The appearance pattern of tooth germs in the round crucian carp,Carassius auratus grandoculis

Cyprinid fishes generally replace their teeth alternately and cephalad. The larvae ofCarassius auratus grandoculis also replace their teeth alternately and cephalad, in a pattern of 4-2-3-1-. However, adults ofCarassius species replace their teeth from anterior to posterior, in a pattern of 1-2-3-4-1-. So I analyzed the appearance pattern of tooth germs in larvae and juveniles inCarassius auratus grandoculis. At stage 5 of the post-larval period, developmental difference is made between both sides. In the pharyngeal dentition on one side developing poorly, the anterior tooth on the fifth replacement wave, tooth₄[An2] appeared later than the central teeth on following replacement wave, tooth₅[Pol]. Moreover, the anterior tooth on the seventh replacement wave, tooth₆[An2], appeared later than the central teeth on the following replacement wave, tooth₇[Pol], on both sides. The reverse of tooth germ appearance between anterior teeth and central teeth makes a change of replacement pattern from 4-2-3-1-4- to 1-2-3-4-1-. The change of replacement pattern is caused by the confusion of tooth germs of anterior teeth on both sides.Mylopharyngodon piceus andCyprinus carpio make a change of replacement patterns in the early juvenile period, too. This change of replacement pattern may be a specialized character among the subfamily Cyprininae.     

Development of the pharyngeal teeth in the big head,Aristichthys nobilis (Cyprinidae)

The development of pharyngeal dentition was observed in the big head,Aristichthys nobilis, which is one of the hypophthalmichthyines of the cyprinids. This fish has the C-type larval dentition, in which no teeth ever occur at the position An3, and in which the first tooth at the position An2 is on the third replacement wave. So the positions Pol, Ce0, Ani and An2 in the larval dentition correspond to the positions A4, A3, A2 and A1 in the adult dentition, respectively. The initial tooth at each position is a conical one. The conical teeth are then changed to ones bearing a narrow grinding surface with a hook at the tip and some denticles on the margins. These teeth are of theLeuciscus stage. Tn the following teeth, the grinding surface is expanded, and the denticles are increased in number and distributed on not only the margins but also the whole grinding surface. These teeth bearing a very broad grinding surface characterize the hypophthalmichthyines. At the positions A2 to A4, the teeth become the hypophthalmichthyine type in the larval period. But the tooth at the position A1 becomes the hypophthalmichthyine type in the juvenile period. The morphological change of teeth in this species is simple although their teeth are highly specialized. We think that this phenomenon gives a hint on their phylogeny.     

Tooth development and replacement in the Atlantic Cutlassfish,Trichiurus lepturus,with comparisons to other Scombroidei

Atlantic Cutlassfish, Trichiurus lepturus, have large, barbed, premaxillary and dentary fangs, and sharp dagger-shaped teeth in their oral jaws. Functional teeth firmly ankylose to the dentigerous bones. We used dry skeletons, histology, SEM, and micro-CT scanning to study 92 specimens of T. lepturus from the western North Atlantic to describe its dentition and tooth replacement. We identified three modes of intraosseous tooth replacement in T. lepturus depending on the location of the tooth in the jaw. Mode 1 relates to replacement of premaxillary fangs, in which new tooth germs enter the lingual surface of the premaxilla, develop horizontally, and rotate into position. We suggest that growth of large fangs in the premaxilla is accommodated by this horizontal development. Mode 2 occurs for dentary fangs: new tooth germs enter the labial surface of the dentary, develop vertically, and erupt into position. Mode 3 describes replacement of lateral teeth, in which new tooth germs enter a trench along the crest of the dentigerous bone, develop vertically, and erupt into position. Such distinct modes of tooth replacement in a teleostean species are unknown. We compared modes of replacement in T. lepturus to 20 species of scombroids to explore the phylogenetic distribution of these three replacement modes. Alternate tooth replacement (in which new teeth erupt between two functional teeth), ankylosis, and intraosseous tooth development are plesiomorphic to Bluefish + other Scombroidei. Our study highlights the complexity and variability of intraosseous tooth replacement. Within tooth replacement systems, key variables include sites of formation of tooth germs, points of entry of tooth germs into dentigerous bones, coupling of tooth germ migration and bone erosion, whether teeth develop horizontally or immediately beneath the tooth to be replaced, and how tooth eruption and ankylosis occur. Developmentally different tooth replacement processes can yield remarkably similar dentitions.     

