Ultrastructural data on the scales of the dipnoan Protoptems annectens (Sarcopterygii, Osteichthyes)

The structure and the mineralization of the scales of the living dipnoan (lungfish) Protoptems annectens have been investigated by transmission electron microscopy (TEM). The thin and imbricated scales are composed of two layers: the squamulae and the basal plate. At the outer surface, the squamulae form isolated plates superficially ornamented with spines and concretions and made up of acellular bone. After demineralization, the squamulae show a heterogeneous organic matrix composed of thin randomly oriented collagen fibrils forming a loose network within which the concretions appear as electronlucent circular areas. Abundant and aggregated concretions are located within the spines. The crystallites are oriented by the collagen fibrils except in the concretions. Anchoring bundles composed of parallel collagen fibrils arise from the squamulae and connect the scales to the overlying dermis.
The basal plate, the most developed part of the scale, is made up of isopedine. Its main component consists of thick, closely packed collagen fibrils organized in a 'double twisted plywood-like structure'. Fibroblasts are present in the basal plate. Mineralization occurs only in few plies located beneath the squamulae. Mandl's corpuscles are found in front of the mineralization front. The mineral deposit is oriented by the collagen fibrils.
The scales of Protoptems annectens differ from the typical elasmoid scales of the teleosts by the peculiar structure of the squamulae, nevertheless they show enough structural characteristics to support the hypothesis that they can be considered as scales of the elasmoid grade, which have retained some plesiomorphic characteristics.     

Scale diversity in bigeye tuna (Thunnus obesus): Fat‐filled trabecular scales made of cellular bone

Tunas of the genus Thunnus possess many morphological and physiological adaptations for their high‐performance epipelagic ecology. Although Thunnus anatomy has been studied, there are no quantitative studies on the structure of their scales. We investigated the scales of bigeye tuna (Thunnus obesus) from ten regions of the body using micro computed tomography (µCT)‐scanning and histology to quantitatively and qualitatively compare regional scale morphology. We found a diversity of scale sizes and shapes across the body of bigeye tuna and discriminant function analysis on variables derived from µCT‐data showed that scales across the body differ quantitatively in shape and size. We also report the discovery of a novel scale type in corselet, tail, and cheek regions. These modified scales are ossified shells supported by internal trabeculae, filled with fat, and possessing an internal blood supply. Histological analysis showed that the outer lamellar layers of these thickened scales are composed of cellular bone, unexpected for a perciform fish in which bone is typically acellular. In the fairing region of the anterior body, these fat‐filled scales are stacked in layers up to five scales deep, forming a thickened bony casing. Cheek scales also possess a fat‐filled internal trabecular structure, while most posterior body scales are more plate‐like and similar to typical teleost scales. While the function of these novel fat‐filled scales is unknown, we explore several possible hypotheses for their function.     

Upper Ordovician chondrichthyan‐like scales from North America

Studies of Ordovician micromeric fish scales from the Sandbian of North America have identified a number of scale‐based taxa potentially referable to the chondrichthyans and therefore can be among the stratigraphically oldest representatives of the clade described to date. Two of these, Tezakia hardingensis gen. et sp. nov. and Canyonlepis smithae gen. et sp. nov., are formally described herein. Tezakia gen. nov. scales are composed exclusively of tubular dentine and possess polyodontocomplex crowns with a characteristically large primordial odontode. Similar scale crown architecture has been reported only in the reputed chondrichthyan Altholepis composita (Lower Devonian of Podolia, Ukraine), and on these grounds, the two are united within the newly erected Altholepidiformes ordo nov. Multiple odontocomplexes are also a feature of Canyonlepis gen. nov. scale crowns; however, the latter do not demonstrate prominent primordial odontodes and are supported by a base composed of acellular bone. Additional data suggest that both taxa possess a combination of characteristics (areal crown growth, scale symmetry, linear odontocomplex architecture and absence of enamel, osteons, cancellous bone and hard‐tissue resorption) previously documented to occur only in chondrichthyan scales. This study contributes to a growing body of evidence that reveals the presence of diverse tissue types (bone, tubular and atubular dentine) and morphogenetic patterns (odontocomplex and non‐odontocomplex type of scale crown growth) in the dermal skeleton of putative Ordovician chondrichthyans.     

