Evolution and development of the chordates: collagen and pharyngeal cartilage

Chordates evolved a unique body plan within deuterostomes and are considered to share five morphological characters, a muscular postanal tail, a notochord, a dorsal neural tube, an endostyle, and pharyngeal gill slits. The phylum Chordata typically includes three subphyla, Cephalochordata, Vertebrata, and Tunicata, the last showing a chordate body plan only as a larva. Hemichordates, in contrast, have pharyngeal gill slits, an endostyle, and a postanal tail but appear to lack a notochord and dorsal neural tube. Because hemichordates are the sister group of echinoderms, the morphological features shared with the chordates must have been present in the deuterostome ancestor. No extant echinoderms share any of the chordate features, so presumably they have lost these structures evolutionarily. We review the development of chordate characters in hemichordates and present new data characterizing the pharyngeal gill slits and their cartilaginous gill bars. We show that hemichordate gill bars contain collagen and proteoglycans but are acellular. Hemichordates and cephalochordates, or lancelets, show strong similarities in their gill bars, suggesting that an acellular cartilage may have preceded cellular cartilage in deuterostomes. Our evidence suggests that the deuterostome ancestor was a benthic worm with gill slits and acellular gill cartilages.     

Innate immunity in early chordates and the appearance of adaptive immunity

In the urochordate Ciona intestinalis some membrane Immunoglobulin superfamily members with ancestral features of antigen receptors are homologs of vertebrate adhesion molecules acting as virus receptors. They include the following: the junction adhesion molecule (reovirus receptor) (JAM), the Cortical thymocyte marker of Xenopus (CTX family) (Coxsackie's virus receptor) and the poliovirus receptor (PVR). In humans these genes belong to the same linkage group, of which 4 paralogous groups exist. This situation is consistent with the notion that the Ciona set of genes would correspond to a preduplication state. In addition, the human region 3q13 and its paralogs, harbour genes remotely related to the nectin family that can be detected in Protostomes (human CRTAM and CD80-86 related to Drosophila Beat). In addition, this linkage group contains several CDs important for the immune system CD166, CD47 and many members of the tetraspanin family. The VC1-like core of the nectin is homologous to the VCI core of the MHC-linked tapasin and to the VC1 segments of, for example, specific antigen receptors of vertebrates, and could be related to a primitive antigen receptor gene. It is suggested that the virus binding property of the members of this family was exploited, and that they were recruited in the vertebrate immune system following the introduction of the somatic rearrangement machinery. In this way the adaptive immune system could have developed from a set of receptors involved in a primitive local innate immunity involving NF-kappaB-mediated apoptosis.     

Some ultrastructural observations on the microfilaria of Breinlia sergenti-the pharyngeal thread and Innenkorper

Some ultrastructural observations on the microfilaria of Breinlia sergenti—the pharyngeal thread and Innenkorper.International Journal for Parasitology4: 375–382. The fine structure of the pharyngeal thread and the Innenkorper (inner body) of the microfilaria of Breinlia sergenti is described. The pharyngeal thread is seen as an elongated cuticular structure with associated nervous elements while the Innenkorper is sac-like in appearance. A possible function related to the transfer and storage of nutrient material is assigned to these structures.     

On the phylogenetic significance of sagittocysts and copulatory organs in acoel turbellarians

Viewed by SEM and TEM, sagittocysts of Convoluta bifoveolata Mamkaev, 1971, and needles of C. sagittifera Ivanov, 1952, have the same structure. Both are capsule-form extrusomes containing a protrusible needle. Only seven similar species of convolutimorph acoels symbiotic with green algae and C. sagittifera, without algae, possess extrusomes of this peculiar and complicated type. The sagittocyst is a clear synapomorphy of all these species. A sacciform ciliated antrum lacking a seminal vesicle is also characteristic of these species and also of three Japanese species of green (algae-symbiotic) convolutimorph acoels lacking sagittocysts. We suggest schemes of the possible evolution of male and female copulatory organs to provide a basis for better using such organs as phylogenetic characters. We regard the formation of a ciliated sacciform antrum as an independent evolutionary trend. This conclusion forms the basis for establishing the separate family Sagittiferidae. Species of this family seem to have originated in the West Pacific.     

