The junction zone: Initial site of mineralization in radula teeth of the chiton Cryptoplax striata (Mollusca: Polyplacophora)

Elemental composition and distribution in individual teeth of the whole radula of the chiton Cryptoplax striata were analyzed using energy-dispersive spectroscopy. Both the element deposited and its position within the tooth vary according to the stage of mineralization. The initial site of mineralization is the junction zone, the region between the tooth cusp and base. In this region, the first element to be deposited is iron, followed by phosphorus and then calcium. Iron deposition next commences in the tooth cusp cap, where it proceeds rapidly, being virtually complete within 12 tooth rows. By contrast, mineralization in the core of the tooth cusp does not commence until well down the radula and consists initially of iron and phosphorus with the addition of a small amount of calcium 6 rows later. While mineralization in the tooth base commences early in radula development, it continues right through to the fully mature end of the radula. A number of minor elements are also found at various stages of mineralization. The data obtained have been used to construct a schematic of the progression of mineralization along the length of the radula. © 1996 Wiley-Liss, Inc.     

Structural Organisation of the Cusps of the Radular Teeth of the Chiton Plaxiphora albida

Abstract The structure, morphology and organisation of the cusps of the major lateral radula teeth of the chiton Plaxiphora albida have been examined using light, transmission and scanning electron microscopy, together with energy dispersive X-ray analysis and Mössbauer spectroscopy. In this chiton species, both the anterior and posterior surfaces of the major lateral teeth are composed of magnetite, which is indicated to be non-stoichiometric and associated with some maghemite, together with small amounts of phosphorus and silicon. This outer layer surrounds an inner core region of the tooth, which only reaches the surface through a small window zone on the anterior surface and which contains large amounts of iron and phosphorus presumably in the form of iron(III) phosphate. The organic matrix, on which the teeth are constructed, consists of a zone of densely packed fine fibres at the surface of the tooth, underlain by larger fibres which become sparser deeper into the cusp. The core region is characterized by the presence of densely packed short fibres. In contrast to the situation found in most other species of chiton, large fibres of the organic matrix extend throughout the region of magnetite mineralization, leading to the suggestion that the matrix exerts more control over the mineralization of magnetite than has previously been thought.     

红条毛肤石鳖齿舌主侧齿中铁还原酶分布及与生物矿化的关系

以红条毛肤石鳖Acanthochiton rubrolineatus(Lischke)齿舌为材料,通过切片和酶组织化学技术,在光镜和电镜下对齿舌主侧齿的微结构及高铁还原酶的存在进行观察,从微观角度了解齿舌主侧齿齿尖的矿化机理。结果显示,成熟主侧齿由齿尖和齿基组成。齿尖结构由外至内分为三层,最外层为磁铁矿层,前后齿面磁铁矿层的厚度不等,后齿面约50μm,前齿面约5-10μm。向内依次为棕红色的纤铁矿层,厚约10μm,及略显黄色的有机基质层,有机基质层占据着齿尖内部的大部分结构。高分辨透射电镜下显示磁铁矿由条状四氧化三铁颗粒组成,长约2-3μm,宽约100-150nm。齿舌的矿化是一个连续过程,不同部段处于不同的矿化阶段,齿舌囊上皮细胞沿囊腔分布,并形成齿片。未矿化的新生主侧齿齿尖中存在由有机基质构成的网状结构。随矿化的进行,有机基质内出现矿物颗粒。初始矿化的齿尖外表面有一个细胞微突层,微突的另一端为囊上皮细胞,矿物质经由微突层达齿尖并沉积于有机基质中,齿尖随之矿化并成熟。初始矿化齿尖的外围有大量的三价铁化物颗粒,稍成熟的齿尖外围同时还出现二价铁化物。新生或初始矿化主侧齿齿尖外围的囊上皮细胞中有大量球形类似于铁蛋白聚集体的内容物,直径0.6-0.8μm,球体由膜包围。齿舌囊上皮组织中存在三价高铁还原酶,此酶分布于上皮细胞的膜表面,可能与齿尖表面磁铁矿的生成有一定的关系。       

A new biomineral identified in the cores of teeth from the chiton <Emphasis Type="Italic"> Plaxiphora albida</Emphasis>

