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1.	Anderson, WG, et al. (författare) Purification, characterization, and biological activity of insulins from dogfish, Scyliohinus canicula, and the hammerhead shark, Sphyrna lewini. 2002 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 126:1, s. 113-122 Tidskriftsartikel (refereegranskat)abstract Insulin was purified from pancreatic extracts of two elasmobranch species belonging to different families in the order Carcharhiniformes, the European spotted dogfish, Scyliorhinus canicula (Scyliorhinidae), and the hammerhead shark, Sphyrna lewini (Carcharhinidae). The amino acid sequence of dogfish insulin was established as A-chain GIVDHCCRNT(10) CSLYDLEGYC(20)NQ and B-chain LPSQHLCGSH(10)LVETLYFVCG(20)QKGFYYVPKV(30). The primary structure of hammerhead shark insulin was similar to that of dogfish insulin with only 2 amino acid substitutions at A8 (R --> H) and B30 (V --> I). The elasmobranch insulins were markedly different from human insulin (17 amino acid substitutions) but all the residues in human insulin that are believed to be important in determining the receptor binding conformation (B6, B8, B11, B13, B23, B24, B25, A2, A3, and A19) have been conserved in the elasmobranch insulins with the exception of the conservative substitution Phe --> Tyr at B25. Consistent with this, dogfish and human insulin showed almost identical binding affinity to the recombinant solubilized human insulin receptor (K-D values of 14.0 and 18.6 pM, respectively; relative potency 133%). Previous studies have shown that bovine insulin produces severe and sustained hypoglycemia in elasmobranchs but the effect is of slow onset. Bolus arterial injections of dogfish insulin (10 nmol (.) kg(-1)) into unanesthetized, fasting dogfish (n = 9) produced no changes in blood glucose, 3-hydroxybutyrate, and acetoacetate concentrations over a 4-h period. In a second series of experiments (n = 7), dogfish insulin (10 nmol (.) kg(-1)) produced a significant (P < 0.05) fall in blood glucose after 12 h that persisted for at least 48 h, but no change in ketone body concentrations. The data indicate that the metabolic actions of an endogenous elasmobranch insulin in an elasmobranch are similar to those previously described for mammalian insulin. (C) 2002 Elsevier Science (USA).
2.	Björnsson, Björn Thrandur, 1952, et al. (författare) Arrested Development: The Molecular and Endocrine Basis of Flatfish Metamorphosis (ARRDE) 2006 Rapport (övrigt vetenskapligt/konstnärligt)
3.	Björnsson, Björn Thrandur, 1952, et al. (författare) Gastrointestinal functions and food intake regulation in salmonids: impact of dietary lipids 2002 Rapport (övrigt vetenskapligt/konstnärligt)abstract GUTINTEGRITY, Q5RT-2000-31656, Annual progress report for the first reporting period: 2001-01-01 – 2001-12-31
4.	Björnsson, Björn Thrandur, 1952, et al. (författare) Growth hormone endocrinology of salmonids: regulatory mechanisms and mode of action 2002 Ingår i: FISH PHYSIOLOGY AND BIOCHEMISTRY. - 0920-1742. ; 27:3-4, s. 227-242 Tidskriftsartikel (refereegranskat)abstract The focus of this review is on the regulatory mechanisms and the mode of action of GH in salmonids. To stimulate further research, it aims at highlighting areas where numerous important breakthroughs have recently been made, as well as where data are currently lacking. The regulation of GH secretion is under complex hypothalamic control, as well as under negative feedback control by GH and IGF-I. Further, the recently characterized ghrelin is a potent GH secretagogue, and may prove to be a link between feed intake and growth regulation. GH plasma profiles show indications of diurnal changes, but whether salmonids have true pulsatile GH secretion remains to be elucidated. The recent cloning and characterization of the salmon GH receptor (GHR) is a major research break-through which will give new insights into the mechanisms of GH action. It should also stimulate research into circulating GH-binding proteins (GHBPs), as they appear to be a soluble form of the GHR. The salmonid GHR sequences show evolutionary divergence from other fish species, but with a high degree of identity within the salmonid group. Radioreceptorassay studies have found GHR present in all tissues examined, which is in line with the highly pleiotropic action of GH. Data are currently scarce on the plasma dynamics of GH in salmonids, and further studies on GHR and GHBPs dynamics coupled to assessments of GH clearance rates and pathways are needed. The direct versus indirect nature of GH action remains to be clarified, but GH appears to act both locally at the target tissue level to stimulate the autocrine/paracrine action of IGF-I, as well as on the liver to increase plasma IGF-I levels. In addition, GH interacts with other hormones such as cortisol, thyroid hormones, insulin, and reproductive hormones, generating a wide range of physiological effects. GH may act both peripherally and directly at the level of the central nervous system to modify behavior, probably by altering the dopaminergic activity in the brain.