鲤形目鱼类咽齿形态及发育的比较研究

对鲤形目所属６科类的咽龄形态及个体发育模型进行了较详细的比较研究。结果得出除双孔鱼科鱼类未形成咽齿外,其它５科鱼类在咽齿发生的过程中具有齿着生方向完全相反的两种发育模式。以亚口鱼科,鳅科和平鳍鳅为一类的齿后向增长的类型和以鲤科另一类的齿同增长的类型。     

Tooth replacement in the red-backed salamander, Plethodon cinereus

The developmental cycle of the teeth in Plethodon cinereus is analyzed on morphological grounds using alizarin preparations. All the stages in development do not occupy the same proportion of the life cycle time. Functional teeth and germs at an early stage in development occupy a large proportion of the life cycle time, whereas the processes of tooth shedding and ankylosis occur very quickly. The time during which any locus does not bear a functional tooth, and is therefore a non-functional locus, is reduced to a minimum. P. cinereus has a basic pattern of tooth replacement which is consistent with Zahnreihen which are 2.0 tooth spaces apart. Variations in the replacement pattern are common and these are produced by relatively small fluctuations in the spacing of the Zahnreihen around the ?mean? of 2.0. Localized disturbances which produce breaks in the replacement pattern and cause waves to cross also occur. These may be due to the failure of tooth germs to develop, the fusion of tooth germs, or may be the result of the inherent variability in a complex biological system. This variability causes individual tooth germs to develop too slowly or too quickly and hence assume an ?abnormal”? position thus causing breaks in the replacement pattern. Tooth replacement may be controlled by an intra-local mechanism(s) rather than by stimuli which travel along the jaw.     

Structure, attachment, replacement and growth of teeth in bluefish, Pomatomus saltatrix (), a teleost with deeply socketed teeth

Tooth replacement poses many questions about development, pattern formation, tooth attachment mechanisms, functional morphology and the evolution of vertebrate dentitions. Although most vertebrate species have polyphyodont dentitions, detailed knowledge of tooth structure and replacement is poor for most groups, particularly actinopterygians. We examined the oral dentition of the bluefish, Pomatomus saltatrix, a pelagic and coastal marine predator, using a sample of 50 individuals. The oral teeth are located on the dentary and premaxillary bones, and we scored each tooth locus in the dentary and premaxillary bones using a four-part functional classification: absent (A), incoming (I), functional (F=fully ankylosed) or eroding (E). The homodont oral teeth of Pomatomus are sharp, deeply socketed and firmly ankylosed to the bone of attachment. Replacement is intraosseus and occurs in alternate tooth loci with long waves of replacement passing from rear to front. The much higher percentage of functional as opposed to eroding teeth suggests that replacement rates are low but that individual teeth are quickly lost once erosion begins. Tooth number increases ontogenetically, ranging from 15–31 dentary teeth and 15–39 premaxillary teeth in the sample studied. Teeth increase in size with every replacement cycle. Remodeling of the attachment bone occurs continuously to accommodate growth. New tooth germs originate from a discontinuous dental lamina and migrate from the lingual (dentary) or labial (premaxillary) epithelium through pores in the bone of attachment into the resorption spaces beneath the existing teeth. Pomatomus shares unique aspects of tooth replacement with barracudas and other scombroids and this supports the interpretation that Pomatomus is more closely related to scombroids than to carangoids.     

