Processes of scale formation in fish and Agnatha

The formation of the scale cover in ancient fish and Agnatha was analyzed using paleontology data on some forms and by studying recent species according to the geochronological principle and morphological-genetic coupling approach. The histogenesis of true scale cover was transformed simultaneously but independently from dermal denticles, when the first process was accompanied by reinforcement of the bones and formation of head-body armor and dermal-like external elements of the skullcap. The scale cover is an independent organ system and is characterized by self-supporting conservative ancestral mechanisms of development.     

Formation of dermal skeletal and dental tissues in fish: a comparative and evolutionary approach

Osteichthyan and chondrichthyan fish present an astonishing diversity of skeletal and dental tissues that are often difficult to classify into the standard textbook categories of bone, cartilage, dentine and enamel. To address the question of how the tissues of the dermal skeleton evolved from the ancestral situation and gave rise to the diversity actually encountered, we review previous data on the development of a number of dermal skeletal elements (odontodes, teeth and dermal denticles, cranial dermal bones, postcranial dermal plates and scutes, elasmoid and ganoid scales, and fin rays). A comparison of developmental stages at the tissue level usually allows us to identify skeletogenic cell populations as either odontogenic or osteogenic on the basis of the place of formation of their dermal papillae and of the way of deposition of their tissues. Our studies support the evolutionary affinities (1) between odontodes, teeth and denticles, (2) between the ganoid scales of polypterids and the elasmoid scales of teleosts, and (3) to a lesser degree between the different bony elements. There is now ample evidence to ascertain that the tissues of the elasmoid scale are derived from dental and not from bony tissues. This review demonstrates the advantage that can be taken from developmental studies, at the tissue level, to infer evolutionary relationships within the dermal skeleton in chondrichthyans and osteichthyans.     

Cell polarity: intrinsic or externally imposed?

A basic question in studies of the genesis of cell polarity is whether the polarity is an intrinsic and permanent property of cells or whether it is externally imposed by signals at the cell periphery. Current models favor the possibility that an external signal selectively imposes a polarized cell morphology. However, recent data from different experimental systems are discussed here that support the idea that an intrinsic polarity in animal cells is maintained through a dynamic process involving specific activities of the cortical microfilament system and the centrosome-microtubule complex. In this view, external signals capable of modulating cell polarity, for example, during chemotaxis or histogenesis, do so by acting on mechanisms that maintain cells permanently polarized. The contribution of the cytoskeleton to the genesis of cell polarity is discussed, with particular reference to experimental evidence for global cytoskeletal dynamics, and it is suggested that critical advances in our understanding of the maintenance of cell polarity will depend on our obtaining further knowledge of the molecular mechanisms controlling interactions between microtubules and microfilaments. Microtubules appear to exert an inhibitory control on the recruitment of cytoplasmic myosin into the cortex, and there are data indicating that the centrosome and centrioles could actively contribute to the establishment of cell polarity.     

Histology of “placoderm” dermal skeletons: Implications for the nature of the ancestral gnathostome

The vertebrate dermal skeleton has long been interpreted to have evolved from a primitive condition exemplified by chondrichthyans. However, chondrichthyans and osteichthyans evolved from an ancestral gnathostome stem‐lineage in which the dermal skeleton was more extensively developed. To elucidate the histology and skeletal structure of the gnathostome crown‐ancestor we conducted a histological survey of the diversity of the dermal skeleton among the placoderms, a diverse clade or grade of early jawed vertebrates. The dermal skeleton of all placoderms is composed largely of a cancellar architecture of cellular dermal bone, surmounted by dermal tubercles in the most ancestral clades, including antiarchs. Acanthothoracids retain an ancestral condition for the dermal skeleton, and we record its secondary reduction in antiarchs. We also find that mechanisms for remodeling bone and facilitating different growth rates between adjoining plates are widespread throughout the placoderms. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Analysis of morphogenesis and keratinization in transfilter recombinants of feather-forming skin