Euconodont hard tissue: preservation patterns of the basal body

Conodont elements, consisting of crown and basal tissue are the well-known fossilized hard parts of Conodonta (extinct marine chordates), but the taphonomic processes leading to decomposition or remineralization of the basal tissue are not well understood. Here we focus on the taphonomy of basal tissue, reviewing the published record and describing new material from Asia and Europe (248 occurrences globally). These include crown and basal tissue in conjunction, and isolated basal bodies showing different stages of preservation. Some isolated specimens resemble phosphate rings similar to those assigned to Phosphannulus universalis. High-resolution biostratigraphy indicates that the lamellar type of conodont basal tissue is found in all facies and depositional environments. Other basal tissue types, described in the literature as tubular, mesodentine, spherulitic or lamellar with canalules, are limited to the early Palaeozoic and found exclusively in siliciclastic deposits (with the exception of spherulitic tissue). Although the stratigraphic record of basal tissue spans the range of Euconodonta (Cambrian–Triassic), this study shows that most of the isolated plate and ring-like structures are derived from early Palaeozoic coniform conodonts. Basal tissue of platform-type elements has a much more fragile shape and is therefore rarely preserved as a recognizable isolated unit.     

革鲀科鳞片构造的扫描电镜观察和比较研究

革鲀科的鳞片构造不同于一般真骨鱼类的骨鳞,由特化的基板和鳞棘两部分组成。中国有革鲀科鱼类12属19种,本文研究了其中10属14种的鳞片构造,运用扫描电镜进行了观察和比较。根据基板表面棱突的特征,可将革鲀科基板构造分为3个式型、2个亚型;根据鳞棘数目、形状及排列方式的不同,可以划分为4个式型。革鲀科各种间的鳞片具有不同的性状,而且比较稳定,因此可以作为分类鉴定的依据之一。     

Observations on the fetus and placenta of a proboscis monkey (Nasalis Iarvatus)

The villous bidiscoid placenta in utero of a proboscis monkey with a fetus of 9.0 cm crown rump length was examined. The immature male fetus weighed 56 g and had a 5.4 cm head circumference. Of particular interest was the stomach, consisting of three parts, large fundus, a narrow tubular portion, and a small pyloric region. The fetal adrenal glands were remarkable in that they were larger than the kidneys. Thus, the adrenal/kidney weight ratio approximated 1.18. The marginally inserted umbilical cord measured 15 cm. Owing to the superficial implantation in the proboscis monkey, penetration of syncytiotrophoblast and a fibrin layer were not observed in the basal plate of the placenta; cytotrophoblastic cell columns attached to the basal plate, consisting of decidua basalis without invasion of trophoblastic giant cells. Intraarterial trophoblastic migration was found in the uterus.     

Cytoskeletal organization and collagen orientation in the fish scales

Summary Immunofluorescence and electron microscopy were used to analyze the relationships between the organization of collagen fibrils in elasmoid scales, and the orientation of microtubules and actin microfilaments in the scleroblasts producing this collagenous stroma. Attention was focused on the basal plate of the scales because of the highly ordered three-dimensional arrangement of the collagen fibrils in superimposed plies forming an acellular plywood-like structure. The collagen fibrils are synthesized by the scleroblasts forming a monolayered pseudo-epithelium, the hyposquama, at the lowest surface of the scale. Fully developed scales with a low collagen deposition rate were compared with regenerating scales active in fibrillogenesis. When an ordered array of the collagen fibrils is found, the innermost collagen fibrils are coaligned with microtubules and actin microfilaments. Thus, because of this coalignment, microtubules and actin microfilaments of the hyposquamal scleroblasts are subjected to consecutive alterations during the formation of the plies of the basal plate. The sequence of events when the collagen fibrils change their direction from one ply to the other in the basal plate is deduced from immunofluorescence and phase-contrast-microscopic observations. During the formation of the orthogonal plywood-like structure in the regenerating scales, first microtubules may change their curse with a rotating angle of about 90°; then, actin microfilaments are disorganized and reorganized by interacting mechanically with the microtubules with which they are coaligned. Collagen fibrils are synthesized in a direction that is roughly perpendicular to that of the preceding ply. The unknown signals inducing the change in direction of the cytoskeleton may be transmitted throughout the hyposquama via gap junctions.This work is dedicated to the memory of Jacques Escaig     