扁圆吻鲴下咽齿的个体发生及替换规律

扁圆吻鲴下咽齿的个体发生可分为三个阶段：初齿期、过渡齿期和成齿期。初步期符合鲤科鱼类的一般规律;过渡齿期相当延长,产生全部齿位,６或７枚齿,齿的发生存在两种类型;成齿与幼齿的替换规律完全不同,发育进入成齿阶段后,主行齿由奇数齿位与偶数齿位交错替换转变为相二枚齿进行替换,替换公式为１－４,２－５,３－６或１－４－７,２－５,３－６（主行齿６枚或７枚）,全部替换一次分三列替换波完成,可将扁圆吻鲴下咽齿的发育模式视为新的类型,副行齿在过渡齿期出现,与主行齿的发展模式不同,替换形式始终为相令齿位交错进行,本文还探讨了咽骨的发育及其对下咽发生的影响。     

Structure, attachment, replacement and growth of teeth in bluefish, Pomatomus saltatrix (Linnaeus, 1776), a teleost with deeply socketed teeth

Tooth replacement poses many questions about development, pattern formation, tooth attachment mechanisms, functional morphology and the evolution of vertebrate dentitions. Although most vertebrate species have polyphyodont dentitions, detailed knowledge of tooth structure and replacement is poor for most groups, particularly actinopterygians. We examined the oral dentition of the bluefish, Pomatomus saltatrix, a pelagic and coastal marine predator, using a sample of 50 individuals. The oral teeth are located on the dentary and premaxillary bones, and we scored each tooth locus in the dentary and premaxillary bones using a four-part functional classification: absent (A), incoming (I), functional (F=fully ankylosed) or eroding (E). The homodont oral teeth of Pomatomus are sharp, deeply socketed and firmly ankylosed to the bone of attachment. Replacement is intraosseus and occurs in alternate tooth loci with long waves of replacement passing from rear to front. The much higher percentage of functional as opposed to eroding teeth suggests that replacement rates are low but that individual teeth are quickly lost once erosion begins. Tooth number increases ontogenetically, ranging from 15–31 dentary teeth and 15–39 premaxillary teeth in the sample studied. Teeth increase in size with every replacement cycle. Remodeling of the attachment bone occurs continuously to accommodate growth. New tooth germs originate from a discontinuous dental lamina and migrate from the lingual (dentary) or labial (premaxillary) epithelium through pores in the bone of attachment into the resorption spaces beneath the existing teeth. Pomatomus shares unique aspects of tooth replacement with barracudas and other scombroids and this supports the interpretation that Pomatomus is more closely related to scombroids than to carangoids.     

On the control of tooth replacement in reptiles and its relationship to growth

The teeth of nearly all non-mammalian vertebrates are replaced in waves which sweep through alternate tooth positions. It is argued that tooth replacement in these animals represents growth of the dentition. It is shown that the pattern of tooth replacement could be described by the exponential equation t_(n)r, = k e^ar+bn when t(n)r is the time at which the rth replacement erupts in the nth position and k, a and b are constants. The length of a replacement wave (w) which is visible in the mouth, can be calculated from the equation w = 2(a?b)/a?2b for forward travelling waves. The effect of different ratios, $\text{a}\text{b}$, on wavelength is described. The model can be interpreted as describing the effect of a zone of inhibition which (it is argued) temporarily surrounds any newly initiated tooth. The increasing time required to dissipate the inhibition around successive replacement teeth is related to the age of the animal. This increasing time permits successive teeth to grow for longer periods than their predecessors and can account for a gradual increase in the size of successive teeth. A similar mechanism could account for the phasic nature of bone growth. It is indicated that the model could be difficult to test.     

