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The antigen-receptor genes of vertebrates are rearranged by a specialized somatic recombination mechanism in developing lymphocytes - and, unexpectedly, also in the germline of cartilaginous fishes. The recombination system that carries out these DNA rearrangements may thus be a significant evolutionary force, perhaps not limited to rearrangements at antigen-receptor loci.
In all vertebrate species examined to date, rearrangement and somatic modification of gene segmental elements that encode portions of the antigen-combining sites of immunoglobulins are integral components of the generation of antibody diversity. In the phylogenetically primitive cartilaginous fishes, gene segments encoding immunoglobulin heavy and light chain loci are arranged in multiple clusters, in which segmental elements are separated by only 300-400 bp. In some cases, segmental elements are joined in the germline of nonlymphoid cells (joined genes). Both genomic library screening and direct amplification of genomic DNA have been used to characterize at least 89 different type I light chain gene clusters in the skate, Raja. Analyses of predicted nucleotide sequences and predicted peptide structures are consistent with the distribution of genes into different sequence groups. Predicted amino acid sequence differences are preferentially distributed in complementarity-determining versus framework regions, and replacement-type substitutions exceed neutral substitutions. When specific germline sequences are related to the sequences of individual cDNAs, it is apparent that the joined genes are expressed and are potentially somatically mutated. No evidence was found for the presence of any type I light chain gene in Raja that is not germline joined. The type I light chain gene clusters in Raja appear to represent a novel gene system in which combinatorial and junctional diversity are absent.
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This study investigated the nature of vasodilator mechanisms in the dorsal aorta of the giant shovelnose ray, Rhinobatus typus. Anatomical techniques found no evidence for an endothelial nitric oxide synthase, but neural nitric oxide synthase was found to be present in the perivascular nerve fibres of the dorsal aorta and other arteries and veins using both NADPH-diaphorase staining and immunohistochemistry with a specific neural NOS antibody. Arteries and veins both contained large nNOS-positive nerve trunks from which smaller nNOS-positive bundles branched and formed a plexus in the vessel wall. Single, varicose nNOS-positive nerve fibres were present in both arteries and veins. Within the large bundles of both arteries and veins, groups of nNOS-positive cell bodies forming microganglia were observed. Double-labelling immunohistochemistry using an antibody to tyrosine hydroxylase showed that nearly all the NOS nerves were not sympathetic. Acetylcholine always caused constriction of isolated rings of the dorsal aorta and the nitric oxide donor, sodium nitroprusside, did not mediate any dilation. Addition of nicotine (3 x 10(-4) M) to preconstricted rings caused a vasodilation that was not affected by the nitric oxide synthase inhibitor, L-NNA (10(-4) M), nor the soluble guanylyl cyclase inhibitor, ODQ (10(-5) M). This nicotine-mediated vasodilation was, therefore, not due to the synthesis and release of NO. Disruption of the endothelium significantly reduced or eliminated the nicotine-mediated vasodilation. In addition, indomethacin (10(-5) M), an inhibitor of cyclooxygenases, significantly increased the time period to maximal dilation and reduced, but did not completely inhibit the nicotine-mediated vasodilation. These data support the hypothesis that a prostaglandin is released from the vascular endothelium of a batoid ray, as has been described previously in other groups of fishes. The function of the nitrergic innervation of the blood vessels is not known because nitric oxide does not appear to regulate vascular tone.
The intracellular localization of a tetraphenylporphyrin bearing four long lipophilic alkyl chains in electrocytes from Psammobatis extenta(Rajidae) is described. In contrast to what is usually the case, this porphyrin derivative is localized in the electromotor nerves and the nuclear chromatin of the electrocytes. Both structures exhibited intense fluorescence, whereas, the mitochondria were only slightly fluorescent. These data are discussed in relation to electrocyte death in a weakly electric fish. Additionally, electron probe X-ray microanalysis suggests a migration of chloride and cationic ions, which might be implicated in chloride and cationic channel activation in the electrocyte.