Structure of the skin of Agonus cataphractus (Teleostei)

The skin of the scuted teleost Agonus cataphractus has been investigated by histochemical methods, SEM and TEM. The anterior dorsal skin bears tubercles of epidermis overlying tiny ossifications (scutelets) superficial to the main scutes. The epidermis secretes a cuticular layer containing acidic non-sulphated glycoproteins, but there are no mucous goblet cells in the external skin. Non-mucous sacciform cells of two types are present in the epidermis, also numerous chloride cells. Scanning electron microscopy reveals variation in the microridge pattern of superficial epithelial cells, thought to relate to arrival at the surface and secretion of the cuticle. The major scutes overlap anteriorly, contrary to the normal arrangement of scales, indicating that they are secondary ossifications. The type of mineralization is similar to that of acellular bone. The scutes are set directly in the collagen of the dermis. They have a girdered structure with radial and cross bars, inserting on both faces of a thin plate. The interstices are occupied by unmineralized collagen, and extrinsic collagen bundles impinge on the bone. Non-mineralized parts of the dermis contain tracts of microfibrils in addition to collagen; these are best developed in the flexible gular skin and in the barbels and are interpreted as elastic tissue, although an amorphous component was not seen. The barbels have a core of connective tissue without a cartilaginous skeleton and bear taste buds and numerous chloride cells.     

Histology of the heterostracan dermal skeleton: Insight into the origin of the vertebrate mineralised skeleton

Living vertebrates are divided into those that possess a fully formed and fully mineralised skeleton (gnathostomes) versus those that possess only unmineralised cartilaginous rudiments (cyclostomes). As such, extinct phylogenetic intermediates of these living lineages afford unique insights into the evolutionary assembly of the vertebrate mineralised skeleton and its canonical tissue types. Extinct jawless and jawed fishes assigned to the gnathostome stem evidence the piecemeal assembly of skeletal systems, revealing that the dermal skeleton is the earliest manifestation of a homologous mineralised skeleton. Yet the nature of the primitive dermal skeleton, itself, is poorly understood. This is principally because previous histological studies of early vertebrates lacked a phylogenetic framework required to derive evolutionary hypotheses. Nowhere is this more apparent than within Heterostraci, a diverse clade of primitive jawless vertebrates. To this end, we surveyed the dermal skeletal histology of heterostracans, inferred the plesiomorphic heterostracan skeleton and, through histological comparison to other skeletonising vertebrate clades, deduced the ancestral nature of the vertebrate dermal skeleton. Heterostracans primitively possess a four‐layered skeleton, comprising a superficial layer of odontodes composed of dentine and enameloid; a compact layer of acellular parallel‐fibred bone containing a network of vascular canals that supply the pulp canals (L1); a trabecular layer consisting of intersecting radial walls composed of acellular parallel‐fibred bone, showing osteon‐like development (L2); and a basal layer of isopedin (L3). A three layered skeleton, equivalent to the superficial layer L2 and L3 and composed of enameloid, dentine and acellular bone, is possessed by the ancestor of heterostracans + jawed vertebrates. We conclude that an osteogenic component is plesiomorphic with respect to the vertebrate dermal skeleton. Consequently, we interpret the dermal skeleton of denticles in chondrichthyans and jawless thelodonts as independently and secondarily simplified. J. Morphol. 276:657–680, 2015. © 2015 The Authors Journal of Morphology Published by Wiley Periodicals, Inc.     

