Renin-angiotensin system in antarctic fishes

• 1.1. The presence of a renin-angiotensin-like system has been investigated in the Antarctic fishes Chionodraco hamatus (Fam. Channichthydae) and Pagothenia (Trematomus) bernacchii (Fam. Notothenidae).
• 2.2. A renin-like activity is present in plasma and kidney of both the white blooded (Chionodraco) and the red blooded (Pagothenia) species.
• 3.3. An angiotensin converting enzyme-like activity has been demonstrated in plasma, gills and kidneys of both species. The activity is inhibited by high temperature.
• 4.4. From our data a renin-angiotensin-like system is present in the Antarctic fishes studied but the cascade of enzymes is active only at low temperatures.

     

Ocular morphology in antarctic notothenioid fishes

Beneath the sea ice at McMurdo Sound, Antarctica, notothenioid fishes are subject to extreme seasonal variation in the annual light cycle including 4 months of continual darkness. Gross and microscopic anatomy of the eyes of 18 species revealed ocular morphology that was generally similar to that of coastal fishes elsewhere in the world, and unlike that of deep sea fishes living in perpetual darkness. The spectacle was well developed as were hyaloid arteries at the vitreoretinal interface. Fourteen species had a choroid body, and its presence was considered a primitive character state for notothenioids. The choroid body was absent in phyletically derived groups. The choroid body was especially large in Dissostichus mawsoni, the only species with a rod dominated retina. Retinae were 154–279 μm thick with layering and sublayering typical for teleosts. Although all species had both rods and cones, there was marked interspecific variation in the ratio of cones:rods and in the total number of visual cells. Non-Antarctic notothenioids from New Zealand had more visual cells than most species from McMurdo Sound. Retinae appeared balanced for vision under dim but seasonally variable light conditions and not specially adapted to the 4-month period of winter darkness. Retinal histology reflected the ecology and depth range of most species. Based on ecology and retinal histology, four groups of species were recognized: (1) Non-Antarctic, (2) cryopelagic (including two visually oriented benthic species), (3) pelagic and benthopelagic, and (4) benthic.     

Evolution and adaptive radiation of antarctic fishes

There are few instances where a knowledge of the thermal physiology, habitats and lifestyles of a group of closely related species can be mapped onto a well-supported phylogeny and a detailed climatic history. The unique fish fauna of the Southern Ocean, dominated by a single group of fish whose phylogeny is known from traditional and molecular techniques, provides one such opportunity. Furthermore, these fish are living at an extreme temperature for marine organisms. Physiological and molecular studies are revealing details of the mechanisms of temperature compensation and, combined with knowledge of the thermal history, are throwing new light on the process of evolution in this unique group of fish.     

Adaptive evolution of hepcidin genes in antarctic notothenioid fishes

Hepcidin is a small bioactive peptide with dual roles as an antimicrobial peptide and as the principal hormonal regulator of iron homeostasis in human and mouse. Hepcidin homologs of very similar structures are found in lower vertebrates, all comprise approximately 20-25 amino acids with 8 highly conserved cysteines forming 4 intramolecular disulfide bonds, giving hepcidin a hairpin structure. Hepcidins are particularly diverse in teleost fishes, which may be related to the diversity of aquatic environments with varying degree of pathogen challenge, oxygenation, and iron concentration, factors known to alter hepcidin expression in mammals. We characterized the diversity of hepcidin genes of the Antarctic notothenioid fishes that are endemic to the world's coldest and most oxygen-rich marine water. Notothenioid fishes have at least 4 hepcidin variants, in 2 distinctive structural types. Type I hepcidins comprise 3 distinct variants that are homologs of the widespread 8-cysteine hepcidins. Type II is a novel 4-cysteine variant and therefore only 2 possible disulfide bonds, highly expressed in hematopoietic tissues. Analyses of d(N)/d(S) substitution rate ratios and likelihood ratio test under site-specific models detected significant signal of positive Darwinian selection on the mature hepcidin-coding sequence, suggesting adaptive evolution of notothenioid hepcidins. Genomic polymerase chain reaction and Southern hybridization showed that the novel type II hepcidin occurs exclusively in lineages of the Antarctic notothenioid radiation but not in the basal non-Antarctic taxa, and lineage-specific positive selection was detected on the branch leading to the type II hepcidin clade under branch-site models, suggesting adaptive evolution of the reduced cysteine variant in response to the polar environment. We also isolated a structurally distinct 4-cysteine (4cys) hepcidin from an Antarctic eelpout that is unrelated to the notothenioids but inhabits the same freezing water. Neighbor-Joining (NJ) analyses of teleost hepcidins showed that the eelpout 4cys variant arose independently from the notothenioid version, which lends support to adaptive evolution of reduced cysteine hepcidin variants on cold selection. The NJ tree also showed taxonomic-specific expansions of hepcidin variants, indicating that duplication and diversification of hepcidin genes play important roles in evolutionary response to diverse ecological conditions.     