The hydrated iron(III) oxide limonite is reported for the first time as a biomineral. In situ laser Raman spectra of the tooth cores from major lateral teeth of the chiton Plaxiphora albida are compared with those of synthetic and mineral iron phosphates and iron oxides. Raman spectra measured on iron phosphate and iron oxide standard materials are shown to be easily distinguishable from one another. The central tooth cores of mature P. albida teeth do not show any evidence for the presence of a separate iron phosphate mineral. Rather, in each tooth a narrow band of the hydrated iron(III) oxide limonite is shown to separate the magnetite of the tooth surface from a central core region comprising both lepidocrocite and limonite. The high concentration of phosphorus in P. albida tooth cores, previously observed by energy dispersive spectroscopy, is not associated with a separate iron phosphate mineral, indicating that this element may be adsorbed onto the surface of the iron oxide minerals present. The failure to detect a separate iron(III) phosphate is discussed with reference to other chiton species that display high levels of iron and phosphorus in the cores of their mature major lateral teeth.     

Characterization of the calcium biomineral in the radular teeth of Chiton pelliserpentis

The radula in a group of molluscan invertebrates, the chitons (Polyplacophora), is a ribbon-like apparatus used for feeding and which bears a series of distinctive mineralized teeth called the major lateral teeth. While some chiton species deposit only iron biominerals in these teeth, many others deposit both iron and calcium. In this study, the calcium biomineral in the teeth of one of the latter types of species, the Australian east-coast chiton, Chiton pelliserpentis, has been isolated and examined for the first time. Spectroscopic and crystallographic techniques have identified the biomineral as a carbonate-substituted apatite with significant fluoride substitution also likely. Fourier-transform infrared and laser Raman spectroscopy indicated that the carbonate content was less than that of either bovine tibia cortical bone or human tooth enamel. X-ray diffraction analysis showed the biomineral to be poorly crystalline due to small crystal size and appreciable anionic substitution. The lattice parameters were calculated to be a=9.382?Å and c=6.883?Å, which are suggestive of a fluorapatite material. It is postulated that structural and biochemical differences in the tooth organic matrix of different chiton species will ultimately determine if the teeth become partly calcified or iron mineralized only.     

Fine structure of the mineralized teeth of the chiton Acanthopleura echinata (Mollusca: Polyplacophora)

The major lateral teeth of the chiton Acanthopleura echinata are composite structures composed of three distinct mineral zones: a posterior layer of magnetite; a thin band of lepidocrocite just anterior to this; and apatite throughout the core and anterior regions of the cusp. Biomineralization in these teeth is a matrix-mediated process, in which the minerals are deposited around fibers, with the different biominerals described as occupying architecturally discrete compartments. In this study, a range of scanning electron microscopes was utilized to undertake a detailed in situ investigation of the fine structure of the major lateral teeth. The arrangement of the organic and biomineral components of the tooth is similar throughout the three zones, having no discrete borders between them, and with crystallites of each mineral phase extending into the adjacent mineral zone. Along the posterior surface of the tooth, the organic fibers are arranged in a series of fine parallel lines, but just within the periphery their appearance takes on a "fish scale"-like pattern, reflective of the cross section of a series of units that are overlaid, and offset from each other, in adjacent rows. The units are approximately 2 microm wide and 0.6 microm thick and comprise biomineral plates separated by organic fibers. Two types of subunits make up each "fish scale": one is elongate and curved and forms a trough, in which the other, rod-like unit, is nestled. Adjacent rod and trough units are aligned into large sheets that define the fracture plane of the tooth. The alignment of the plates of rod-trough units is complex and exhibits extreme spatial variation within the tooth cusp. Close to the posterior surface the plates are essentially horizontal and lie in a lateromedial plane, while anteriorly they are almost vertical and lie in the posteroanterior plane. An understanding of the fine structure of the mineralized teeth of chitons, and of the relationship between the organic and mineral components, provides a new insight into biomineralization mechanisms and controls.     