Ultrastructure of the flagellar apparatus inPleurochrysis (classPrymnesiophyceae)

Summary The ultrastructure of the flagellar apparatus of aPleurochrysis, a coccolithophorid was studied in detail. Three major fibrous connecting bands and several accessory fibrous bands link the basal bodies, haptonema and microtubular flagellar roots. The asymmetrical flagellar root system is composed of three different microtubular roots (referred to here as roots 1,2, and 3) and a fibrous root. Root 1, associated with one of the basal bodies, is of the compound type, constructed of two sets of microtubules,viz. a broad sheet consisting of up to twenty closely aligned microtubules, and a secondary bundle made up of 100–200 microtubules which arises at right angles to the former. A thin electron-dense plate occurs on the surface of the microtubular sheet opposite the secondary bundle. The fibrous root arises from the same basal body and passes along the plasmalemma together with the microtubular sheet of root 1. Root 2 is also of the compound type and arises from one of the major connecting bands (called a distal band) as a four-stranded microtubular root and extends in the opposite direction to the haptonema. From this stranded root a secondary bundle of microtubules arises at approximately right angle. Root 3 is a more simple type, composed of at least six microtubules which are associated with the basal body. The flagellar transition region was found to be unusual for the classPrymnesiophyceae. The phylogenetic significance of the flagellar apparatus in thePrymnesiophyceae is discussed.     

Structural peculiarities of the tubercles in the skin of the turbot,Scophthalmus maximus (L., 1758) (Osteichthyes,Pleuronectiformes, Scophthalmidae)

The structure of the bony tubercles of the turbot, Scophthalmus maximus (L., 1758), was examined using ground sections, microradiography, SEM, and TEM. The tubercles are small, isolated, mineralized conical plates randomly distributed in the eyed side of the body. They are composed of three layers: the outer limiting layer, the external layer, and the basal plate, which make up the thin and flat elasmoid scales of Teleostei. The main difference between regular elasmoid scales and bony tubercles lies in the organization and the growth of the basal plate. Indeed, the conical shape of the tubercle is the result of a prominent thickening of the central part of the basal plate where the collagen matrix is organized in a complicated three-dimensional network. Densely packed thick collagen fibrils form superimposed plies organized in a plywood-like structure that resembles that of the elasmoid scales but it is criss-crossed by numerous vertical sheets of thin collagen fibrils. The tubercles originate from thin and flat plates located in the skin of larvae and juveniles, whose structure is that of regular-developing elasmoid scales. Thus, the tubercles of Scophthalmus maximus could be considered as modified elasmoid scales rather than bony structures. They might be the result of specific arrangements related to the general trend of reduction of the dermal skeleton in the teleostean lineage.     

Structure and formation of ankylosis in Xenopus laevis

The structure of ankylotic teeth in Xenopus laevis was studied by light, transmission, and scanning electron microscopy as well as by microradiography in decalcified and undecalcified specimens. The mature teeth of Xenopus laevis are calcified from the crown to the base, fused to the jaw bone, and have no uncalcified area, such as a fibrous ring separating the tooth into the crown and pedicle. Microradiography shows that the mature tooth and jaw bone appear as an X-ray opaque area, except for the basal region of the dentine. This region is composed of an X-ray translucent area and an X-ray opaque thin layer on the lingual side of the translucent area. The mature tooth is composed of two differently calcified areas: (1) a highly calcified area, which makes up almost all of the tooth and contains a thin layer of the basal dentine on the lingual side, and (2) a lowly calcified basal dentine, which is fused to the jaw bone. Therefore, the lowly calcified area does not completely separate the dentine and jaw bone. Repeating banding patterns among the collagen fibrils differ among the dentine-forming area and the matrices of dentine and jaw bone. During the formation of ankylosis of the tooth germ, collagen bundles in the dentine-forming area accumulate directly on the surface of the jaw bone. Consequently, the mature teeth of Xenopus laevis fuse to the jaw bone directly without the mediation of the other structures.     