Four rows of pharyngeal teeth in an aberrant specimen of the small African barb Barbus paludinosus(Cyprinidae): novelty or atavistic alteration?

An extra fourth row of pharyngeal teeth was found on both pharyngeal arches in a specmen of the small African barb, Barbus paludinosus , a species normally exhibiting three rows of pharyngeal teeth. Possible phylogenetic significance of this alteration is discussed in connection with occurrence of extra fourth rows of pharyngeal teeth in other cyprinid lineages.     

Functional morphology of the pharyngeal jaw apparatus in moray eels

Moray eels (Muraenidae) are a relatively large group of anguilliform fishes that are notable for their crevice-dwelling lifestyle and renowned for their ability to consume large prey. Morays apprehend their prey by biting and then transport prey by extreme protraction and retraction of their pharyngeal jaw apparatus. Here, we present a detailed interpretation of the mechanisms of pharyngeal jaw transport based on work with Muraena retifera. We also review what is known of the moray pharyngeal jaw apparatus from the literature and provide comparative data on the pharyngeal jaw elements and kinematics for other moray species to determine whether interspecific differences in morphology and behavior are present. Rather than comprising broad upper and lower processing tooth plates, the pharyngeal jaws of muraenine and uropterygiine morays, are long and thin and possess large, recurved teeth. Compared with the muraenines, the pharyngobranchials of the uropterygiines do not possess a horn-shaped process and their connection to the fourth epibranchial is dorsal rather than medial. In addition, the lower tooth plates do not exhibit a lateral groove that serves as a site of muscle attachment for the pharyngocleitheralis and the ventral rather than the lateral side of the lower tooth plate attaches to the fourth ceratobranchial. In all morays, the muscles positioned for protraction and retraction of the pharyngeal apparatus have undergone elongation, while maintaining the generalized attachment sites on the bones of the skull and axial skeleton. Uropterygiines lack a dorsal retractor muscle and we presume that retraction of the pharyngeal jaws is achieved by the pharyngocleitheralis and the esophagus. The fifth branchial adductor is greatly hypertrophied in all species examined, suggesting that morays can strongly adduct the pharyngeal jaws during prey transport. The kinematics of biting behavior during prey capture and transport resulted in similar magnitudes of cranial movements although the timing of kinematic events was significantly different and the duration of transport was twice as long as prey capture. We speculate that morays have evolved this alternative prey transport strategy as a means of overcoming gape constraints, while hunting in the confines of coral reefs.     