5.	Björnsson, Björn Thrandur, 1952, et al. (författare) Is salmon smoltification an example of vertebrate metamorphosis? Lessons learnt from work on flatfish larval development 2012 Ingår i: Aquaculture. - : Elsevier BV. - 0044-8486. ; 362-363, s. 264-272 Tidskriftsartikel (refereegranskat)abstract The terms metamorphosis and smoltification both describe developmental processes. However, the question on what specific criteria define these terms continues to engage scientists. At the same time, various views have been expressed on whether or not smoltification of anadromous salmonids should be regarded as an example of vertebrate metamorphosis. This short overview tries to summarize some of these discussions and starts by determining if smoltification meets any of the criteria used to define metamorphosis. In particular, it broadly compares the process of flatfish metamorphosis with that of salmonid smoltification from a morphological, endocrine, molecular and behavioral perspective. Tools and approaches developed and used in metamorphosis research which could be useful in continued work on smoltification are highlighted. (c) 2011 Elsevier B.V. All rights reserved.
6.	Björnsson, Björn Thrandur, 1952, et al. (författare) The Impact of Initial Energy Reserves on Growth Hormone Resistance and Plasma Growth Hormone-Binding Protein Levelsin Rainbow Trout Under Feeding and Fasting Conditions 2018 Ingår i: Frontiers in Endocrinology. - : Frontiers Media SA. - 1664-2392. ; 9 Tidskriftsartikel (refereegranskat)abstract The growth hormone (GH)-insulin-like growth factor I (IGF-I) system regulates important physiological functions in salmonid fish, including hydromineral balance, growth, and metabolism. While major research efforts have been directed toward this complex endocrine system, understanding of some key aspects is lacking. The aim was to provide new insights into GH resistance and growth hormone-binding proteins (GHBPs). Fish frequently respond to catabolic conditions with elevated GH and depressed IGF-I plasma levels, a condition of acquired GH resistance. The underlying mechanisms or the functional significance of GH resistance are, however, not well understood. Although data suggest that a significant proportion of plasma GH is bound to specific GHBPs, the regulation of plasma GHBP levels as well as their role in modulating the GH-IGF-I system in fish is virtually unknown. Two in vivo studies were conducted on rainbow trout. In experiment I, fish were fasted for 4 weeks and then refed and sampled over 72 h. In experiment II, two lines of fish with different muscle adiposity were sampled after 1, 2, and 4 weeks of fasting. In both studies, plasma GH, IGF-I, and GHBP levels were assessed as well as the hepatic gene expression of the growth hormone receptor 2a (ghr2a) isoform. While most rainbow trout acquired GH resistance within 4 weeks of fasting, fish selected for high muscle adiposity did not. This suggests that GH resistance does not set in while fat reserves as still available for energy metabolism, and that GH resistance is permissive for protein catabolism. Plasma GHBP levels varied between 5 and 25 ng ml(-1), with large fluctuations during both long-term (4 weeks) fasting and short-term (72 h) refeeding, indicating differentiated responses depending on prior energy status of the fish. The two opposing functions of GHBPs of prolonging the biological half-life of GH while decreasing GH availability to target tissues makes the data interpretation difficult, but nutritional regulatory mechanisms are suggested. The lack of correlation between hepatic ghr2a expression and plasma GHBP levels indicate that ghr2a assessment cannot be used as a proxy measure for GHBP levels, even if circulating GHBPs are derived from the GH receptor molecule.