The relationships between feather morphogenesis, histogenesis, and biochemical differentiation were examined by recombining backskin epidermis and dermis, from chick embryos (Hamburger-Hamilton stages 27-31), with an intervening Nucleopore filter (pore size of 0.4 micron). The filter inhibited normal feather morphogenesis and histogenesis of barb ridges, yet feather-like filaments, which were free of dermal cells, formed from the epidermal cells. Using indirect immunofluorescence, with antiserum against alpha- and beta-keratins, the biochemical differentiation of the feather-like filaments was compared to normal feathers. In the feather-like filaments resulting from tissues of stages 27-29, cells containing beta keratins were occasionally seen at the periphery of the filaments, yet cells containing alpha-keratins were inappropriately located throughout the filaments. In a few feather-like filaments on recombinants resulting from tissues of stages 29.5-31, cells positive for beta-keratins were found in the center of the filament, but again alpha-keratins were also found. Surrounding these cells there were several layers of cells, arranged circumferentially, resembling sheath cells. Some sheath-like cells contained beta-keratins. We conclude that although feather epidermal cells, which are separated from their dermis by a Nuclepore filter, can undergo limited morphogenesis and the production of alpha- and beta-keratins, normal feather morphogenesis, histogenesis, and biochemical differentiation require the intimate associations of epidermis and dermis.     

Models of epithelial histogenesis

Epithelial tissues emerge from coordinated sequences of cell renewal, specialization and assembly. Like corresponding immature tissues, adult epithelial tissues are provided by stem cells which are responsible for tissue homeostasis. Advances in epithelial histogenesis has permitted to clarify several aspects related to stem cell identification and dynamics and to understand how stem cells interact with their environment, the so-called stem cell niche. The development and maintenance of epithelial tissues involves epithelial-mesenchymal signalling pathways and cell-matrix interactions which control target nuclear factors and genes. The tooth germ is a prototype for such inductive tissue interactions and provides a powerful experimental system for the study of genetic pathways during development. Clonogenic epithelial cells isolated from developing as well mature epithelial tissues has been used to engineer epithelial tissue-equivalents, e.g. epidermal constructs, that are used in clinical practise and biomedical research. Information on molecular mechanisms which regulate epithelial histogenesis, including the role of specific growth/differentiation factors and cognate receptors, is essential to improve epithelial tissue engineering.     

The histologists of the S. M. Kirov Military Medical Academy--their contribution to practical medicine]

This is a short review of investigations of an applied character, performed at the Department of Histology and Embryology of the Academy. Most of the investigations have been carried out as joint works with clinicians and are aimed to solve some actual tasks of practical medicine. Their main contents have been investigations on reactivity and degeneration of tissues under various experimental and clinical conditions. Some new data have been obtained concerning cellular and ultrastructural mechanisms of the wound process, scientifically grounded influences have been undertaken in order to regulate this process. The material presented has served as a base for elaboration of a number of fundamental principles of histology and first of all some new ideas on histogenesis, cellular-differon organization and reparative regeneration of tissues. Regularities of histo- and organogenesis and intertissue interrelation are the base for understanding regeneration mechanisms of organs and tissues. In what form the reparative regeneration is manifested depends on histogenesis and is specific for each tissue. To know regularities of histogenesis and peculiarities of its manifestation is necessary to understand problems of pathology, since regularities of the normal histogenesis serve as a foundation for understanding the essence of pathological processes.     