Divergence in skeletal mass and bone morphology in antarctic notothenioid fishes

Although notothenioid fishes lack swim bladders, some species live temporarily or permanently in the water column. Given its relatively high density, skeletal mass is a key determinant of buoyancy. Notothenioids have reduced skeletal ossification, but there is little quantitative data on the phylogenetic distribution of this trait. We obtained dry skeletal masses for 54 specimens representing 20 species from six notothenioid families. Although comparative data are sparse, notothenioid skeletons comprise a smaller percentage of body mass, <3.5%, than those of three non‐notothenioid perciforms. With relatively high skeletal mass, the non‐Antarctic Bovichtus diacanthus is similar in skeletal mass to some non‐notothenioids. Eleginops maclovinus, the non‐Antarctic sister group of the Antarctic clade, has a relatively light skeleton (<2% of body mass) similar to many species in the Antarctic clade. Low skeletal mass is therefore a synapomorphy shared by Eleginops plus the Antarctic clade. We provide gross, histological, and micro‐CT documentation of the structure and location of bone and cartilage in skulls, pectoral girdles, and vertebrae, with emphasis on the bovichtid B. diacanthus, the eleginopsid E. maclovinus, and the channichthyid Chaenodraco wilsoni. In Eleginops and the Antarctic clade, most bone is spongy and most species have persisting cartilage in the skull and appendicular skeleton. We also measured the relative size of the notochordal canal in adult vertebral centra of 38 species representing all eight families. There is considerable interspecific variation in this pedomorphic trait and all species show an ontogenetic reduction in the relative size of the canal. However, large persisting canals are present in adults of the Antarctic clade, especially in the nototheniids Pleuragramma and Aethotaxis and in a number of bathydraconid and channichthyid genera. J. Morphol. 275:841–861, 2014. © 2014 Wiley Periodicals, Inc.     

Nervous and sensory system correlates of an epibenthic evolutionary radiation in antarctic notothenioid fishes, genus Trematomus (Perciformes; Nototheniidae)

The perciform suborder Notothenioidei consists of 120 species, with 94 confined to the Antarctic Region of the Southern Ocean. On the Antarctic shelf, this phyletic radiation has been accompanied by a substantial morphological and ecological diversification towards a pelagic existence. For example, the primarily benthic genus Trematomus contains an epibenthic radiation that includes T. loennbergii, T. lepidorhinus, and T. eulepidotus. By comparing these epibenthic species with three congeneric benthic species (T. scotti, T. pennellii, and T. bernacchii) we tested three null hypotheses regarding brain variation in Antarctic trematomids: 1) that there is no difference in brain morphology among the six species; 2) that phylogenetic and ecological factors do not influence brain morphology; and 3) that peripheral sensory structures do not influence brain morphology. We rejected each of these hypotheses, leading us to conclude that Trematomus brains vary interspecifically, between benthic and epibenthic species, and with a species' depth distribution. Further, we conclude that brain variation is correlated with differences in peripheral sensory systems and motor activity. Specifically, epibenthic Trematomus have larger percentages of their brain volume devoted to lateral line mechanoreceptive and motor (cerebellar) structures. Species living at greater depths have low ratios of cones:rods in the retina and larger olfactory structures.     