7.	Breves, J. P., et al. (författare) Cortisol regulates insulin-like growth-factor binding protein (igfbp) gene expression in Atlantic salmon parr 2020 Ingår i: Molecular and Cellular Endocrinology. - : Elsevier BV. - 0303-7207 .- 1872-8057. ; 518 Tidskriftsartikel (refereegranskat)abstract © 2020 Elsevier B.V. The growth hormone (Gh)/insulin-like growth-factor (Igf)/Igf binding protein (Igfbp) system regulates growth and osmoregulation in salmonid fishes, but how this system interacts with other endocrine systems is largely unknown. Given the well-documented consequences of mounting a glucocorticoid stress response on growth, we hypothesized that cortisol inhibits anabolic processes by modulating the expression of hepatic igfbp mRNAs. Atlantic salmon (Salmo salar) parr were implanted intraperitoneally with cortisol implants (0, 10, and 40 μg g−1 body weight) and sampled after 3 or 14 days. Cortisol elicited a dose-dependent reduction in specific growth rate (SGR) after 14 days. While plasma Gh and Igf1 levels were unchanged, hepatic igf1 mRNA was diminished and hepatic igfbp1b1 and -1b2 were stimulated by the high cortisol dose. Plasma Igf1 was positively correlated with SGR at 14 days. Hepatic gh receptor (ghr), igfbp1a, -2a, -2b1, and -2b2 levels were not impacted by cortisol. Muscle igf2, but not igf1 or ghr, levels were stimulated at 3 days by the high cortisol dose. As both cortisol and the Gh/Igf axis promote seawater (SW) tolerance, and particular igfbps respond to SW exposure, we also assessed whether cortisol coordinates the expression of branchial igfbps and genes associated with ion transport. Cortisol stimulated branchial igfbp5b2 levels in parallel with Na+/K+-ATPase (NKA) activity and nka-α1b, Na+/K+/2Cl--cotransporter 1 (nkcc1), and cystic fibrosis transmembrane regulator 1 (cftr1) mRNA levels. The collective results indicate that cortisol modulates the growth of juvenile salmon via the regulation of hepatic igfbp1s whereas no clear links between cortisol and branchial igfbps previously shown to be salinity-responsive could be established.
8.	Breves, J. P., et al. (författare) Hepatic insulin-like growth factor binding protein (igfbp) responses to food restriction in Atlantic salmon smolts 2016 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 233, s. 79-87 Tidskriftsartikel (refereegranskat)abstract The growth hormone (Gh)/insulin-like growth-factor (Igf) system plays a central role in the regulation of growth in fishes. However, the roles of Igf binding proteins (Igfbps) in coordinating responses to food availability are unresolved, especially in anadromous fishes preparing for seaward migration. We assayed plasma Gh, Igf1, thyroid hormones and cortisol along with igfbp mRNA levels in fasted and fed Atlantic salmon (Salmo solar). Fish were fasted for 3 or 10 days near the peak of smoltification (late April to early May). Fasting reduced plasma glucose by 3 days and condition factor by 10 days. Plasma Gh, cortisol, and thyroxine (T-4) were not altered in response to fasting, whereas Igf1 and 3,5,3'-triiodo-L-thyronine (T-3) were slightly higher and lower than controls, respectively. Hepatic igfbp1b1, -1b2, -2a,-2b1 and -2b2 mRNA levels were not responsive to fasting, but there were marked increases in igfbp1a1 following 3 and 10 days of fasting. Fasting did not alter hepatic igf1 or igf2; however, muscle igf1 was diminished by 10 days of fasting. There were no signs that fasting compromised branchial ionoregulatory functions, as indicated by unchanged Na+/K+-ATPase activity and ion pump/transporter mRNA levels. We conclude that dynamic hepatic igfbp1a1 and muscle igf1 expression participate in the modulation of Gh/Igf signaling in smolts undergoing catabolism.