Cellular succession in the epithelium lining the pharyngeal cavity of planarians

Regardin cell renewal of planarian epithelial tissue, little is known at present except for the epidermis (Tyler, 1984). According to Skaer (1965), the definitive epidermal cells in planarian embryos originate in the parenchyma where they develop as precursor cells which then migrate into the epidermis. This process of development is also reported in regenerating planarians (Spiegelman & Dudley, 1973; Morita & Best, 1974; Hori, 1978). Recently the validity of Skaer's conclusion has clearly been confirmed by Ishii & Sakurai (1998) using SEM and TEM.The present study deals with the origin and renewal of cells of another epithelium — that lining the pharyngeal cavity of a freshwater planarian Dugesia japonica. This epithelium is known to consist of cells with secretory granduls of fingerprint-like structure (Ishii, 1966; Bowen & Ryder, 1973; Gamo & Garcia-Corrales, 1988; Asai, 1991). The characteristic feature of these specific granules, as well as epitheliosomes (Tyler, 1984) of the epidermis, is very useful as a cell marker for distinguishing cell types and following cell differentiation. Gamo & Garcia-Corrales (1988) claimed the cell populations of the pharyngeal epithelium of Dugesia gonocephala is morphologically and functionally heterogeneous. The sequence of morphological alteration offers the oppurtunity to distinguish at least three functional stages. These represent different adaptive forms of a single cell type, and indicate transformation from one to the next induced by cell renewal. It suggests a beta cell (noeblast) origin of the epithelial cells.In the present study SEM observations revealed theat there are morphologically three distinct cell types which are considered to be juvenile, mature, and senile cells respectively, judging from the differences in cells and the many transitional forms among them (Fig. 1). When observed by TEM, the cytoplasmic patterns of these cell types are correspondingly different from one another, similarly the surface specializations of the plasma membranes of each cell type which facilitate their identification by TEM also vary. Apparently alterations of cell morphology are involved in ageing in a single cell type. Morphological features of each cell type are summarized. Juveniles (R) rarely occur, are small cells provided with prominent apical ruffles and considered to be phagocytic because the cells often contain newly formed phagosomes. Mature cells (F) predominate, are large, polygonal, flat cells, presumably with an intense secretory activity. The cells contain many profiles of granular ER and a small number of specific (secretory) granules. Senile cells (M) are intermittently distributed, irregularly contoured cells with heavy surface microvilli. The specific granules are dense in the apical cytoplasm, and profiles of granular ER decease in number.The precursor epithial cells that are seen in the parenchyma exhibit a considerable degree of differentiation before their translocation into the lining. The primordial cells observed are neoblast-like, with several cohromatoid bodies and many free ribosomes (Morita et al., 1969). These results obtain thus support the evidence and prediction of Gamo & Garcia-Corrales (1988). The observed mode of cell replacement appears to be the same as found in the epidermis (Lentz, 1967).     

Morphological analysis of the pharyngeal jaws in two populations of Lebias fasciata Valenciennes, 1821 (Teleostei: Cyprinodontidae)

The study of the pharyngeal jaws in two geographically isolated Italian populations of Lebias fasciata indicated the presence of two phenotypes: the Adriatic phenotype with a large ceratobranchial V and upper and lower pharyngeal jaws bearing few large teeth and the Sicilian phenotype with a smaller ceratobranchial V and pharyngeal jaws with smaller and more numerous teeth. The morphological variations of pharyngeal jaws should be interpreted as a result of the geographical isolation of these two populations. J. Morphol. 241:107–114, 1999. © 1999 Wiley‐Liss, Inc.     

二氢叶酸还原酶基因在斑马鱼咽弓发育过程中作用的研究

叶酸缺乏可导致胚胎先天性发育异常,二氢叶酸还原酶是叶酸生物学作用通路中的关键因子,其功能阻抑将抑制叶酸生物学作用的发挥.咽弓是脊椎动物胚胎发育中头面部结构、心脏流出道等的共同前体.在模式生物斑马鱼中,利用基因表达阻抑以及过表达技术,探讨二氢叶酸还原酶基因(DHFR)在斑马鱼咽弓发育过程中的作用.石蜡切片以及软骨染色结果显示,DHFR表达阻抑导致斑马鱼咽弓以及腭发育明显异常,而DHFR过表达可部分挽救上述发育异常表型.TBX1和HAND2在咽弓发育中有重要作用.通过胚胎整体原位杂交以及Real-timePCR技术检测TBX1和HAND2表达水平.DHFR表达阻抑后TBX1和HAND2的表达降低,DHFR过表达可使TBX1和HAND2的表达增加.上述结果表明,DHFR在斑马鱼咽弓发育过程中扮演重要角色,DHFR通过影响TBX1和HAND2的表达而调控咽弓的形成和分化.     