Evolution of the cytoskeleton

The eukaryotic cytoskeleton appears to have evolved from ancestral precursors related to prokaryotic FtsZ and MreB. FtsZ and MreB show 40-50% sequence identity across different bacterial and archaeal species. Here I suggest that this represents the limit of divergence that is consistent with maintaining their functions for cytokinesis and cell shape. Previous analyses have noted that tubulin and actin are highly conserved across eukaryotic species, but so divergent from their prokaryotic relatives as to be hardly recognizable from sequence comparisons. One suggestion for this extreme divergence of tubulin and actin is that it occurred as they evolved very different functions from FtsZ and MreB. I will present new arguments favoring this suggestion, and speculate on pathways. Moreover, the extreme conservation of tubulin and actin across eukaryotic species is not due to an intrinsic lack of variability, but is attributed to their acquisition of elaborate mechanisms for assembly dynamics and their interactions with multiple motor and binding proteins. A new structure-based sequence alignment identifies amino acids that are conserved from FtsZ to tubulins. The highly conserved amino acids are not those forming the subunit core or protofilament interface, but those involved in binding and hydrolysis of GTP.     

Moving into shape: cell migration during the development and histogenesis of the cerebellum

The enormous expansion the vertebrate nervous system goes through from its first anlage to its adult shape and organization goes along with extensive rearrangements of its constituent cells and typical cellular migrations, often over long distances, and by convoluted pathways. Here, I try to summarize how the cells that form the cerebellum move and migrate during normal cerebellar histogenesis. The cerebellum is made up of a limited set of clearly distinguishable classes of cells, some of which are also readily accessible by genetic tools. Its structure and development have been the focus of studies dating back to at least Ramon y Cajal which have yielded fundamental insights into basic mechanisms of the development of the nervous. During cerebellar histogenesis, several distinct and well-discernable modes of migration may be recognized, some of which have been studied in considerable morphological and molecular detail. Still, often grace to the detail known, a wealth of open questions remains, and the cerebellar anlage remains a highly accessible and promising paradigm for those interested in nervous system development and cell migration in general. I also point out some of the issues that may warrant consideration when results from technically distinct studies are compared and integrated.     

Expression and role of E- and P-cadherin adhesion molecules in embryonic histogenesis. II. Skin morphogenesis

Expression and the role of E- and P-cadherin in the histogenesis of the surface epidermis and hair follicles were examined using the upper lip skin of the mouse. P-cadherin is expressed exclusively in the proliferating region of these tissues, that is in the germinative layer of the surface epidermis, the outer root sheath and the hair matrix. E-cadherin is coexpressed in these layers but this molecule was also detected in non-proliferating regions such as the intermediate layer of the surface epidermis and the immature regions of the inner root sheath. Neither P- nor E-cadherin was detected in fully keratinized layers such as the horny layer of the surface epidermis, the outermost layer of the outer root sheath and the mature hair fibres. These two cadherins were not detected in dermal cells. We cultured pieces of the upper lip skin in vitro in the absence or presence of a monoclonal antibody to E-cadherin (ECCD-1) or to P-cadherin (PCD-1). In control cultures, skin morphogenesis normally occurred in a pattern whereby the hair follicles grew and dermal cells were condensed to form the dermal sheath. A mixture of ECCD-1 and PCD-1, however, induced abnormal morphogenesis in the skin in several respects. (1) The cuboidal or columnar arrangement of basal epithelial cells was distorted. (2) Hair follicles were deformed. (3) Condensation of dermal cells was suppressed, causing a homogeneous distribution of these cells. These results suggest that cadherins present in epidermal cells are involved not only in maintaining the arrangement of these cells but also in inducing dermal condensation.     

Differences in the histogenesis and keratin expression of avian extraembryonic ectoderm and endoderm recombined with dermis