9.	Breves, J. P., et al. (författare) In vivo effects of 17α-ethinylestradiol, 17β-estradiol and 4-nonylphenol on insulin-like growth-factor binding proteins (igfbps) in Atlantic salmon 2018 Ingår i: Aquatic Toxicology. - : Elsevier BV. - 0166-445X. ; 203, s. 28-39 Tidskriftsartikel (refereegranskat)abstract Feminizing endocrine disrupting compounds (EDCs) affect the growth and development of teleost fishes. The major regulator of growth performance, the growth hormone (Gh)/insulin-like growth-factor (Igf) system, is sensitive to estrogenic compounds and mediates certain physiological and potentially behavioral consequences of EDC exposure. Igf binding proteins (Igfbps) are key modulators of Igf activity, but their alteration by EDCs has not been examined. We investigated two life-stages (fry and smolts) of Atlantic salmon (Salmo salar), and characterized how the Gh/Igf/Igfbp system responded to waterborne 17α-ethinylestradiol (EE2), 17β-estradiol (E2) and 4-nonylphenol (NP). Fry exposed to EE2 and NP for 21 days had increased hepatic vitellogenin (vtg) mRNA levels while hepatic estrogen receptor α (erα), gh receptor (ghr), igf1 and igf2 mRNA levels were decreased. NP-exposed fry had reduced body mass and total length compared to controls. EE2 and NP reduced hepatic igfbp1b1, -2a, -2b1, -4, -5b2 and -6b1, and stimulated igfbp5a. In smolts, hepatic vtg mRNA levels were induced following 4-day exposures to all three EDCs, while erα only responded to EE2 and E2. EDC exposures did not affect body mass or fork length; however, EE2 diminished plasma Gh and Igf1 levels in parallel with reductions in hepatic ghr and igf1. In smolts, EE2 and E2 diminished hepatic igfbp1b1, -4 and -6b1, and stimulated igfbp5a. There were no signs of compromised ionoregulation in smolts, as indicated by unchanged branchial ion pump/transporter mRNA levels. We conclude that hepatic igfbps respond (directly and/or indirectly) to environmental estrogens during two key life-stages of Atlantic salmon, and thus may modulate the growth and development of exposed individuals.
10.	Breves, J. P., et al. (författare) Variation in branchial expression among insulin-like growth-factor binding proteins (igfbps) during Atlantic salmon smoltification and seawater exposure 2017 Ingår i: BMC Physiology. - : Springer Science and Business Media LLC. - 1472-6793. ; 17:1 Tidskriftsartikel (refereegranskat)abstract Background: In preparation for migration from freshwater to marine habitats, Atlantic salmon (Salmo salar L.) undergo smoltification, a transformation that includes the acquisition of hyposmoregulatory capacity. The growth hormone (Gh)/insulin-like growth-factor (Igf) axis promotes the development of branchial ionoregulatory functions that underlie ion secretion. Igfs interact with a suite of Igf binding proteins (Igfbps) that modulate hormone activity. In Atlantic salmon smolts, igfbp4,-5a,-5b1,-5b2,-6b1 and-6b2 transcripts are highly expressed in gill. We measured mRNA levels of branchial and hepatic igfbps during smoltification (March, April, and May), desmoltification (July) and following seawater (SW) exposure in March and May. We also characterized parallel changes in a broad suite of osmoregulatory (branchial Na+/K+-ATPase (Nka) activity, Na + /K + /2Cl - cotransporter 1 (nkcc1) and cystic fibrosis transmembrane regulator 1 (cftr1) transcription) and endocrine (plasma Gh and Igf1) parameters. Results: Indicative of smoltification, we observed increased branchial Nka activity, nkcc1 and cftr1 transcription in May. Branchial igfbp6b1 and -6b2 expression increased coincidentally with smoltification. Following a SW challenge in March, igfbp6b1 showed increased expression while igfbp6b2 exhibited diminished expression. igfbp5a,-5b1 and-5b2 mRNA levels did not change during smolting, but each had lower levels following a SW exposure in March. Conclusions: Salmonids express an especially large suite of igfbps. Our data suggest that dynamic expression of particular igfbps accompanies smoltification and SW challenges; thus, transcriptional control of igfbps may provide a mechanism for the local modulation of Igf activity in salmon gill. © 2017 The Author(s).
