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Jawed Vertebrates (jawed + vertebrate)
Selected AbstractsThe ancestral complement system in sea urchinsIMMUNOLOGICAL REVIEWS, Issue 1 2001L. Courtney Smith Summary: The origin of adaptive immunity in the vertebrates can be traced to the appearance of the ancestral RAG genes in the ancestral jawed vertebrate; however, the innate immune system is more ancient. A central subsystem within innate immunity is the complement system, which has been identified throughout and seems to be restricted to the deuterostomes. The evolutionary history of complement can be traced from the sea urchins (members of the echinoderm phylum), which have a simplified system homologous to the alternative pathway, through the agnathans (hagfish and lamprey) and the elasmobranchs (sharks and rays) to the teleosts (bony fish) and tetrapods, with increases in the numbers of complement components and duplications in complement pathways. Increasing complexity in the complement system parallels increasing complexity in the deuterostome animals. This review focuses on the simplest of the complement systems that is present in the sea urchin. Two components have been identified that show significant homology to vertebrate C3 and factor B (Bf), called SpC3 and SpBf, respectively. Sequence analysis from both molecules reveals their ancestral characteristics. Immune challenge of sea urchins indicates that SpC3 is inducible and is present in coelomic fluid (the body fluids) in relatively high concentrations, while SpBf expression is constitutive and is present in much lower concentrations. Opsonization of foreign cells and particles followed by augmented uptake by phagocytic coelomocytes appears to be a central function for this simpler complement system and important for host defense in the sea urchin. These activities are similar to some of the functions of the homologous proteins in the vertebrate complement system. The selective advantage for the ancestral deuterostome may have been the amplification feedback loop that is still of central importance in the alternative pathway of complement in higher vertebrates. Feedback loop functions would quickly coat pathogens with complement leading to phagocytosis and removal of foreign cells, a system that would be significantly more effective than an opsonin that binds upon contact as a result of simple diffusion. An understanding of the immune response of the sea urchin, an animal that is a good estimator of what the ancestral deuterostome immune system was like, will aid us in understanding how adaptive immunity might have been selected for during the early evolution of the vertebrates and how it might have been integrated into the pre-existing innate immune system that was already in place in those animals. The authors are grateful to Drs Sham Nair and Paul Gross for their critique of the manuscript and helpful suggestions. This work was supported by the National Science Foundation (MCB 9603086). [source] The adaptive phenotype of cortical thymic epithelial cellsEUROPEAN JOURNAL OF IMMUNOLOGY, Issue 4 2009Thomas Boehm Abstract Epithelial cells in the thymus are required for positive and negative selection of developing thymocytes. Although medullary epithelial cells play a major role in negative selection owing to their facility of expressing peripheral self-antigens, the adaptive features of cortical epithelial cells are largely unknown. A paper in this issue of the European Journal of Immunology shows that the putative serine protease Prss16 affects positive selection of a subset of CD4+ T cells. A survey of chordate genomes indicates that the Prss16 gene emerged in vertebrates as a paralogue of evolutionarily older members of the serine carboxypeptidase 28 family; thus, Prss16 is not associated with the appearance of the adaptive immune system and the thymus in jawed vertebrates. Nevertheless, it appears that Prss16 has later evolved as an essential contributor to the MHC class II peptide/ligand repertoire. [source] The plasticity of immunoglobulin gene systems in evolutionIMMUNOLOGICAL REVIEWS, Issue 1 2006Ellen Hsu Summary:, The mechanism of recombination-activating gene (RAG)-mediated rearrangement exists in all jawed vertebrates, but the organization and structure of immunoglobulin (Ig) genes, as they differ in fish and among fish species, reveal their capability for rapid evolution. In systems where there can exist 100 Ig loci, exon restructuring and sequence changes of the constant regions led to divergence of effector functions. Recombination among these loci created hybrid genes, the strangest of which encode variable (V) regions that function as part of secreted molecules and, as the result of an ancient translocation, are also grafted onto the T-cell receptor. Genomic changes in V-gene structure, created by RAG recombinase acting on germline recombination signal sequences, led variously to the generation of fixed receptor specificities, pseudogene templates for gene conversion, and ultimately to Ig sequences that evolved away from Ig function. The presence of so many Ig loci in fishes raises interesting questions not only as to how their regulation is achieved but also how successive whole-locus duplications are accommodated by a system whose function in other vertebrates is based on clonal antigen receptor expression. [source] The phylogenetic origins of the antigen-binding receptors and somatic diversification mechanismsIMMUNOLOGICAL REVIEWS, Issue 1 2004John P. Cannon Summary:, The adaptive immune system