The responses of the chorionic ectoderm and allantoic endoderm (from 8-day chick embryos) to dermal induction were compared through tissue recombinants grafted onto the chorioallantoic membrane. The chorionic epithelium formed the appropriate epidermis with a fully developed stratum corneum in response to both spur and scutate scale dermises. Analysis of these recombinant epidermal tissues by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE) demonstrated that tissue-specific expression of the alpha (alpha) and beta (beta) keratin polypeptides occurred. In addition, indirect immunofluorescence studies with antisera to alpha or beta keratins showed that the beta stratum, which characterizes the epidermis of spurs and scutate scales, was formed, and the alpha keratins were distributed as in the normal epidermal tissues. In contrast, although the allantoic endoderm became stratified in association with either spur or scutate scale dermis, a stratum corneum with a beta stratum did not develop. SDS-PAGE analysis demonstrated that while the characteristic beta keratins of scutate scales and spur were not detected, most of the alpha keratins normally elaborated by these structures were present, suggesting that even without histogenesis of a stratum corneum the expression of alpha keratins of endoderm could be regulated in a tissue-specific manner by dermis. This study also demonstrated that there are differences in the abilities of the chorionic and allantoic epithelia to respond to the same dermal cues, which may reflect earlier restrictions in their developmental potentials.     

Voltage clamp analysis of two inward current mechanisms in the egg cell membrane of a starfish

Ionic mechanisms of excitation were studied in the immature egg cell membrane of a starfish, Mediaster aequalis, by analyzing membrane currents during voltage clamp. The cell membrane shows two different inward current mechanisms. One is activated at a membrane potential of -55 approximately -50 mV and the other at -7 approximately -6 mV. They are referred to as channels I and II, respectively. A similar difference is also found in the membrane potential of half inactivation. Currents of the two channels can, therefore, be separated by selective inactivation. The currents of both channels depend on Ca++ (Sr++ or Ba++) but only the current of channel I depends on Na+. The time-course of current differs significantly between the two channels when compared at the same membrane potential. The relationship between the membrane current and the concentration of the permeant ions is also different between the two channels. The result suggests that channel II is a more saturable system. The sensitivity of the current to blocking cations such as Co++ or Mg++ is substantially greater in channel II than in channel I. Currents of both channels depend on the external pH with an apparent pK of 5.6. They are insensitive to 3 muM tetrodotoxin (TTX) but are eliminated totally by 7.3 mM procaine. The properties of channel II are similar to those of the Ca channel found in various adult tissues. The properties of channel I differ, however, from those of either the typical Ca or Na channels. Although the current of the channel depends on the external Na the amplitude of the Na current decreases not only with the Na concentration but also with the Ca concentration. No selectivity is found among Li+, Na+, Rb+, and Cs+. The experimental result suggests that Na+ does not carry current but modifies the current carried by Ca in channel I.     

Developmental genetics and evolution of symbiotic structures in nitrogen-fixing nodules and arbuscular mycorrhiza.

Genetic and molecular mechanisms of development are compared for two major plant-microbe endosymbioses: N(2)-fixing nodules (with rhizobia or actinomycetes Frankia) and arbuscular mycorrhiza (with Glomales fungi). Development from the primordia formed de novo in root tissues is common for all known types of N(2)-fixing nodules. However, their structure varies greatly with respect to: (i) tissue topology (location of vascular bundles is peripherical in legumes or central in non-legumes); (ii) position of nodule primordium (inner or outer cortex in legumes, pericycle in non-legumes); (iii) stability of apical meristem (persistent in the indeterminate nodules, transient in the determinate ones). In addition, legumes vary in ability to form compartments harboring endosymbiotic rhizobia and located intercellularly (infection threads) and intracellularly (symbiosomes). Using pea (Pisum sativum) symbiotic mutants, the nodule developmental program is dissected into a range of spatially and temporarily differentiated steps comprising four sub-programs (development of endosymbiotic compartments; nodule histogenesis; autoregulation of nodulation; bacteroid differentiation). The developmental mutations are suggested in some cases to reverse the endosymbiotic system into the morphologically simpler forms some of which may correspond to the ancestral stages of nodule evolution. The origin of legume-rhizobial and actinorhizal symbioses is suggested to be based on a set of preadaptations many of which had been evolved in angiosperms during coevolution with arbuscular mycorrhizal fungi (e.g., inter- and intracellular maintenance of symbionts, their control via defence-like reactions and recognition of chitin-like molecules). An analysis of parallel morphological variation in symbiotic mutants and wild-growing legume species enables us to reconstruct the major stages of evolution for N(2)-fixing symbioses.     