Tooth replacement pattern of Coloborhynchus robustus (Pterosauria) from the Lower Cretaceous of Brazil

The well preserved anterior upper and lower jaw fragment of an adult specimen of Coloborhynchus robustus (Pterosauria: Ornithocheiridae), SMNK 2302 PAL, allowed investigations of the replacement pattern of the dentition macroscopically and by using CT scans. The quantification of the dentition by Zahnreihen, Z-Spacing, and replacement waves indicates a complex pattern of different replacement stages in which large gaps within the dentition were avoided. The specialized prey-catching apparatus of Coloborhynchus thus could retain its function even following tooth replacement. The replacement process in the specimen took about 2/3 of the total life-time of a tooth, and damaged teeth in the anterior jaw region may have been replaced more rapidly than posterior teeth. The distolingual replacement of the functional teeth delayed the time of their shedding in comparison with the circular resorption present in crocodiles. In contrast to these, the distolingual position of the replacement tooth did not decrease the biomechanical stability of the functional tooth, which can also be observed as a convergence in other thecodont dentitions, e.g., recent carnivore mammals. Teeth were shed when their replacement had reached about 60% of the full-grown height. A comparison of the observed pattern is constricted by the preservation and preparation of other specimens. Unfortunately, no known specimen in public collections reaches the quality of Coloborhynchus robustus, SMNK 2302 PAL, so that comparable patterns in other specimens are not likely to be detected.     

Mosquito species distribution and larval breeding habitats with taxonomic identification of Anopheline mosquitoes in Korea

In 2005, adult and larval mosquito surveillance was conducted at selected sites in Korea to associate larval habitats with species distribution of mosquitoes of the Anopheles Hyrcanus Group (An. sinensis, An. lesteri, An. pullus, An. belenrae and An. kleini) and other mosquito species. Anopheles specimens belonging to the Anopheles Hyrcanus Group were identified to species level by molecular confirmation using the internal transcribed spacer (ITS)‐2 within nuclear ribosomal (r)DNA. A total of 6644 mosquitoes from resting and light trap collections (4451; 67.0%) and larval collections (2193; 33.0%) comprising 32 species and nine genera (Culex [11], Anopheles [8], Ochlerotatus [5], Aedes [3], Armigeres [1], Coquillettidia [1], Mansonia [1], Tripteroides [1] and Lutzia [1]) were collected. Larval habitats were characterized into 14 categories. Of a total of 4534 Anopheles spp. collected (3766 resting and light trap collections and 768 larval collections), Anopheles sinensis (3194; 70.4%) was the most frequently captured, followed by An. kleini (813; 17.9%), An. pullus (299; 6.6%) and An. belenrae (129; 2.8%). Four species of Anopheles (An. lesteri, An. sineroides, An. koreicus and An. lindesayi) were infrequently collected (<3.0%) at all sites surveyed by all methods of collection. Anopheles kleini, An. pullus and An. belenrae were collected in greater proportions in malaria high‐risk areas north of Seoul, and were infrequently collected in other parts of Korea, where An. sinensis was the predominant Anopheles spp. captured. A total of 2110 culicine mosquitoes (685 adult collections and 1425 larval collections) comprising 24 species and eight genera were collected.     

Enameloid/enamel transition through successive tooth replacements in <Emphasis Type="Italic">Pleurodeles waltl</Emphasis> (Lissamphibia,Caudata)

Study of the evolutionary enameloid/enamel transition suffers from discontinuous data in the fossil record, although a developmental enameloid/enamel transition exists in living caudates, salamanders and newts. The timing and manner in which the enameloid/enamel transition is achieved during caudate ontogeny is of great interest, because the caudate situation could reflect events that have occurred during evolution. Using light and transmission electron microscopy, we have monitored the formation of the upper tooth region in six successive teeth of a tooth family (position I) in Pleurodeles waltl from late embryos to young adult. Enameloid has only been identified in embryonic tooth I₁ and in larval teeth I₂ and I₃. A thin layer of enamel is deposited later by ameloblasts on the enameloid surface of these teeth. From post-metamorphic juvenile onwards, teeth are covered with enamel only. The collagen-rich enameloid matrix is deposited by odontoblasts, which subsequently form dentin. Enameloid, like enamel, mineralizes and then matures but ameloblast participation in enameloid matrix deposition has not been established. From tooth I₁ to tooth I₃, the enameloid matrix becomes ever more dense and increasingly comes to resemble the dentin matrix, although it is still subjected to maturation. Our data suggest the absence of an enameloid/enamel transition and, instead, the occurrence of an enameloid/dentin transition, which seems to result from a progressive slowing down of odontoblast activity. As a consequence, the ameloblasts in post-metamorphic teeth appear to synthesize the enamel matrix earlier than in larval teeth.     