Atomic force microscopy study of tooth surfaces

Atomic force microscopy (AFM) was used to study tooth surfaces in order to compare the pattern of particle distribution in the outermost layer of the tooth surfaces. Human teeth and teeth from a rodent (Golden hamster), from a fish (piranha), and from a grazing mollusk (chiton) with distinct feeding habits were analyzed in terms of particle arrangement, packing, and size distribution. Scanning electron microscopy and transmission electron microscopy were used for comparison. It was found that AFM gives high-contrast, high-resolution images and is an important tool as a source of complementary and/or new structural information. All teeth were cleaned and some were etched with acidic solutions before analysis. It was observed that human enamel (permanent teeth) presents particles tightly packed in the outer surface, whereas enamel from the hamster (continuously growing teeth) shows particles of less dense packing. The piranha teeth have a thin cuticle covering the long apatite crystals of the underlying enameloid. This cuticle has a rough surface of particles that have a globular appearance after the brief acidic treatment. The similar appearance of the in vivo naturally etched tooth surface suggests that the pattern of globule distribution may be due to the presence of an organic material. Elemental analysis of this cuticle indicated that calcium, phosphorus, and iron are the main components of the structure while electron microdiffraction of pulverized cuticle particles showed a pattern consistent with hydroxyapatite. The chiton mineralized tooth cusp had a smooth surface in an unabraded region and a very rough structure with the magnetite crystals (already known to make part of the structure) protruding from the surface. It was concluded that the structures analyzed are optimized for efficiency in feeding mechanism and life span of the teeth.     

In situ Raman spectroscopic studies of the teeth of the chiton Acanthopleura hirtosa

 In situ Raman spectroscopy, in combination with energy dispersive spectroscopy, has been used for the first time to determine the identities and locations, at the micron level, of mineral phases present in single chiton teeth that have been extensively mineralized. At the later stages of development the major lateral teeth of the chiton Acanthopleura hirtosa show characteristic spectroscopic evidence for the presence of lepidocrocite (γ-FeOOH), magnetite (Fe₃O₄), and an apatitic calcium phosphate. Goethite (α-FeOOH) and ferrihydrite (5 Fe₂O₃·9 H₂O), which have been detected previously in teeth at the early stages of mineralization, were not detected in this mature tooth. The spatial distribution of these phases was determined, providing evidence for the presence of a discrete layer of lepidocrocite between the magnetite and apatite regions, illustrating the complexity of the biomineralization process. The technique of laser Raman microscopy is shown to be ideal for the examination of small biomineralized structures in situ, such as chiton teeth. Received: 6 July 1998 / Accepted: 19 August 1998     

The mineralization and hardness of the radular teeth of the limpet Patella vulgata L.

Summary A study of the Patella vulgata radula has been made using: the scanning electron microscope in its normal and compositional contrast modes of operation, the electron microprobe analyser, ion etching with argon ions and microhardness testing.Only iron, silicon and small amounts of sulphur were detected in the radula. The teeth can be subdivided into a cusp, a junctional area where the cusp is joined to the base, and the base which is embedded in the radular membrane. From a study of longitudinal vertical and transverse sections of the mature teeth it was found that the cusp could be subdivided into a posterior iron-rich area (44–51% Fe, 1–6% Si) and an anterior silicon-rich area (22–30% Fe, 27–32% Si). The junctional zone consisted of a poorly mineralised layer at its border with the cusp and an iron-rich layer where it joined the base. The upper part of the base (5% Fe, 16% Si) could be clearly differentiated from the silicon-rich anterior and lower parts of the base (3–4% Fe, 28–35% Si). No minerals were detected in the membrane. The changes in the mineral content of the teeth cusps along the length of the radula were studied. Iron appeared in the cusps at the 25th row and the concentration increased to 28% at the 50th row. The iron was here evenly distributed throughout the cusp. Silicon appeared in the anterior part of the cusp at the 50th row and as it increased in concentration so the iron was displaced, and at the same time the concentration of iron increased in the posterior part of the cusp. Mineralization appeared to be complete by the 150th row.The teeth cusps appear to consist of 800 Å fibres grouped into 1 thick bundles and the tooth appears to be covered by a thin enamel-like layer. It is suggested that the fibres contain the silicon-rich phase and the matrix the iron-rich phase.The significance of the arrangement of the fibres and the distribution of the minerals are discussed with relation to the function of the teeth.We wish to thank Mr. A. Rees and Mr. A. Davies for their technical assistance; Prof. Lewis and Dr. James for the use of the Electron Microprobe; and the S.R.C. for their financial support.     