Hypothesis: cyclical histogenesis is the basis of circannual timing

Circannual rhythms are innately timed long-term (tau ≈ 12 months) cycles of physiology and behavior, crucial for life in habitats ranging from the equator to the Poles. Here the authors propose that circannual rhythm generation depends on tissue-autonomous, reiterated cycles of cell division, functional differentiation, and cell death. They see the feedback control influencing localized stem cell niches as crucial to this cyclical histogenesis hypothesis. Analogous to multi-oscillator circadian organization, circannual rhythm generation occurs in multiple tissues with hypothalamic and pituitary sites serving as central pacemakers. Signals including day length, nutrition, and social factors can synchronize circannual rhythms through hormonal influences, notably via the thyroid and glucocorticoid axes, which have profound effects on histogenesis. The authors offer 4 arguments in support of this hypothesis: (1) Cyclical histogenesis is a prevalent process in seasonal remodeling of physiology. It operates over long time domains and exhibits tissue autonomy in its regulation. (2) Experiments in which selected peripheral endocrine signals are held constant indicate that circannual rhythms are not primarily the product of interacting hormonal feedback loops. (3) Hormones known to control cell proliferation, differentiation, and organogenesis profoundly affect circannual rhythm expression. (4) The convergence point between photoperiodic input pathways and circannual rhythm expression occurs in histogenic regions of the hypothalamus and pituitary. In this review, the authors discuss how testing this hypothesis will depend on the use of cellular/molecular tools and animal models borrowed from developmental biology and neural stem cell research.     

Testing models of dental development in the earliest bony vertebrates, Andreolepis and Lophosteus

Theories on the development and evolution of teeth have long been biased by the fallacy that chondrichthyans reflect the ancestral condition for jawed vertebrates. However, correctly resolving the nature of the primitive vertebrate dentition is challenged by a dearth of evidence on dental development in primitive osteichthyans. Jaw elements from the Silurian-Devonian stem-osteichthyans Lophosteus and Andreolepis have been described to bear a dentition arranged in longitudinal rows and vertical files, reminiscent of a pattern of successional development. We tested this inference, using synchrotron radiation X-ray tomographic microscopy (SRXTM) to reveal the pattern of skeletal development preserved in the sclerochronology of the mineralized tissues. The tooth-like tubercles represent focal elaborations of dentine within otherwise continuous sheets of the dermal skeleton, present in at least three stacked generations. Thus, the tubercles are not discrete modular teeth and their arrangement into rows and files is a feature of the dermal ornamentation that does not reflect a polarity of development or linear succession. These fossil remains have no bearing on the nature of the dentition in osteichthyans and, indeed, our results raise questions concerning the homologies of these bones and the phylogenetic classification of Andreolepis and Lophosteus.     

Evolutionary genetics of vertebrate tissue mineralization: the origin and evolution of the secretory calcium-binding phosphoprotein family