Wake dynamics and locomotor function in fishes: interpreting evolutionary patterns in pectoral fin design

The great anatomical diversification of paired fins within theActinopterygii (ray-finned fishes) can be understood as a suiteof evolutionary transformations in design. At a broad taxonomicscale, two clear trends exist in the morphology of the anteriorlysituated pectoral fins. In comparing basal to more derived clades,there are general patterns of (i) reorientation of the pectoralfin base from a nearly horizontal to more vertical inclination,and (ii) migration of the pectoral fin from a ventral to mid-dorsalbody position. As yet, the functional significance of thesehistorical trends in pectoral fin design remains largely untestedby experiment. In this paper we test the proposal that variationin pectoral fin structure has an important influence on themagnitude and orientation of fluid forces generated during maneuveringlocomotion. Using digital particle image velocimetry for quantitativewake visualization, we measure swimming forces in ray-finnedfishes exhibiting the plesiomorphic and apomorphic pectoralfin anatomy. Our experiments focus on rainbow trout (Oncorhynchusmykiss), a lower teleost with pectoral fins positioned ventrallyand with nearly horizontally inclined fin bases, and bluegillsunfish (Lepomis macrochirus), a relatively derived perciformfish with more vertically oriented pectoral fins positionedmid-dorsally on the body. In support of hypotheses arising fromour prior wake studies and previously untested models in theliterature, we find that the pectoral fins of sunfish generatesignificantly higher forces for turning and direct braking forcescloser to the center of mass of the body than the pectoral finsof trout. These results provide insight into the hydrodynamicimportance of major evolutionary transformations in pectoralfin morphology within the Actinopterygii.     

Molecular evolution of hemoglobins of antarctic fishes (Notothenioidei)

Amino acid sequences of α- and β-chains of human hemoglobin and of hemoglobins of coelacanth and 24 teleost fish species, including 11 antarctic and two temperate Notothenioidei, were analyzed using maximum parsimony. Trees were derived for the α- and β-chains separately and for tandemly arranged sequences, using the human and coelacanth sequences as outgroups in all analyses. The topologies of the trees of the α-and β-chains are highly congruent and indicate a specific pattern of gene duplications and gene expression of teleost hemoglobins which has not yet been investigated into more detail. The Notothenioid fish generally contain a single major hemoglobin and often a second minor component. The α- and β-chains of the major components form a monophyletic group in all investigated trees, with the nonantarctic Pseudaphritis as their sister taxon. The minor chains also are a monophyletic group and form an unresolved cluster with the major chains and the hemoglobins of tuna and red gurnard. The Notothenioid families Nototheniidae and Bathydraconidae appear to be paraphyletic. Received: 26 March 1997 / Accepted: 7 May 1997     

The phylogeny of polar fishes and the structure,function and molecular evolution of hemoglobin

Fishes thriving in polar habitats offer many opportunities for comparative approaches to understanding protein thermal adaptations. Investigations on the remarkable evolutionary adaptations to these environments of basic proteins such as hemoglobin, the oxygen carrier, can provide new insights into the mechanisms studied in temperate organisms and can shed light on convergent processes evolved in response to thermal adaptations. At the molecular level, hemoglobins are one of the most intriguing systems for studying the relationships between environmental conditions and adaptations. This review summarizes the current knowledge on molecular structure, biological function and phylogeny of hemoglobins of fish species living in both polar habitats but having different evolutionary histories. In benthic, non-migratory, cold-adapted fishes, the stability of thermal conditions may have generated no or few variations in selective pressures on globin sequences through evolutionary time, so that sequences retain the species phylogenetic “signal”. In pelagic, migratory, cold-adapted or temperate fishes, variations in selective pressures on globin sequences caused by variations in temperature accompanying the dynamic life style may have disrupted the phylogenetic “signal” in phenetic trees.     