1. The phospholipid composition of the main proteolipid complexes of the nervous system was studied in myelin and synaptosomal membranes from brains of representatives of various vertebrate classes. 2. The relative content of acid phospholipids was much higher in proteolipid complexes from myelin and synaptosomal membranes of all vertebrates studied as compared to their content in the initial lipid extract (28-80% and 11-20% of total phospholipid content, respectively). 3. The relative content of acid phospholipids in proteolipid complexes of myelin membranes was much lower in brain of fishes and amphibia as compared to higher vertebrates. 4. The main acid phospholipids of proteolipid complexes was phosphatidylserine, phosphatidic acid being characteristic for myelin proteolipids and diphosphatidyl glycerol for synaptosomal proteolipids of all vertebrates studied.
In mammals, the gastric H(+)-K(+)-ATPase (HKalpha1) mediates acid secretion in the stomach and kidneys. Like mammals, elasmobranchs also secrete acid from their stomachs, but unlike mammals they primarily use their gills for systemic acid excretion instead of their kidneys. The purpose of this study was to determine if an HKalpha1 orthologue exists in an elasmobranch (Atlantic stingray, Dasyatis sabina), to determine if it is expressed in gills and, if so, to localize its expression and determine if its expression is regulated during hypercapnia or freshwater acclimation. A polyclonal antibody made against an HKalpha1 peptide detected HKalpha1 immunoreactivity in protein isolates and tissue sections of stingray stomachs and gills. Immunohistochemistry demonstrated that HKalpha1 immunoreactivity was present in a subpopulation of epithelial cells in both organs. Double-labeling experiments in the gills showed that HKalpha1 immunoreactivity occurred in Na(+)-K(+)-ATPase-rich cells and not in V-type H(+)-ATPase-rich cells. RT-PCRs were used to deduce the primary sequence of a putative H(+)-K(+)-ATPase from the stomach of Atlantic stingrays. The 3,421-base pair cDNA includes a coding region for a 1,025-amino acid protein that is over 80% identical to HKalpha1 of mammals. RT-PCRs were then used to demonstrate that this transcript is also expressed in the gills. To our knowledge, this is the first H(+)-K(+)-ATPase sequence reported for any elasmobranch and the first full-length sequence for any fish. We also provide the first evidence for its expression in the gills of any fish and demonstrate that its expression increased during freshwater acclimation but not exposure to hypercapnia.
It has been shown that the dependence of glycogenphosphorylase kinase activity on the quantity of glycogen added corresponds to the glycogenphosphorylase saturation curve by glycogen obtained during investigation of complex formation with the muscle glycogenphosphorylase of the animals: rat, frog Rana temporaria and fish Raja clavata under identical conditions. We suggest that glycogenolytic enzymes organize regular structure on glycogen particle. Glycogenphosphorylases of heat-loving animals have high affinity to glycogen that is accounted for necessity to keep the structure in natural conditions.
An immunohistochemical study was made of the octavolateral centres in the medulla of a cartilaginous fish, using an antibody specific for dopamine. No immunopositive cell bodies were observed but dopaminergic fibres were present in parts of the region. The overlying cerebellar crest was devoid of dopamine; the dorsal (electroreceptive) nucleus was poorly innervated, but the medial (mechanoreceptive) nucleus received a rich dopaminergic innervation comprising very fine, varicose fibres.
The white-edge freshwater whip ray Himantura signifer is ammonotelic in freshwater, but retains the capacities of urea synthesis and ureosmotic osmoregulation to survive in brackish water. The first objective of this study was to examine whether exposure to brackish water would lead to increases in food intake, and/or conservation of nitrogen in H. signifer upon daily feeding. Results obtained showed that a progressive increase in ambient salinity, from 1 per thousand to 15 per thousand over a 10-day period, did not lead to an increase in daily food intake. However, there were significant reductions in daily rates of ammonia and urea excretion in H. signifer during salinity changes, especially between day 5 (in 10 per thousand water) and day 10 (in 15 per thousand water) when compared to those of the control kept in 1 per thousand water. Consequently, there was a significant decrease in the percentage of nitrogen (N) from the food being excreted as nitrogenous waste (ammonia-N+urea-N) during this period. On day 10, the tissue urea contents in fish exposed to 15 per thousand water were significantly greater than those of fish kept in 1 per thousand water, and the excess urea-N accumulated in the former fish could totally account for the cumulative deficit in excretion of urea-N+ammonia-N during the 10-day period. Thus, it can be concluded that H. signifer is N-limited, and conserved more N from food when exposed to brackish water. The conserved N was converted to urea, which was retained in tissues for osmoregulation. The second objective of this study was to elucidate whether the retention of the capacity of N conservation in H. signifer would lead to an accumulation of urea in fish exposed to not only 15 per thousand water, but also 1 per thousand water, upon feeding. For fish pre-acclimated to 1 per thousand water or 15 per thousand water for 10 days and then fasted for 48 h, the rate of ammonia excretion in fish exposed to 15 per thousand water was consistently lower than that of fish exposed to 1 per thousand water, throughout the 36-h post-feeding period. In addition, the hourly rate of urea excretion in the former was significantly lower than that of the latter between hours 12 and 36. There were postprandial increases in ammonia contents in the muscle, liver, stomach, intestine, brain and plasma of fish kept in 1 per thousand water; but postprandial increases in ammonia occurred only in the liver and brain of fish exposed to 15 per thousand water, and the magnitudes of increases in the latter were smaller than those in the former. Indeed, postprandial increases in tissue urea contents occurred in both groups of fish, but the greatest increase in urea content was observed in the muscle of fish exposed to 15 per thousand water. Taken together, these results indicate that H. signifer in freshwater could be confronted with postprandial osmotic stress because of its capacity of conserving N and increasing urea synthesis upon feeding.