Proteomic analysis from the mineralized radular teeth of the giant Pacific chiton, Cryptochiton stelleri (Mollusca)

The biomineralized radular teeth of chitons are known to consist of iron-based magnetic crystals, associated with the maximum hardness and stiffness of any biomineral. Based on our transmission electron microscopy analysis of partially mineralized teeth, we suggest that the organic matrix within the teeth controls the iron oxide nucleation. Thus, we used Nano-LC-MS to perform a proteomic analysis of the organic matrix in radular teeth of the chiton Cryptochiton stelleri in order to identify the proteins involved in the biomineralization process. Since the genome sequence of C. stelleri is not available, cross-species similarity searching and de novo peptide sequencing were used to screen the proteins. Our results indicate that several proteins were dominant in the mineralized part of the radular teeth, amongst which, myoglobin and a highly acidic peptide were identified as possibly involved in the biomineralization process.     

Matrix Heterogeneity in the Radular Teeth of the Chiton Acanthopleura hirtosa

The structure and organization of the organic matrix of the cusps of the major lateral teeth of the chiton Acanthopleura hirtosahave been examined using conventional light and transmission electron microscopy techniques and by using the protein ferritin as an ultrastructural probe. The results show major structural differences in the organic matrix between the surface layers of the anterior (calcified) region and the posterior (magnetite-mineralized) region and their respective underlying regions. In addition, the central (lepidocrocite-mineralized) region of the tooth has been examined and shown to consist of bundles of fibres arranged such that they display a tightly interwoven pattern. It is suggested that while the structural organization of surface fibres readily permits the passage of ions required for mineralization, the architecturally discrete distribution of biominerals found in mature chiton teeth is due mostly to spatial delineation of the tooth by matrix macromolecules in the central region of the tooth.     

Ultrastructure and the importance of wear in the dentition of the halfbeak (Pisces: Hemiramphidae) pharyngeal mill

To assess how tooth microstructure and composition might facilitate the pharyngeal mill mechanism of halfbeaks, apatite structure and iron content were determined by scanning electron microscopy and energy dispersive X‐ray analysis for Hyporhamphus regularis ardelio, Arrhamphus sclerolepis krefftii, and Hemiramphus robustus. Iron was present in developing teeth and was concentrated along the shearing edge of spatulate incisiform teeth, which dominate the occlusive wear zone in all three species. A model based on tooth structure and wear rate is proposed to explain how halfbeaks maintain a fully functional occlusion zone throughout growth and consequent tooth addition and replacement. Replacement teeth erupt and wear rapidly so that a constant occlusion plane is always present. Iron within the tooth tissue reduces the wear rate of the cutting edge while simultaneously maintaining its sharpness and efficiency. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

The mosasaur tooth attachment apparatus as paradigm for the evolution of the gnathostome periodontium

SUMMARY Vertebrate teeth are attached to jaws by a variety of mechanisms, including acrodont, pleurodont, and thecodont modes of attachment. Recent studies have suggested that various modes of attachment exist within each subcategory. Especially squamates feature a broad diversity of modes of attachment. Here we have investigated tooth attachment tissues in the late cretaceous mosasaur Clidastes and compared mosasaur tooth attachment with modes of attachment found in other extant reptiles. Using histologic analysis of ultrathin ground sections, four distinct mineralized tissues that anchor mosasaur teeth to the jaw were identified: (i) an acellular cementum layer at the interface between root and cellular cementum, (ii) a massive cone consisting of trabecular cellular cementum, (iii) the mineralized periodontal ligament containing mineralized Sharpey's fibers, and (iv) the interdental ridges connecting adjacent teeth. The complex, multilayered attachment apparatus in mosasaurs was compared with attachment tissues in extant reptiles, including Iguana and Caiman . Based on our comparative analysis we postulate the presence of a quadruple-layer tissue architecture underlying reptilian tooth attachment, comprised of acellular cementum, cellular cementum, mineralized periodontal ligament, and interdental ridge (alveolar bone). We propose that the mineralization status of the periodontal ligament is a dynamic feature in vertebrate evolution subject to functional adaptation.     