The cryoprotective effect of antifreeze glycopeptides from antarctic fishes.

Apparently vitrified cells and tissues often fail to survive, probably from damage from growth of microscopically invisible ice crystals. Special biological antifreezes from some polar fishes have been shown to adsorb to specific faces of ice crystals and inhibit crystal growth. Vitrification in the presence of antifreezes therefore may help enhance postvitrification viability of cells and tissues. We report here that the addition of fish antifreeze glycopeptides (AFGPs) to vitrifying solutions increases post-thaw viability in cultured immature pig oocytes and two-cell stage embryos of mice and pigs after rapid cooling to cryogenic temperatures. The criterion for viability is maturation to metaphase for the oocytes and the ability to develop into the four-cell stage for the pig embryo and the blastocyst stage for the mouse embryo. Without AFGPs, or with addition of antifreeze peptides (AFPs), the particular vitrifying solution and cooling/warming/culturing regime used in this study produced zero viability. In the presence of the AFGPs (40 mg/ml), survival of pig oocytes and embryos was increased to about 25%, and that of mouse embryos to 82%. Dose-response studies for the mouse embryos showed that the protective effect of AFGPs shows saturation kinetics and levels off at 20 mg/ml. The AFGPs appeared to preserve cell membrane structural integrity; however, an intact cell membrane did not always lead to viability. The absence of protective effect by AFPs suggests that protection by the AFGPs is unrelated to their common antifreeze property, i.e., inhibition of ice crystal growth, but probably results from interaction with and stabilization of the cell membranes unique to the AFGPs.     

The physiological function of the coelom in starfish larvae and its evolutionary implications

The coelom in the bipinnaria larva of Asterias acts as a buoyancy tank. The concentrations of magnesium and sulphate in the coelomic fluid are lower than in seawater, reducing the density. The coelomic epithelium is a secretory epithelium, probably secreting sodium or chloride ions that then draw in the counter ion and water. The rate of urine production is very high for an isotonic marine animal, compensating for the large surface/volume ratio of the coelom. This function would account for the precocious development of the coelom and its association with an excretory duct. It is proposed that the coeloms of other pelagic larvae such as the actinotroch of Phoronis and of echiuran larvae have a similar function and that this may have been an original function of the coelom, although in many phyla this function has been modified or lost.     

Ecological and behavioural correlates of intracellular buffering capacity in the muscles of antarctic fishes

Summary Five species of antarctic fishes can be arranged in order of increasing anaerobic capacity of the white muscles for burst swimming: Rhigophila dearborni (Zoarcidae), icefish (Channichthyidae), Dissostichus mawsoni, Trematomus centronotus, and Pagothenia borchgrevinki (Nototheniidae). This order reflects increasing dependence on anaerobic work done during short bursts of speed during prey capture or predator avoidance. Buffer capacity () for white muscle was lower than that of behaviourally equivalent fish from lower latitudes and is itself temperature-dependent.     

Diversification of brain morphology in antarctic notothenioid fishes: Basic descriptions and ecological considerations