Overlapping origins of pharyngeal arch crest cells on the postotic hind-brain

The developing hind-brain of vertebrates consists of segmental units called rhombomeres. Although crest cells emigrate from the hind-brain, they are subsequently subdivided into several cell populations that are attached to restricted regions of the hind-brain. At the preotic level, only even-numbered rhombomeres are accompanied by crest cells, while the odd-numbered ones are not. At the postotic level, such the birhombomeric repetition becomes obscure. In order to map the origins and distributions of postotic crest cells, focal injections of Dil were made into various axial levels of the postotic neural tube. Cephalic crest cells at the postotic level first form a single cell population deposited by cells along the dorsolateral pathway. They are called the circumpharyngeal crest cells (CP cells) and are secondarily subdivided into each pharyngeal arch ectomesenchyme. The neural tube extending from r5 to the somite 3/4 boundary gave rise to CP cells. The neuraxial origins of each pharyngeal ectomesenchyme extended for more than three somite lengths, most of which overlapped with the other. Unlike in the preotic region, there is no segmental registration between neuraxial levels and pharyngeal arches. Caudal portions of the CP cell population show a characteristic distribution pattern that circumscribes the postotic pharyngeal arches caudally. Heterotopic transplantation of the Dil-labeled neural crest into the somite 3 level resulted in a distribution of labeled cells similar to that of CP cells, suggesting that the pattern of distribution depends upon dynamic modification of the body wall associated with pharyngeal arch formation.     

Development of the pharyngeal arch skeleton in Catostomus commersonii (Teleostei: Cypriniformes)

Skeletal elements of the gill arches of adult cypriniform fishes vary widely in number, size, and shape and are important characters in morphologically based phylogenetic studies. Understanding the developmental basis for this variation is thus phylogenetically significant but also important in relation to the many developmental genetic and molecularly based studies of the early developing and hence experimentally tractable gill arches in the zebrafish, a cyprinid cypriniform. We describe the sequence of the chondrification and ossification of the pharyngeal arches and associated dermal bones from Catostomus commersonii (Catostomidae, Cypriniformes) and make selected comparisons to other similarly described pharyngeal arches. We noted shared spatial trends in arch development including the formation of ventral cartilages before dorsal and anterior cartilages before posterior. Qualitatively variable gill arch elements in Cypriniformes including pharyngobranchial 1, pharyngobranchial 4, and the sublingual are the last such elements to chondrify in C. commersonii. We show that the sublingual bone in C. commersonii has two cartilaginous precursors that fuse and ossify to form the single bone in adults. This indicates homology of the sublingual in catostomids to the two sublingual bones in the adults of cobitids and balitorids. Intriguing patterns of fusion and segmentation of the cartilages in the pharyngeal arches were discovered. These include the individuation of the basihyal and anterior copula through segmentation of a single cartilage rod, fusion of cartilaginous basibranchials 4 and 5, and fusion of hypobranchial 4 with ceratobranchial 4. Such “fluidity” in cartilage patterning may be widespread in fishes and requires further comparative developmental studies. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Anatomy of a hemiramphid pharyngeal mill with reference to Arrhamphus sclerolepis krefftii (Steindachner) (Teleostei: Hemiramphidae)

The structure and function of the pharyngeal jaw apparatus (PJA) and postpharyngeal alimentary tract of Arrhamphus sclerolepis krefftii, an herbivorous hemiramphid, were investigated by dissection, light and scanning electron microscopy, and X-ray analysis of live specimens. A simple model of PJA operation is proposed, consisting of an adductive power stroke of the third pharyngobranchial that draws it posteriorly while the fifth ceratobranchial is adducted, and a return stroke in which the third pharyngobranchial bone is drawn anteriorly during abduction of the fifth ceratobranchial. Teeth in the posteromedial region of the PJA are eroded into an occlusion zone where the teeth of the third pharyngobranchial are spatulate incisiform and face posteriorly in opposition to the rostrally oriented spatulate incisiform teeth in the wear zone of the fifth ceratobranchial. The shape of the teeth and their pedestals (bone of attachment) is consistent with the model and with the forces likely to operate on the elements of the PJA during mastication. The role of pharyngeal tooth replacement in maintaining the occlusal surfaces in the PJA during growth is described. The postpharyngeal alimentary tract of A. sclerolepis krefftii comprises a stomachless cylinder that attenuates gradually as it passes straight to the anus, interrupted only by a rectal valve. The ratio of gut length to standard length is about 0.5. Despite superficial similarities to the cichlid PJA (Stiassny and Jensen [1987] Bull Mus Comp Zool 151:269-319), the hemiramphid PJA differs in the fusion of the third pharyngobranchial bones, teeth in the second pharyngobranchials and the fifth ceratobranchial face anteriorly, the presence of a slide-like diarthroses between the heads of the fourth epibranchials and the third pharyngobranchial, the occlusion zone of constantly wearing teeth, and the unusual form of the muscularis craniopharyngobranchialis. The functional relationship between these structures is explained and the consequence for the fish of a complex PJA and a simple gut is discussed.     