11.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Atlantic halibut growth hormone: structure and plasma levels of sexually mature males and females during photoperiod-regulated annual cycles 2002 Ingår i: General and Comparative Endocrinology. - 0016-6480 .- 1095-6840. ; 127:1, s. 94-104 Tidskriftsartikel (refereegranskat)abstract The main objectives of this study were to obtain the amino acid sequence of Atlantic halibut (Hippoglossus hippoglossus) growth hormone (hhGH) and compare it with other teleost species, to establish a radioimmunoassay to assess plasma hhGH levels and thus to gain information about possible biological functions and regulation by photoperiod. The hhGH gene was cloned and its amino acid sequence deduced from the cDNA. The mature hhGH protein consists of 186 amino acids. Comparison with other flatfish species as well as a species from a different order, the pufferfish, reveals that the sequence similarities of the mature hhGH with that of the barfin flounder, the Japanese flounder, the sole and the pufferfish are 99.5, 81.7, 74.2, and 65.2%, respectively. The sequence similarities appear to correctly reflect the gross phylogenetic relationships among these telcost species. A specific GH-RIA was developed for measurements of Atlantic halibut GH levels. Assessment of plasma GH levels in adult halibut revealed large gender differences, with GH levels frequently being an order of magnitude higher in males than females. The mean (+/-SEM) plasma GH for males kept on normal annual photoperiod were 25.2 +/- 6.11 ng ml(-1) and for females were 5.14 +/- 1.94 ng ml(-1). It appears likely that plasma growth hormone levels in Atlantic halibut can be inversely correlated to growth and metabolism. Shifting of the annual photoperiod cycles demonstrated that photoperiod in not a regulator of plasma GH levels in the Atlantic halibut. but further research is needed to assess whether GH plays a role in the reproduction of this marine teleost species. (C) 2002 Elsevier Science (USA). All rights reserved.
12.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Distribution of ghrelin-producing cells and Na+,K+-ATPase in tissues of Atlantic halibut (Hippoglossus hippoglossus) during early development. 2010 Ingår i: European Aquaculture Scociety, Porto, Potugal, October 5-8, 2010. ; Conference Program Konferensbidrag (övrigt vetenskapligt/konstnärligt)
13.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Effects of rearing stress on Atlantic salmon (Salmo salar L.) antibody response to a non-pathogenic antigen. 2000 Ingår i: Aquaculture Research. - : Hindawi Limited. - 1355-557X .- 1365-2109. ; 31:12, s. 932-930 Tidskriftsartikel (refereegranskat)abstract This study was carried out to test the effects of acute stress on the primary and the secondary antibody responses in Atlantic salmon (Salmo salar L.). Fish received a primary immunization with sheep red blood cells (SRBC) and received a second immunization 11 weeks later. At priming, a group of fish was stressed by water level reduction for 30 min (moderate stress), and another group was stressed severely by applying the stressor twice, with an interval of 6 h, 3 days in a row (severe stress). At the time of the booster, another two groups of previously unstressed, primed fish were treated likewise. Unstressed fish mounted a primary antibody response to SRBC and a secondary response, which was improved compared with the primary response., The primary antibody response against SRBC was impaired by severe, but not moderate, acute stress at priming, whereas the secondary antibody response was unaffected. Both moderate and severe stress at the time of the booster was immunosuppressive in fish that had been primed under non-stress conditions. It is concluded that cortisol release as a result of stress exposure is most immunosuppressive at the time of the second immunization.
14.	Einarsdottir, Ingibjörg, 1951, et al. (författare) GH-receptor derived plasma GH binding protein in rainbow trout : Effects of fasting and a single-meal refeeding 2012 Ingår i: 7th International Symposium of Fish Endocrinology, Buenos Aires, Argentina, 3-6 September 2012. ; Program and Abstract Book Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract Plasma growth hormone binding proteins (GHBPs) transport circulating growth hormone (GH), function as a GH pool and prevent GH from degradation. In mammals, high-affinity GHBPs are identical to the extracellular domain of the growth hormone receptor (GHR). GHBPs have been demonstrated in fish, but their molecular identity has not been confirmed. To examine a possible GHR origin of plasma GHBPs in rainbow trout, a GHBP radioimmunoassay was established, using antibodies raised against a synthesized peptide sequence of the salmonid GHR extracellular domain, and recombinant proteins for iodination and standards. In order to examine effects of nutritional state on GHBP levels, rainbow trout were fasted for four weeks while a control group was fed to satiation. Then, all fish received a single meal and were sampled 2, 4, 8, 24 and 72 h postprandially. During the 4-week fasting, GHBP levels did not change, while plasma GH levels increased, as has been shown in earlier studies. Following the meal, a transient decrease in both GH and GHBP was seen in the fasted fish, lasting 2 to 4 h. This is the first time plasma GHBP levels have been measured in fish. The data indicate that GHBP and GH levels may be regulated independently or in concert. Supported by EU (Project LIFECYCLE FP7-222719).