The Notothenioidei, a perciform suborder of 120 species, dominates the ichthyofauna of the Southern Ocean around Antarctica. Unlike most teleost groups, notothenioids have undergone a corresponding ecological and phyletic diversification and therefore provide an excellent opportunity to study the divergence of the nervous system in an unusual environment. Our goal is to evaluate notothenioid brain variation in light of this diversification. To provide a baseline morphology, we examine the gross morphology and histology of the brain of Trematomus bernacchii, a generalized member of the family Nototheniidae. We then examine the variation in brain gross anatomy (32 species) and histology (10 species) of other notothenioids. Our sample represents about 27% of the species in this group and includes species from each of the six families, as well as species representing diverse ecologies. For comparison we reference the well-studied brains of two species of temperate perciformes (Perca flavescens and Lepomis humilis). Our results show that, in general, notothenioid brains are more similar to the brains of temperate perciforms than to the unusual brains of cave-dwelling and deep-sea fishes. Interspecific variation in gross brain morphology is comparable to that in Old World cyprinids and is illustrated for 17 species. Variation is especially noteworthy in the ecologically and geographically diverse family Nototheniidae. Measurements indicate that sensory regions (olfactory bulbs, eminentia granularis, and crista cerebellaris) exhibit the most pronounced variation in relative surface area. Association areas, including the corpus cerebelli and the telencephalon, exhibit moderate variation in size, shape, and lobation patterns. Regulatory areas of the brain, including the saccus vasculosus and the subependyma of the third ventricle, are also variable. These regions are best developed in species living in the subfreezing water close to the continent. In some species the expanded ependymal lining forms ventricular sacs, not previously described in any other vertebrate. Three species, including two nototheniids (Eleginops maclovinus and Pleuragramma antarcticum) and the only artedidraconid in our sample, have distinctive brains. The unique brain morphology of Pleuragramma is probably related to a sensory (lateral line) specialization for feeding. Within the Nototheniidae, a phyletic effect on cerebellar morphology is evident in the Coriiceps group and in the Pleuragramminae. Neither phyletic position nor ecological factors (water temperature, position in the water column, dietary habits) alone fully expalin the pattern of notothenioid brain diversification. © 1995 Wiley-Liss, Inc.     

Local phylogenetic divergence and global evolutionary convergence of skull function in reef fishes of the family Labridae

The Labridae is one of the most structurally and functionally diversified fish families on coral and rocky reefs around the world, providing a compelling system for examination of evolutionary patterns of functional change. Labrid fishes have evolved a diverse array of skull forms for feeding on prey ranging from molluscs, crustaceans, plankton, detritus, algae, coral and other fishes. The species richness and diversity of feeding ecology in the Labridae make this group a marine analogue to the cichlid fishes. Despite the importance of labrids to coastal reef ecology, we lack evolutionary analysis of feeding biomechanics among labrids. Here, we combine a molecular phylogeny of the Labridae with the biomechanics of skull function to reveal a broad pattern of repeated convergence in labrid feeding systems. Mechanically fast jaw systems have evolved independently at least 14 times from ancestors with forceful jaws. A repeated phylogenetic pattern of functional divergence in local regions of the labrid tree produces an emergent family-wide pattern of global convergence in jaw function. Divergence of close relatives, convergence among higher clades and several unusual 'breakthroughs' in skull function characterize the evolution of functional complexity in one of the most diverse groups of reef fishes.     

Comparative RNomics and modomics in Mollicutes: prediction of gene function and evolutionary implications

Stable RNAs are central to protein synthesis. Ribosomal RNAs make the core of the ribosome and provide the scaffold for accurate translation of mRNAs by a set of tRNA molecules each carrying an activated amino acid. To fulfill these important cellular functions, both rRNA and tRNA molecules require more than the four canonical bases and have recruited enzymes that introduce numerous modifications on nucleosides. Mollicutes are parasitic unicellular bacteria that originated from gram-positive bacteria by considerably reducing their genome, reaching a minimal size of 480 kb in Mycoplasma genitalium. By analyzing the complete set of tRNA isoacceptors (tRNomics) and predicting the tRNA/rRNA modification enzymes (Modomics) among all sequenced Mollicutes (15 in all), our goal is to predict the minimal set of RNA modifications needed to sustain accurate translation of the cell's genetic information. Building on the known phylogenetic relationship of the 15 Mollicutes analyzed, we demonstrate that the solutions to reducing the RNA component of the translation apparatus vary from one Mollicute to the other and often rely on co-evolution of specific tRNA isoacceptors and RNA modification enzymes. This analysis also reveals that only a few modification enzymes acting on nucleotides of the anticodon loop in tRNA (the wobble position 34 as well as in position 37, 3'-adjacent to anticodon) and of the peptidyltransferase center of 23S rRNA appear to be absolutely essential and resistant to gene loss during the evolutionary process of genome reduction.