锦鲤咽齿替换及齿胚的解剖学观察,前插1

从鱼类解剖学和口腔医学的角度,观察和讨论了锦鲤咽齿、齿胚的形态结构和咽齿的替换,并对咽齿和咬合板做了扫描电镜观察.发现锦鲤咽齿替换与齿胚有密切关系;咽齿替换分为两侧不同名齿同时替换、同侧两枚不同名齿同时替换和单侧一枚齿自主替换3种类型;咽齿替换是终生发生的,遵循着"被替换咽齿的骨性支持从基骨骨孔开始吸收-同名齿胚的移行与就位-旧齿脱落与新齿支持组织改建为骨性"的过程.认为齿胚的发育是启动咽齿替换的起始因素;锦鲤与野生鲤有近缘关系.     

Development and fine structure of pharyngeal replacement teeth in juvenile zebrafish (Danio rerio) (Teleostei, Cyprinidae)

Teeth are commonly used model systems for the study of epithelial-mesenchymal interactions during organogenesis. We describe here the ultrastructural characteristics of developing pharyngeal replacement teeth in juvenile zebrafish, an increasingly important model organism for vertebrate development. Replacement teeth develop in close association with the dental organ of a functional tooth. Morphogenesis is well advanced prior to the start of cytodifferentiation. Fibrillar enameloid matrix is formed first, followed by the deposition of predentine. Initial mineralization of the enameloid proceeds quickly; maturation involves the presence of ruffled-bordered ameloblasts. Dentine mineralization is inotropic and is mediated by matrix vesicles. Woven-fibred attachment bone matrix is deposited before completion of dentine mineralization. Eruption of fully ankylosed teeth is a fast process and may involve degenerative changes in the pharyngeal epithelium. Mononucleated osteoclasts and clastic cells located in the pulp cavity intervene in tooth resorption prior to shedding. Structural differences with larval, first-generation zebrafish teeth include the presence of dentinal tubules and the absence of an electron-dense covering membrane. Part of these differences may relate to size differences of the teeth. Others, like the site of the replacement tooth bud, suggest that initiation may take place in already committed epithelium from the first initiation event in the larval stage.     

Ontogenetic transition from unicuspid to multicuspid oral dentition in a teleost fish: Astyanax mexicanus, the Mexican tetra (Ostariophysi: Characidae)

Teleost fishes display a remarkable diversity of adult dentitions; this diversity is all the more remarkable in light of the uniformity of first-generation dentitions. Few studies have quantitatively documented the transition between generalized first-generation dentitions and specialized adult dentitions in teleosts. We investigated this transition in the Mexican tetra, Astyanax mexicanus (Characidae), by measuring aspects of the dentition in an ontogenetic series of individuals from embryos to 160 days old, in addition to adults of unknown age. The first-generation dentition and its immediate successors consist of small, unicuspid teeth that develop extraosseously. Multicuspid teeth first appear during the second tooth replacement event, and are derived from single tooth germs, rather than from the fusion of multiple conical tooth germs. We document that the transition from unicuspid to multicuspid teeth corresponds to a change in the location of developing tooth germs (from extraosseous to intraosseous) and in patterns of tooth replacement (from haphazard to simultaneous within a jaw quadrant). In addition, while the size of the largest teeth scales with positive allometry to fish size, the transition to multicuspid teeth is accompanied by an exceptionally large increase in tooth size.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 145 , 523–538.     