A previously healthy adult male suffered a wound to the dorsal ulnar aspect of his right hand by a stingray barb while fishing off the East coast of Florida. Two weeks after the imbedded barb had been surgically removed, an erythematous lesion developed around the wound. Histopathologic and microbiological studies revealed infection caused by Fusarium solani. The patient was successfully treated with debridement and skin grafting in conjunction with ketoconazole therapy.
The mapping of Hox clusters and many duplicated genes in zebrafish indicated an extra whole-genome duplication in ray-fined fish. However, to reconstruct the preduplication chromosomes (proto-chromosomes), the comparative genomic studies of more distantly related teleosts are essential. Medaka and zebrafish are ideal for this purpose, because their lineages separated from their last common ancestor approximately 140 million years ago. To reconstruct ancient vertebrate chromosomes, including the chromosomes of the vertebrate ancestor of humans from 450 million years ago, we mapped 818 genes and expressed sequence tags (ESTs) on a single meiotic backcross panel obtained from inbred strains of the medaka, Oryzias latipes. Comparisons of linkage relationships of orthologous genes among three species of vertebrates (medaka, zebrafish, and human) indicate the number and content of the chromosomes of the last common ancestor of ray-fined fish and lobe-fined fish (including humans), and the extra whole genome duplication event in the ray-fin lineage occurred in the common ancestor of perhaps all teleosts.
The genetic mechanisms regulating tetrapod limb development are well characterized, but how they were assembled during evolution and their function in basal vertebrates is poorly understood. Initial studies report that chondrichthyans, the most primitive extant vertebrates with paired appendages, differ from ray-finned fish and tetrapods in having Sonic hedgehog (Shh)-independent patterning of the appendage skeleton. Here we demonstrate that chondrichthyans share patterns of appendage Shh expression, Shh appendage-specific regulatory DNA, and Shh function with ray-finned fish and tetrapods. These studies demonstrate that some aspects of Shh function are deeply conserved in vertebrate phylogeny, but also highlight how the evolution of Shh regulation may underlie major morphological changes during appendage evolution.