arose in ancestors of the jawed vertebrates approximately 500 million years ago. Homologs of immunoglobulins (Igs), T-cell antigen receptors (TCRs), major histocompatibility complex I (MHC I) and MHC II, and the recombination-activating genes (RAGs) have been identified in all extant classes of jawed vertebrates; however, no definitive homolog of any of these genes has been identified in jawless vertebrates or invertebrates. RAG-mediated recombination and associated junctional diversification of both Ig and TCR genes occurs in all jawed vertebrates. In the case of Igs, somatic variation is expanded further through class switching, gene conversion, and somatic hypermutation. Although the identity of the ,primordial' receptor that was interrupted by the recombination mechanism in jawed vertebrates may never be established, many different families of genes that exhibit predicted characteristics of such a receptor have been described both within and outside the jawed vertebrates. Recent data from various model systems point toward a continuum of immune receptor diversity, encompassing many different families of recognition molecules whose functions are integrated in an organism's response to pathogenic invasion. Various approaches, including both genomic and protein-functional analyses, currently are being applied in jawless vertebrates, protochordates, and other invertebrate deuterostome systems and may yield definitive evidence regarding the presence or absence of adaptive immune homologs in species lacking adaptive immune systems. Such studies have the potential for uncovering previously unknown mechanisms of generating receptor diversity. [source] The lectin-complement pathway , its role in innate immunity and evolutionIMMUNOLOGICAL REVIEWS, Issue 1 2004Teizo Fujita Summary:, Innate immunity was formerly thought to be a non-specific immune response characterized by phagocytosis. However, innate immunity has considerable specificity and is capable of discriminating between pathogens and self. Recognition of pathogens is mediated by a set of pattern recognition receptors, which recognize conserved pathogen-associated molecular patterns (PAMPs) shared by broad classes of microorganisms, thereby successfully defending invertebrates and vertebrates against infection. Lectins, carbohydrate-binding proteins, play an important role in innate immunity by recognizing a wide range of pathogens. Mannose-binding lectin (MBL) and ficolin are lectins composed of a lectin domain attached to collagenous region. However, they use a different lectin domain: a carbohydrate recognition domain (CRD) is responsible for MBL and a fibrinogen-like domain for ficolin. These two collagenous lectins are pattern recognition receptors, and upon recognition of the infectious agent, they trigger the activation of the lectin-complement pathway through attached serine proteases, MBL-associated serine proteases (MASPs). A similar lectin-based complement system, consisting of the lectin,protease complex and C3, is present in ascidians, our closest invertebrate relatives, and functions in an opsonic manner. We isolated several lectins homologous to MBLs and ficolins and several MASPs in invertebrates and lower vertebrates, and herein we discuss the molecular evolution of these molecules. Based on these findings, it seems likely that the complement system played a pivotal role in innate immunity before the evolution of an acquired immune system in jawed vertebrates. [source] Primitive complement system of invertebratesIMMUNOLOGICAL REVIEWS, Issue 1 2004Masaru Nonaka Summary:, Most components of the human complement system have unmistakable domain architectures, making evolutionary tracing feasible. In contrast to the major genes of the adaptive immune system, which are present only in jawed vertebrates, complement component genes with unique domain structures are present not only in jawed vertebrates but also in jawless fish and non-vertebrate deuterostomes. Recent progress in genome analysis in several eukaryotes, occupying the phylogenetically critical positions, showed that most individual domains found in the complement components are metazoa specific, being found both in deuterostomes and in protostomes but not in yeast or plant. However, unique domain architecture of complement components is not present in protostomes, suggesting that the complement system has been established in the deuterostome lineage not by invention of new domains but by innovation of unique combination of the pre-existing domains. The recently assembled Ciona intestinalis draft genome contained the most modular complement genes, except for factor I. However, some possible C. intestinalis complement components show critical structural divergence from the mammalian counterparts, casting doubt on their mutual interaction. Thus, another integrative step seems to have been required to establish the modern complement system of higher vertebrates. [source] An architectural perspective on signaling by the pre-, ,, and ,, T cell receptorsIMMUNOLOGICAL REVIEWS, Issue 1 2003Sandra M. Hayes Summary: The T cell antigen receptor (TCR) is a multimeric complex composed of an antigen-binding clonotypic heterodimer and a signal transducing complex consisting of the CD3 dimers (CD3,, and CD3,,) and a TCR-, homodimer. In all jawed vertebrates there are two T cell lineages, ,, and ,,, distinguished by the clonotypic subunits contained within their TCRs (TCR-, and -, or TCR-, and -,, respectively). A third receptor complex, the preTCR, is only expressed on immature