15.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Growth hormone profiles and development of somatotrophs in Atlantic halibut larvae 2000 Ingår i: International Congress on the Biology of Fish, July 23-27, 2000. Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract Introduction. The Atlantic halibut is the largest flatfish species, and as other flatfish, has a complicated larval development. The pelagic larvae hatch after about two weeks and feeding starts six weeks later. After three to four months, they start to undergo metamorphosis. Following major changes in body shape, including the migration of the left to the right side, the larvae settle as bottom dwelling. In Atlantic halibut aquaculture, the larval rearing is a critical rearing stage, with high incidence of mortality and abnormal development. Growth hormone in teleost fishes is known to participate in the regulation of several important physiological processes including metabolism, growth, appetite, and osmoregulation. It is therefore likely that the hormone may be important for larval growth and development. Therefore, this study was carried out in order to elucidate the growth hormone (GH) endocrinology of halibut larvae, by measuring GH tissue content as well as the histology of the pituitary somatotrophs. Materials and Methods. The study was carried out with unfertilized eggs and larvae from hatching through metamorphosis (from 22 to 859 day-degrees (D°)), collected at the halibut hatchery of Fiskey Ltd, Northern Iceland, over two consecutive years. In order to study GH profiles during early development, a homologous radioimmunoassay was established. GH was isolated from adult halibut pituitaries collected at Fiskey Ltd, using methods modified from Johnson et al (1997). In the radioimmunoassay, this GH was used for standards and iodination, together with specific antibodies raised in rabbits, courtesy of Dr. P. Swanson. For the immunohistochemistry of GH-producing somatotrophs in the pituitaries, anti-halibut antibodies were raised in rabbits and validated. Results and Conclusions. Tissue GH analysis revealed that GH is detectable in unfertilized eggs. In developing larvae, tissue GH content per body weight increased during development from hatching to metamorphosis. The earliest stage at which GH was localized in the somatotrophs by immunohistochemistry, was at the age of 187 D°. The present study demonstrates that there appears to be a maternal source of growth hormone in halibut eggs, similar to what has been demonstrated for thyroid hormones in eggs of different teleost species (Kobuke et al 1987). The study further demonstrates that endogenous production of growth hormone is initiated early in larval development, during the yolk-sack stage, prior to first feeding. The study establishes that GH can play a regulatory role during early development of the Atlantic halibut
16.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Isolation of Atlantic halibut pituitary hormones by continuous-elution electrophoresis followed by fingerprint identification, and assessment of growth hormone content during larval development 2007 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 150:2, s. 355-363 Tidskriftsartikel (refereegranskat)abstract Continuous-elution electrophoresis (CEE) has been applied to separate putative hormones from adult Atlantic halibut pituitaries. Soluble proteins were separated by size and charge on Model 491 Prep Cell (Bio-Rad), where the homogenate runs through a cylindrical gel, and protein fractions are collected as they elute from the matrix. Protein fractions were assessed by SDS–PAGE and found to contain purified proteins of molecular size from 10 to 33 kDa. Fractions containing proteins with molecular weights of approximately 21, 24, 28 and 32 kDa, were identified as putative growth hormone (GH), prolactin, somatolactin and gonadotropins, respectively. These were analyzed further by mass spectrometry and identified with peptide mass protein fingerprinting. The CEE technique was used successfully for purification of halibut GH with a 5% yield, and appears generally well suited to purify species-specific proteins often needed for research in comparative endocrinology, including immunoassay work. Thus, the GH obtained was subsequently used as standards and iodination label in a homologous radioimmunoassay, applied to analyze GH content through larval development in normally and abnormally metamorphosing larvae. As GH is mainly found in the pituitary, GH contents were analyzed in tissue extracts from the heads only. The pituitary GH content increases proportionally to increased larval weight from first feeding to metamorphic climax. No difference in relative GH content was found between normal and abnormal larvae and it still remains to be established if GH has a direct role in metamorphosis.