The objective of this study was to elucidate whether the marine blue-spotted fantail ray, Taeniura lymma, and the freshwater white-edge whip ray, Himantura signifer, injected with NH(4)Cl intraperitoneally would excrete the majority of the excess ammonia as ammonia per se to ameliorate ammonia toxicity despite being ureogenic. To examine the roles of urea and the ornithine-urea cycle, experimental fishes were exposed to salinity changes after being injected with NH(4)Cl. The ammonia excretion rates of the marine ray, T. lymma, injected with NH(4)Cl followed by exposure to seawater (30 per thousand) or diluted seawater (25 per thousand) increased 13-fold and 10-fold, respectively, within the first 3 h. Consequently, the respective percentage of nitrogenous wastes excreted as ammonia were 55% and 65% compared with 21% of the saline-injected control, indicating that T. lymma became apparently ammonotelic after injection with NH(4)Cl. By hour 6, large portions (70%-85%) of the ammonia injected into T. lymma exposed to seawater or diluted seawater had been excreted, and T. lymma excreted much more nitrogenous wastes (135%-180%), in excess of the ammonia injected into the fish, during the 24-h period. For T. lymma exposed to seawater, a small portion (30%) of the ammonia injected into the fish was detoxified to urea during the first 6 h, but there was an apparent suppression of urea synthesis thereafter, contributing partially to the large decrease (19%) in urea contents in its muscle at hour 24. A major contributing factor to the decrease in urea content was a reduction in ammonia production, as indicated by a large deficit between urea loss in the muscle and excess ammonia accumulated plus excess nitrogen excreted in the experimental fish. The freshwater ray, H. signifer, injected with NH(4)Cl followed by exposure to freshwater (0.7 per thousand) or brackish water (10 per thousand) was capable of excreting all the ammonia injected into the body, mainly as ammonia, within 12 h. Like T. lymma, it also excreted the injected ammonia mainly as ammonia during the first 3 h postinjection. During this period, the percentage of the injected ammonia excreted in fish exposed to brackish water (28.4%+/-4.6%) was significantly lower than those exposed to freshwater (56.1%+/-8.26%). In contrast, the percentage of nitrogenous wastes being excreted as urea in the former (38.4%) was significantly greater than that in the latter (14.1%). These results suggest that a portion of the ammonia injected into the fish was turned into urea, and urea synthesis was increased transiently in fish exposed to brackish water during the initial postinjection period. However, urea was not retained effectively by H. signifer. Taken together, these results suggest that the primary function of the ornithine-urea cycle in ureogenic marine and freshwater elasmobranchs is to synthesize urea for osmotic water retention and not for ammonia detoxification.
A repetitive HindIII fragment of DNA from Raja montagui (Rajiformes) was cloned and sequenced for the first time in cartilaginous fishes. This element, which comprises approximately 5% of the whole genome of the spotted ray, is absent in long tandem arrays, being typical of satellite DNA. It appeared constituted by 311 AT-rich bp (61%). The clone was hybridized to the genomic DNA of species with varying phyletic distances, revealing a high degree of conservation.
The morphological and functional characteristics of stingray liver were studied, including the effect of ischaemia/reperfusion. With an isolated perfused model, it was shown that the stingray liver was more resistant than the rat liver to ischaemia/reperfusion injury; this was consistent with the differing partial oxygen tensions usually present in the two species. This study confirmed that whereas stingray hepatocytes form tubules with central bile canaliculi as in other fish, the stingray liver has portal triads and a lobular architecture as in mammals. Apoptosis of hepatocytes, demonstrated in the normal liver, was only marginally enhanced by ischaemia/reperfusion. Resulting apoptotic bodies were phagocytized by macrophage-like cells in hepatocyte tubules. In contrast to rat liver, the stingray liver showed no necrosis after ischaemia-reperfusion.
Analysis of human and mouse immunoglobulins has shown that five of six hypervariable regions that form the antigen binding site have a small repertoire of main chain conformations (canonical structures). Cartilaginous fishes are the most distantly related species to humans known to have an immune system, their evolutionary lines having diverged 450 million years ago. An analysis of VH and V kappa sequences from these fishes shows that all the main chain structures in their L1, L2, H1 and H2 hypervariable regions, and one of those in the L3 region, are the same as those most commonly found in human and mouse. This implies that the canonical structures occurring most commonly in hypervariable regions arose very early in the stages of the evolution of the immune system.
To investigate for the first time the effects of ionizing radiation on thymus of a representative cartilaginous fish, juvenile clearnose skates, Raja eglanteria, were exposed to 0-75 Gy of X-radiation and sacrificed after 12 days. Morphometrics (weight, disc width and total length) and thymus and thymic cyst area were compared to controls using ANOVA. Thymus area declined logarithmically and medullary cysts increased as a function of dose (P < or = 0.05). To assess thymic recovery, skates were exposed to 0, 9, 13.5 or 18 Gy of X-radiation and sacrificed when moribund or on days 10, 20, 30 and 40 post-irradiation. Complete restoration of the thymus was not achieved during the 40-day observation period, although repopulation with pro-thymocytes and partial recovery of thymic architecture were evident histologically. The observed high radiosensitivity of R. eglanteria thymocytes was similar to responses of other vertebrates, but recovery time was prolonged.