T cells. The preTCR, which contains the invariant pre-T, (pT,) chain in lieu of TCR-,, plays a critical role in the early development of ,, lineage cells. The subunit composition of the signal transducing complexes of the pre-, ,,- and ,,TCRs was previously thought to be identical. However, recent data demonstrate that there are significant differences in the signal transducing complexes of these three TCRs. For example, ,,TCRs contain both CD3,, and CD3,, dimers, whereas ,,TCRs contain only CD3,, dimers. Moreover, preTCR function appears to be unaffected in the absence of CD3,, suggesting that CD3,, dimers are dispensable for pre-TCR assembly. In this review, we summarize current data relating to the subunit composition of the pre-, ,,- and ,,TCRs and discuss how these structural differences may impact receptor signaling and ,,/,, lineage determination. [source] Placoderm fishes, pharyngeal denticles, and the vertebrate dentitionJOURNAL OF MORPHOLOGY, Issue 3 2003Zerina Johanson Abstract The correlation of the origin of teeth with jaws in vertebrate history has recently been challenged with an alternative to the canonical view of teeth deriving from separate skin denticles. This alternative proposes that organized denticle whorls on the pharyngeal (gill) arches in the fossil jawless fish Loganellia are precursors to tooth families developing from a dental lamina along the jaw, such as those occurring in sharks, acanthodians, and bony fishes. This not only indicates that homologs of tooth families were present, but also illustrates that they possessed the relevant developmental controls, prior to the evolution of jaws. However, in the Placodermi, a phylogenetically basal group of jawed fishes, the state of pharyngeal denticles is poorly known, tooth whorls are absent, and the presence of teeth homologous to those in extant jawed fishes (Chondrichthyes + Osteichthyes) is controversial. Thus, placoderms would seem to provide little evidence for the early evolution of dentitions, or of denticle whorls, or tooth families, at the base of the clade of jawed fishes. However, organized denticles do occur at the rear of the placoderm gill chamber, but are associated with the postbranchial lamina of the anterior trunkshield, assumed to be part of the dermal cover. Significantly, these denticles have a different organization and morphology relative to the external dermal trunkshield tubercles. We propose that they represent a denticulate part of the visceral skeleton, under the influence of pharyngeal patterning controls comparable to those for pharyngeal denticles in other jawed vertebrates and Loganellia. J. Morphol. 257:289,307, 2003. © 2003 Wiley-Liss, Inc. [source] Evolution of jaw depression mechanics in aquatic vertebrates: insights from GhondrichthyesBIOLOGICAL JOURNAL OF THE LINNEAN SOCIETY, Issue 1 2000CHERYL D. WILGA The widely accepted phylogenctic position of Chondrichthyes as the sister group to all other living gnathostomes makes biomechanical analyses of this group of special significance for estimates of skull function in early jawed vertebrates. We review key findings of recent experimental research on the feeding mechanisms of living elasmobranchs with respect to our understanding of jaw depression mechanisms in gnathostome vertebrates. We introduce the possibility that the ancestral jaw depression mechanism in gnathostomes was mediated by the coracomandibularis muscle and that for hyoid depression by the coracohyoideus muscle, as in modern Chondrichthyes and possibly placoderms. This mechanism of jaw depression appears to have been replaced by the sternohyoideus (homologous to the coracohyoideus) coupling in Osteichthycs following the split of this lineage from Chondrichthyes. Concurrent with the replacement of the branchiomandibularis (homologous to the coracomandibularis) coupling by the sternohyoideus coupling as the dominant mechanism of jaw depression in Osteichthyes was the fusion and shift in attachment of the intcrhyoideus and intermandibularis muscles (producing the protractor hyoideus muscle, mistakenly refereed to as the geniohyoideus), which resulted in a more diversified role of the sternohyoideus coupling in Osteichthyes. The coracohyoideus coupling appears to have been already present in vertebrates where it functioned in hyoid depression, as in modern Chondrichthyes, before it acquired the additional role of jaw depression in Osteichthyes. [source] Developmental anatomy of lampreysBIOLOGICAL REVIEWS, Issue 1 2010Michael K. Richardson Lampreys are a group of aquatic chordates whose relationships to hagfishes and jawed vertebrates are still debated. Lamprey embryology is of interest to evolutionary biologists because it may shed light on vertebrate origins. For this and other reasons, lamprey embryology has been extensively researched by biologists from a range of disciplines. However, many of the key studies of lamprey comparative embryology are relatively inaccessible to the modern scientist. Therefore, in view of the current resurgence of interest in lamprey evolution and development, we present here a review of lamprey developmental anatomy. We identify several features of early organogenesis, including the origin of the nephric duct, that need to be re-examined with modern techniques. The homologies of several structures are also unclear, including the intriguing subendothelial pads in the heart. We hope that this review will form the basis for future studies into the phylogenetic embryology of this interesting group of animals. [source] | |||||||||||||