Conodont affinity of the enigmatic Carboniferous chordate Conopiscius

Conopiscius shares V-shaped myomeres with the co-occurring conodont Clydagnathus but instead of a complex oral apparatus it has only a single pair of conical elements, and structures resembling scales are associated with its myomeres. Moreover, the coarsely crystalline crown tissue typical for conodonts has not been identified in the Conopiscius elements, which show only a finely lamellar skeletal tissue. The gap between conodonts and Conopiscius may be filled by isolated elements of similar morphology and structure occurring in the Late Devonian. They reveal a very thin external layer developed mostly at the tooth tip and resembling conodont crown tissue. The pulp cavity is partially filled with layered or spherulitic phosphatic tissue of the kind known also in conodonts (basal filling tissue) and early vertebrates (lamellin). Conodont elements of similar morphology and representing uni-membrate oral apparatuses have not been previously reported from the Devonian or Carboniferous but occur near the Cambrian–Ordovician transition ( Proconodontus ) and in the Late Permian ( Caenodontus ). It is proposed that Conopiscius represents a mostly cryptic conodont lineage extending from the Early Ordovician to the Permian, instead of being directly related to the agnathans.     

Seed structure and histochemistry in the palm Euterpe edulis

The Euterpe edulis embryo consists of a prominent single cotyledon, a very short radicle-hypocotyl axis and an epicotyl. The epicotyl is obliquely angled with respect to the cotyledon; consequently it corresponds to one of the two categories recognized for palm seeds by DeMason (1988 ). Parenchyma, protoderm and procambium can be distinguished on the basis of position and shape of their cells, which are highly vacuolated with one central vacuole and the cytoplasm restricted to a thin parietal layer. Initial cells from both apical meristems are also vacuolated but they have small vacuoles distributed around the nuclei. Silica occurs in cell walls of some protodermal cells. Raphides, silica bodies and tannins all occur occasionally in vacuoles, especially in the basal cotyledon region. Most embryo cells lack storage reserves and exhibit an active state, with numerous mitochondria, RER cisternae and Golgi apparatus, indicating a strategy of continuous development without the interposition, at maturity, of a dry state. The endosperm consists of living cells with very large nuclei and thickened cell walls. Similar to the endosperm of other studied palm species, their cells exhibit a quiescent appearance with lipid, protein, minerals (in the cytoplasm) and mannans (in the cell walls) as the insoluble storage reserves.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 445–453.     

The ultrastructure of corpus allatum of the earwig Euborellia annulipes lucas (Dermaptera: Labiduridae)

The ultrastructure of corpus allatum of the earwig, Euborellia annulipes has been described. The corpus allatum (CA) is an oval body. The gland is covered by a thin stromal sheath which is wavey and acellular in composition. The gland consists of either parenchymal cells with distinct, double-layered, smooth or slightly undulated plasma-membranes. Golgi bodies and endoplasmic reticulum are poorly developed. The mitochondria are found in abundance and are oval, spherical, elongate, and Y-shaped, with oblique and longitudinal cristae. The gland is innervated by neurosecretory and non-neurosecretory axons. The neurosecretory axons are of two types. The type NS-II, contain electron dense granules of 40...120 nm in diameter and the type NS-I, small less electron dense granules (of mixed nature) 40...90 nm in diameter. Branches of tracheoles also occur that penetrate the gland. The large electron dense granules 100...220 nm in diameter occuring abundantly in aorta, and representing the secretions of the medial A-cells are absent in the CA.     

