

The tesserate pattern of endoskeletal calcification has been investigated in jaws, gill arches, vertebral arches and fins of the sharks Carcharhinus menisorrah, Triaenodon obesus and Negaprion brevirostris by techniques of light and electron microscopy. Individual tesserae develop peripherally at the boundary between cartilage and perichondrium. An inner zone, the body, is composed of calcified cartilage containing viable chondrocytes separated by basophilic contour lines which have been called Liesegang waves or rings. The outer zone of tesserae, the cap, is composed of calcified tissue which appears to be produced by perichondrial fibroblasts more directly, i.e., without first differentiating as chondroblasts. Furthermore, the cap zone is penetrated by acidophilic Sharpey fibers of collagen. It is suggested that scleroblasts of the cap zone could be classified as osteoblasts. If so, the cap could be considered a thin veneer of bone atop the calcified cartilage of the body of a tessera. By scanning electron microscopy it was observed that outer and inner surfaces of tesserae differ in appearance. Calcospherites and hydroxyapatite crystals similar to those commonly seen on the surface of bone are present on the outer surface of the tessera adjacent to the perichondrium. On the inner surface adjoining hyaline cartilage, however, calcospherites of variable size are the predominant surface feature. Transmission electron microscopy shows calcification in close association with coarse collagen fibrils on the outer side of a tessera, but such fibrils are absent from the cartilaginous matrix along the under side of tesserae. Calcified cartilage as a tissue type in the endoskeleton of sharks is a primitive vertebrate characteristic. Calcification in the tesserate pattern occurring in modern Chondrichthyes may be derived from an ancestral pattern of a continuous bed of calcified cartilage underlying a layer of perichondral bone, as theorized by Ørvig (1951); or the tesserate pattern in these fish may itself be primitive.     

Molecular architecture of the N‐type ATPase rotor ring from Burkholderia pseudomallei

The genome of the highly infectious bacterium Burkholderia pseudomallei harbors an atp operon that encodes an N‐type rotary ATPase, in addition to an operon for a regular F‐type rotary ATPase. The molecular architecture of N‐type ATPases is unknown and their biochemical properties and cellular functions are largely unexplored. We studied the B. pseudomallei N₁N_o‐type ATPase and investigated the structure and ion specificity of its membrane‐embedded c‐ring rotor by single‐particle electron cryo‐microscopy. Of several amphiphilic compounds tested for solubilizing the complex, the choice of the low‐density, low‐CMC detergent LDAO was optimal in terms of map quality and resolution. The cryoEM map of the c‐ring at 6.1 Å resolution reveals a heptadecameric oligomer with a molecular mass of ~141 kDa. Biochemical measurements indicate that the c₁₇ ring is H⁺ specific, demonstrating that the ATPase is proton‐coupled. The c₁₇ ring stoichiometry results in a very high ion‐to‐ATP ratio of 5.7. We propose that this N‐ATPase is a highly efficient proton pump that helps these melioidosis‐causing bacteria to survive in the hostile, acidic environment of phagosomes.     

Crystal structure of the C‐terminal domain of the Salmonella type III secretion system export apparatus protein InvA

InvA is a prominent inner‐membrane component of the Salmonella type III secretion system (T3SS) apparatus, which is responsible for regulating virulence protein export in pathogenic bacteria. InvA is made up of an N‐terminal integral membrane domain and a C‐terminal cytoplasmic domain that is proposed to form part of a docking platform for the soluble export apparatus proteins notably the T3SS ATPase InvC. Here, we report the novel crystal structure of the C‐terminal domain of Salmonella InvA which shows a compact structure composed of four subdomains. The overall structure is unique although the first and second subdomains exhibit structural similarity to the peripheral stalk of the A/V‐type ATPase and a ring building motif found in other T3SS proteins respectively.