Dentition of the apron ray Discopyge tschudii (Elasmobranchii: Narcinidae)

The present study provides quantitative and qualitative analyses of the dentition of Discopyge tschudii. Overall, 193 individuals (99 males and 94 females) of D. tschudii were collected on scientific trawl surveys conducted by the National Institute for Fisheries Research and Development (INIDEP) and commercial vessels in Argentina. Discopyge tschudii has rhombic‐shaped teeth, arranged in a semipavement‐like dentition; each tooth has an erect cusp slightly inclined posteriorly and holaulachorized root. Mature males have greater tooth lengths than females and immature specimens. Discopyge tschudii exhibits dignathic homodonty and gradient monognathic heterodonty where teeth of the commissural row are shorter than those of the symphyseal and internal rows.     

Silicification of the medial tooth in the blue crab Callinectes sapidus

Observations of cuticular structures mineralized with silica within the Crustacea have been limited to the opal teeth of copepods, mandibles of amphipods, and recently the teeth of the gastric mill in the blue crab Callinectes sapidus. Copepod teeth are deposited during premolt, with sequential elaboration of organic materials followed by secretion of silica into the tooth mold. The timing of mineralization is in stark contrast to that of the general integument of crustaceans in which calcification is completely restricted to the postmolt period. To determine the timing of molt‐related deposition and silicification of the teeth of the gastric mill, the medial tooth of the blue crab C. sapidus was examined histologically and ultrastructurally across the molt cycle. Histological data revealed deposition of the organic matrix of the epicuticle and exocuticle during premolt. No evidence of postmolt changes in the thickness of the epicuticle and exocuticle, or any deposition of endocuticle, was observed. Scanning electron microscopy revealed degradation of the outer surface of the old tooth during premolt. During premolt, epithelial structures resembling papilla appeared to secrete a fibrous web that coalesces to become the matrix of the new tooth. Semi‐quantitative elemental analyses indicated simultaneous deposition of silica and organic matrix, and demonstrated a homogeneous distribution of silicon throughout the epicuticle of the tooth at all stages. However, there is evidence of deposition (presumably silicification) during postmolt as spaces between the papillae become filled in. Thus, the pattern and timing of deposition and silicification of the tooth are different from both teeth of copepods and the general exoskeleton of decapods, and may facilitate rapid resumption of feeding and consumption of the exuvia in early postmolt. J. Morphol. 277:1648–1660, 2016. © 2016 Wiley Periodicals, Inc.     

部分蜥蜴类牙齿特征补充

The characteristics of modern lizard teeth have often been overlooked as an aid to classification. In order to i-dentify isolated teeth or rows of teeth on the jaws of Quaternary lizard fossils, we observed many modern lizard skulls with complete tooth rows, and thereby discovered that there are different patterns of tooth arrangement which are a significant aid to classification and also valuable in distinguishing lizard tooth fragments or isolated teeth. Our observations suggest that lizard teeth can be divided into three major types: 1 ) Homodont, pleurodont with single-cusp. This kind of teeth is usually slender and closely spaced. Teeth number 20 - 30 or more. The smaller-sized lizards, such as Gekkos gecko, G.Japonicus, Eumeces chinensis (Fig. 1 :A, a), E. xanthi, Leiolopisma tsinlingensis (Fig. 1 :B, b), L. reevesii, Ly-gosoma indicum, Platyurus platyurus and Hemidactylus frenatus, have this kind of arrangement. 2) Heterodont, sub-acrodont or pleurodont, with single-conical cusp teeth at the anterior of the tooth row and with flat-conical bicuspid teeth posteriorly. There are about 18 - 19 check teeth. Eremias argus (Fig. 1:C,c), E. multiocellata and E. brenchltyi have this kind of arrangement. 3 ) Heterodont, with single-conical cusp teeth in the anterior part of the tooth row and with tricuspid, subacrodont teeth posteriorly. There are vertical grooves between the teeth on the external side of the low-er jaw. The fourth tooth in most species is canine-like. There are 16 or less check teeth. The larger-sized lizards, such as Phrynocephalus przewalski, P. frontalis (Fig. 1:D,d), Japalura splendida, J. flaviceos (Fig. 1 : E, e), Calotes versicolor and Leioleps belliana etc. possess this kind of arrangement. Evolutionary trends in lizard teeth are briefly dis-cussed.     