Les relations isopedine - tissu osseux dans le post-temporal et les ecailJes de la ligne laterale de Latimeria chalumnae (Smith)

The scaly part of the posttemporal bone and the lateral-line scales have the same structure as other scales. However, the lateral line with its bony canal induces some local transformations in both structures. The posttemporal bone can be interpreted as a lateral-line scale whose "dermo"-component is extensively developed. With its composite structure, both osseous and scaly, the posttemporal bone affords an instance peculiarly well suited for studying the relationships between isopedine and bone. The observations demonstrate that isopedine whose superposed collagen layers make a double-twisted plywood, is a specialized structure which evolved from a bony tissue. The fibrous basal plate of a Latimeria scale is homologous to the osseous basal plate of a cosmoid scale.     

Ausgangsform und Entwicklung der rhombischen Schuppen der Osteichthyes (Pisces)

Ganoid and cosmoid scales, the two types of rhombic scales within the osteichthyans, can be traced back to a primitive scale similar to the scales ofLophosteus. The primitive rhombic scale did not have a peg-and-socket articulation, it is composed of lamellar bone superposed by many layers of spongy bone + dentine. That kind of superposition of layers of spongy bone + dentine (+ enamel) has been retained in the cosmoid scale; in contrast in the ganoid scale the growth of the dentine has become restricted to the lateral surface, the growth of ganoin to the outer surface and the growth of bone to the inner surface of the scale. The scales ofAndreolepis have a position between the primitive rhombic scale and the ganoid scale. — Scales from the Gedinnian (Lower Devonian) of New Sibirian Islands, USSR, are described asDialipina markae n. sp. The morphological features are very typical for the genus, but the histology is different from the type speciesD. salgueiroensis. Within the two Devonian palaeoniscoid generaDialipina andOrvikuina acellular bone with irregular non-vascular canals of Williamson has developed twice from cellular bone.     

Structure of the accessory reproductive glands of the male migratory grasshopper,Melanoplus sanguinipes

The accessory reproductive glands of Melanoplus sanguinipes comprise two bilateral masses of 16 tubules each, distinguishable in sexually mature insects as four white, ten short hyaline, one long hyaline, and a seminal vesicle. Over most of its length, the wall of each tubule consists of a simple glandular epithelium resting on a basal lamina, surrounded by a thin layer of circular muscle. However, near the junction with the ejaculatory duct, the wall of each tubule has a much thickened circular muscle layer and squamous or cuboidal epithelium, the region serving to regulate movement of secretion into the ejaculatory duct. Interdigitation of adjacent epithelial cells is common, and several kinds of specialized junctions occur. In the glandular region, all epithelial cells appear the same and may be flattened, cuboidal, or columnar depending on the tubule type. Except for those of the seminal vesicle, the glandular epithelial cells share ultrastructural features typical of cells engaged in the synthesis of protein for export. Despite these general similarities, in most instances subtle differences occur in the cellular ultrastructure of the epithelia of each tubule and in the appearance of their luminal secretions, suggesting that the tubules are functionally specialized.     

Phylogenetic position and morphology of Raphidiophrys elongata sp. nov. (Haptista: Centroplasthelida) with notes on cyst wall structure and evolution

The majority of centrohelids bear external coverings consisting of organic spicules or siliceous scales. Cyst coverings are usually reinforced with additional layers of modified scales. The cyst wall of Raphidiophrys heterophryoidea has an unusual and complex structure. It consists of three different types of scales and includes the mosaic scale layer not known in other centrohelids. During excystment, the cyst wall fragments along the sutures of the mosaic layer. For other Raphidiophrys species, cyst coverings are not studied. The present paper describes a new Raphidiophrys species, R. elongata, belonging to the NC7 environmental clade. Trophozoites bore thin plate scales with reduced upper plate. Under starvation, cysts emerged in clonal cultures. Cyst coverings of R. elongata and R. heterophryoidea were studied in comparison with the use of FIB-SEM. Cyst wall of R. elongata was significantly thinner than in R. heterophryoidea and was formed with 3–5 layers of uniform overlapping scales. No mosaic scale layer was present. During excystment, trophozoite exited cyst shell through random fissure. Possible evolutionary events and driving forces behind the complication of cyst wall within Raphidiophrys were discussed.