On the ampullary organs of the South-American paddle-fish Sorubim lima (Siluroidea,Pimelodidae)

Summary Ampullary organs were found in the epidermis of the paddle-fish Sorubim lima; they are distributed all over the skin surface of the fish but are particularly densely grouped in the head region and on the dorsal surface of the paddle. Histological and electron microscopical observations show that their structure is similar to the type of cutaneous ampullary organs characteristic of other Siluroidea. Composed of a relatively large mucus-filled ampulla, the organ possesses a short and narrow canal which leads to the outer epidermal surface. The wall of the ampulla is formed of several layers of flat epidermal cells. In general four sensory cells, each one surrounded by supporting cells, compose the sensory epithelium at the bottom of the ampulla. The inner surface of the sensory cells in contact with the ampullary mucus bears only microvilli. The contact between the nerve endings and the sensory cells show the characteristic structure of an afferent neuro-sensory junction. Two ampullae are innervated in some cases by the same afferent nerve fibre.The author expresses her gratitude to Dr. Szabo for his scientific advice during her stay in Gif sur Yvette     

Ultrastructure of the extensively developed nuchal organs of Laonice bahusiensis (Annelida: Canalipalpata: Spionidae)

The nuchal organs of annelid Laonice bahusiensis (Spionidae) from northern Europe have been studied using scanning and transmission electron microscopy. L. bahusiensis is the first spionid species in which extensively developed, continuous nuchal organs are described. The nuchal organs of this genus are the longest known among polychaete annelids. They consist of paired double bands extending from the prostomium on a mid‐dorsal caruncle for about 24–30 setigers. Their microanatomy corresponds to the general structural plan of nuchal organs: there are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving‐stone‐like cover and innervated by one pair of longitudinally elongated nuchal nerves. These findings clearly favor the hypothesis that the paired, extensively developed ciliated structures found in some Spionidae are homologous with the prostomial nuchal organs characteristic of polychaete annelids. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

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.     

Cellular immune competence of a Drosophila mutant with neoplastic hematopoietic organs

In the Tum^l mutant of Drosophila melanogaster, the larval hematopoietic organs undergo neoplastic changes and release into circulation large numbers of blood cells. The lamellocytes, and to a lesser extent the plasmatocytes from which they are derived, are the cells that encapsulate various endogenous tissues and form melanotic tumors. The mutation is temperature sensitive, with maximum gene expression manifested at 29°C. The ability of Tum^l larvae to encapsulate eggs of the wasp parasite Leptopilina heterotoma is dependent not only on temperature, with host larvae much more immune reactive at 29°C than at lower temperatures (15° or 21°C), but also on the interval of time following infection when temperature shift experiments are performed. When the shift of parasitized larvae from 21° to 29°C is delayed by 18 hr the hosts are not as immune reactive as those shifted immediately after infection. Since Tum^l larvae are potentially highly immune reactive at the time of infection (with sufficient numbers of lamellocytes in circulation to encapsulate parasites), the low degree of immune competence in hosts shifted to 29°C after 18 hr or maintained at lower temperatures suggests that the increased capacity of blood cells to react against foreign surfaces is dependent on the cells acquiring new or altered recognition and adherence properties at 29°C. The 18-hr delay may provide the parasite with an opportunity to interfere with the acquisition of these specific cellular alterations. Differential hemocyte counts from parasitized larvae show abnormally low lamellocyte counts in susceptible hosts, indicating that successfully developing parasites interfere with the differentiation of hemocytes.