Dental homologies in lamniform sharks (Chondrichthyes: Elasmobranchii).

The dentitions of lamniform sharks are said to exhibit a unique heterodonty called the "lamnoid tooth pattern." The presence of an inflated hollow "dental bulla" on each jaw cartilage allows the recognition of homologous teeth across most modern macrophagous lamniforms based on topographic correspondence through the "similarity test." In most macrophagous lamniforms, three tooth rows are supported by the upper dental bulla: two rows of large anterior teeth followed by a row of small intermediate teeth. The lower tooth row occluding between the two rows of upper anterior teeth is the first lower anterior tooth row. Like the first and second lower anterior tooth rows, the third lower tooth row is supported by the dental bulla and may be called the first lower intermediate tooth row. The lower intermediate tooth row occludes between the first and second upper lateral tooth rows situated distal to the upper dental bulla, and the rest of the upper and lower tooth rows, all called lateral tooth rows, occlude alternately. Tooth symmetry cannot be used to identify their dental homology. The presence of dental bullae can be regarded as a synapomorphy of Lamniformes and this character is more definable than the "lamnoid tooth pattern." The formation of the tooth pattern appears to be related to the evolution of dental bullae. This study constitutes the first demonstration of supraspecific tooth-to-tooth dental homologies in nonmammalian vertebrates.     

Tooth and consequences: Heterodonty and dental replacement in piranhas and pacus (Serrasalmidae)

Tooth replacement in piranhas is unusual: all teeth on one side of the head are lost as a unit, then replaced simultaneously. We used histology and microCT to examine tooth‐replacement modes across carnivorous piranhas and their herbivorous pacu cousins (Serrasalmidae) and then mapped replacement patterns onto a molecular phylogeny. Pacu teeth develop and are replaced in a manner like piranhas. For serrasalmids, unilateral tooth replacement is not an “all or nothing” phenomenon; we demonstrate that both sides of the jaws have developing tooth rows within them, albeit with one side more mineralized than the other. All serrasalmids (except one) share unilateral tooth replacement, so this is not an adaptation for carnivory. All serrasalmids have interlocking teeth; piranhas interdigitate lateral tooth cusps with adjacent teeth, forming a singular saw‐like blade, whereas lateral cusps in pacus clasp together. For serrasalmids to have an interlocking dentition, their teeth need to develop and erupt at the same time. We propose that interlocking mechanisms prevent tooth loss and ensure continued functionality of the feeding apparatus. Serrasalmid dentitions are ubiquitously heterodont, having incisiform and molariform dentitions reminiscent of mammals. Finally, we propose that simultaneous tooth replacement be considered as a synapomorphy for the family.     

New ischnacanthid acanthodians from the Early Devonian of Australia, with comments on acanthodian interrelationships

Rockycampacanthus milesi n.gen., n.sp. is described from a single jaw from the Rocky Camp member of Lower Devonian Buchan Group, E Victoria. Rockycampacanthus differs from other ischnacanthiforms in having large multicuspidate teeth with dual rows of secondary cusps forming a posteromesial flange, a mesial tooth row beginning opposite the fourth cusp of the main tooth row, and in the gnathal bone being deepest in the anterior half. Taemasacanthus erroli n. gen., n. sp. is described from several jaw bones from the Lower Devonian Murrumbidgee Group, New South Wales. Taemasacanthus has a well developed posterolabial flange with secondary cusps developed, vertical rows of denticles on the cusps of the main tooth row and a well developed mesial tooth row separated from the main row by a prominent ridge. The labial face of the jaw has a circular ridge which may have supported labial cartilages. The complex mandibular joint in climatiforms, acanthodiiforms and some primitive sharks differs from the simple jaw articulation of ischnacanthids. It is suggested that ischnacanthids are the plesiomorphic sister group to climatiforms plus acanthodiiforms. The interrelationships of ischnacanthids, climatiforms and acanthodiforms are discussed.