Three principal mineralized tissues are present in teeth; a highly mineralized surface layer (enamel or enameloid), body dentin, and basal bone. Similar tissues have been identified in the dermal skeleton of Paleozoic jawless vertebrates, suggesting their ancient origin. These dental tissues form on protein matrix and their mineralization is controlled by distinctive proteins. We have shown that many secretory calcium-binding phosphoproteins (SCPPs) are involved in tetrapod tissue mineralization. These SCPPs all originated from the common ancestral gene SPARCL1 (secreted protein, acidic, cysteine-rich like 1) that initially arose from SPARC. The SCPP family also includes a bird eggshell matrix protein, mammalian milk casein, and salivary proteins. The eggshell SCPP plays crucial roles in rigid eggshell production, milk SCPPs in efficient lactation and in the evolution of complex dentition, and salivary SCPPs in maintaining tooth integrity. A comparative analysis of the mammalian, avian, and amphibian genomes revealed a tandem duplication history of the SCPP genes in tetrapods. Although these tetrapod SCPP genes are fewer in teleost genomes, independent parallel duplication has created distinct SCPP genes in this lineage. These teleost SCPPs are also used for enameloid and dentin mineralization, implying essential roles of SCPPs for dental tissue mineralization in osteichthyans. However, the SCPPs used for tetrapod enamel and teleost enameloid, as well as tetrapod dentin and teleost dentin, are all different. Thus, the evolution of vertebrate mineralized tissues seems to be explained by phenogenetic drift: while mineralized tissues are retained during vertebrate evolution, the underlying genetic basis has extensively drifted.     

The Developmental Basis of Skeletal Cell Differentiation and the Molecular Basis of Major Skeletal Defects

Vertebrate skeletal differentiation retains elements from simpler phyla, and reflects the differentiation of supporting tissues programmed by primary embryonic development. This developmental scheme is driven by homeotic genes expressed in sequence, with subdivision of skeletal primordia driven by a combination of seven transmembrane‐pass receptors responding to Wnt‐family signals, and by bone morphogenetic family signals that define borders of individual bones. In sea‐dwelling vertebrates, an essentially complete form of the skeleton adapted by the land‐living vertebrates develops in cartilage, based on type II collagen and hydrophilic proteoglycans. In bony fishes, this skeleton is mineralized to form a solid bony skeleton. In the land‐living vertebrates, most of the skeleton is replaced by an advanced vascular mineralized skeleton based on type I collagen, which reduces skeletal mass while facilitating use of skeletal mineral for metabolic homeostasis. Regulation of the mammalian skeleton, in this context, reflects practical adaptations to the needs for life on land that are related to ancestral developmental signals. This regulation includes central nervous system regulation that integrates bone turnover with overall metabolism. Recent work on skeletal development, in addition, demonstrates molecular mechanisms that cause developmental bone diseases.     

Scales and Dermal Skeletal Histology of an Early Bony Fish Psarolepis romeri and Their Bearing on the Evolution of Rhombic Scales and Hard Tissues

Recent discoveries of early bony fishes from the Silurian and earliest Devonian of South China (e.g. Psarolepis, Achoania, Meemannia, Styloichthys and Guiyu) have been crucial in understanding the origin and early diversification of the osteichthyans (bony fishes and tetrapods). All these early fishes, except Guiyu, have their dermal skeletal surface punctured by relatively large pore openings. However, among these early fishes little is known about scale morphology and dermal skeletal histology. Here we report new data about the scales and dermal skeletal histology of Psarolepis romeri, a taxon with important implications for studying the phylogeny of early gnathostomes and early osteichthyans. Seven subtypes of rhombic scales with similar histological composition and surface sculpture are referred to Psarolepis romeri. They are generally thick and show a faint antero-dorsal process and a broad peg-and-socket structure. In contrast to previously reported rhombic scales of osteichthyans, these scales bear a neck between crown and base as in acanthodian scales. Histologically, the crown is composed of several generations of odontodes and an irregular canal system connecting cylindrical pore cavities. Younger odontodes are deposited on older ones both superpositionally and areally. The bony tissues forming the keel of the scale are shown to be lamellar bone with plywood-like structure, whereas the other parts of the base are composed of pseudo-lamellar bone with parallel collagen fibers. The unique tissue combination in the keel (i.e., extrinsic Sharpey''s fibers orthogonal to the intrinsic orthogonal sets of collagen fibers) has rarely been reported in the keel of other rhombic scales. The new data provide insights into the early evolution of rhombic (ganoid and cosmoid) scales in osteichthyans, and add to our knowledge of hard tissues of early vertebrates.