Torque generation by one of the motor subunits of heterotrimeric kinesin-2

Heterotrimeric kinesin-2 motors [1] and [2] transport intraflagellar transport (IFT)-particles from the base to the tip of the axoneme to assemble and maintain cilia [3], [4], [5], [6], [7], [8], [9] and [10]. These motors are distinct in containing two non-identical motor subunits together with an accessory subunit [1], [11], [12], [13], [14] and [15]. We evaluated the significance of this organization by comparing purified wild type kinesin-2 holoenzymes that support IFT in vivo, with mutant trimers containing only one type of motor domain that do not support IFT in vivo. In motility assays, wild type kinesin-2 moved microtubules (MTs) at a rate intermediate between the rates supported by the two mutants. Interestingly, one of the mutants, but not the other mutant or the wild type protein, was observed to drive a persistent counter-clock-wise rotation of the gliding MTs. Thus one of the two motor domains of heterotrimeric kinesin-2 exerts torque as well as axial force as it moves along a MT, which may allow kinesin-2 to control its circumferential position around a MT doublet within the cilium.     

The ultrastructure of odontogenesis in larval and adult urodeles; differentiation of the dental epithelial cells

Summary Sections of undemineralized tooth germs ofAmbystoma andTriturus were examined. The ultrastructure of early germs, both larval and adult, and of dentinogenesis, resembled that of mammals. In adult bicuspid teeth, once the dentine of the cusps was mineralized, mineral crystals of a similar size to early mammalian enamel crystals, appeared between the dentine and the inner dental epithelium (i.d.e). Concomitantly, the i.d.e showed features of mammalian secreting ameloblasts. This new layer, regarded as true enamel, lacked collagen, possessed an ordered arrangement of crystals and reached a maximum thickness of 6 m.In larval monocuspid teeth, once dentine mineralization had reached the plasma membranes of the i.d.e at the tip of the cusp, the i.d.e developed a ruffled border. At this stage the dentine of the tip, regarded as enameloid, was very hard and difficult to section. The ruffled border, characteristic of other cells which transport materials, was regarded as indicating that the i.d.e was removing organic matter from the enameloid. The differences in development between larval and adult teeth support the concept that there is a change in cellular activity of the i.d.e which occurs during metamorphosis from the larval to the adult urodele.     

Morphological studies on the mouth cavity of Urodela IX. Teeth of the palate and the splenials in Siren and Pseudobranchus (Sirenidae: Amphibia)

1. Dentition, tooth structure and course of dental laminae of adult and subadult Sirenidae (Siren intermedia, S. lacertina, Pseudobranchus striatus) have been studied by light microscopy and scanning electron microscopy. 2. Splenials, vomers und palatines bear monocuspid unbladed teeth, arranged in a polystichous pattern, whereas praemaxillaries and maxillaries (so far as present) are edentate. Teeth in S. intermedia show a rough basis, which is more prominent in S. lacertina and which has break-throughs in P. striatus. This zone perhaps is homologous to a developing dividing zone typical for teeth in many “Lissamphibia”. 3. With respect to structure and organisation of dentigerous bones and teeth Sirenidae obviously possess a mosaicism of differently developed larval characters in their mouth cavity. 4. The dentition in the recent forms investigated is compared to that of other paedomorphic Urodela und the ancient Habrosaurus dilatus (Sirenidae).     

Tooth Replacement in Late Jurassic Dryolestidae (Eupantotheria, Mammalia)

The discovery of juvenile dentitions of late Jurassic (Kimmeridgian) Dryolestidae (Eupantotheria, Mammalia) from Guimarota, Portugal, yields for the first time information on the mode of tooth replacement in therian mammals prior to the dichotomy of placentals and marsupials. As in extant placentals, tooth replacement occurs at all antemolar positions [incisors (I1–I4), canine (C), premolars (P1–P4)]. P1 and P2 have premolariform milk predecessors, whereas the large premolariform third (P3) and fourth premolars (P4) are preceded by molariform deciduous premolars (dP3, dP4). Tooth replacement takes place in two waves, at least in the lower jaw, with I₂, I₄, P₁, and P₃ in the first series and I₁, I₃, C, P₂, and P₄ in the second. P₄ is the last premolar to erupt, and it is present when the sixth molar (M₆) starts to break through. The reduced tooth replacement pattern of marsupials (with only dP3 being replaced postnatally) evolved secondarily from the primitive or plesiomorphic mammalian condition, which was retained in Dryolestidae and Eutheria.