17.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Occurrence of ghrelin-producing cells, the ghrelin receptor and Na+,K+-ATPase in tissues of Atlantic halibut 2011 Ingår i: Cell and Tissue Resarch. - 0302-766X. ; 344:3, s. 481-498 Tidskriftsartikel (refereegranskat)abstract Ghrelin is a pituitary growth hormone (GH)-secretagogue that also has metabolic, reproductive, proliferative, immunological and brain functions in mammals. Far less is known about its role in fish. We have therefore performed an immunohistochemical determination of its tissue distribution in the developing Atlantic halibut (Hippoglossus hippoglossus) to gain insights into its potential function. Ghrelin immunoreactivity was detected in first-feeding halibut larvae in the skin, urinary bladder, gastrointestinal (GI) tract and olfactory lobe of the brain. In subsequent stages up to metamorphosis, ghrelin immunoreactivity declined in the skin and became evident in the gills. When the stomach developed, ghrelin immunoreactivity declined throughout the GI tract with the exception of the stomach, which exhibited an intense signal. Immunoreactive ghrelin cells were also present in the olfactory lobe, nerve and epithelium and in occasional cells of the buccal cavity and oesophagus. Ghrelin immunoreactivity had an overlapping distribution with that for Na(+),K(+)-ATPase, colocalisation also being observed in some ionocytes of the gill. The co-expression of ghrelin and the GH-secretagogue receptor in the same tissue indicates that ghrelin can exert both endocrine and paracrine actions in the developing halibut. The presence of immunoreactive ghrelin in several osmoregulatory tissues, the GI tract and sensory tissue provides strong evidence that ghrelin has multiple functions during development and also suggests targets for future investigations.
18.	Einarsdottir, Ingibjörg, 1951, et al. (författare) PITUITARY HORMONE PURIFICATION BY CONTINUOUS-ELUTION ELECTROPHORESIS 2006 Ingår i: 7th International Congress on the Biology of Fishes, Book of Abstracts. Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract Widely used methods for peptide hormone purification, such as gel filtration chromatography combined with rpHPLC, are time-consuming and often give low yields and instable proteins. The Model 491 Prep Cell from Bio-Rad provides a continuous-elution electrophoresis technique to separate proteins. The protein mixture runs through a cylindrical gel, proteins are separated by size and charge, as they elute from the matrix and are collected using a fraction collector. Sub-samples are assessed by SDS-PAGE and protein identity is verified by Western blot analysis. We applied this technique to separate putative hormones from adult Atlantic halibut pituitaries. Soluble proteins from homogenized pituitaries were denatured and subjected to continuous-elution electrophoresis. The fractions sampled contained purified proteins with molecular sizes ranging from 10 to 33 kDa. Fractions containing proteins with molecular weights of approximately 21, 24, 28 and 32 kDa were identified as putative growth hormone, prolactin, somatolactin and gonadotropins. These were analyzed further by mass spectrometry (MS) and identified with peptide mass protein fingerprinting. Good yield of highly purified GH was obtained, identified, and found to be biologically stable. Subsequently, it was used for radioactive labeling and as standards in a homologous RIA. Using this assay, the GH was measured in halibut larvae from start-feeding through metamorphosis, showing a stable size-related GH content of the head region. Funded by the EU.
19.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Plasma growth hormone-binding protein levels in Atlantic salmon Salmo salar during smoltification and seawater transfer 2014 Ingår i: Journal of Fish Biology. - : Wiley. - 0022-1112. ; 85:4, s. 1279-1296 Tidskriftsartikel (refereegranskat)abstract Specific growth hormone (GH)-binding protein (Ghbp) was purified from Atlantic salmon Salmo salar and rainbow trout Oncorhynchus mykiss plasma with immunoprecipitation and characterized in cross-linking studies using autoradiography and western blots. The size of the Ghbp was estimated to be c. 53 kDa. A radioimmunoassay was established to measure Ghbp in salmonids, using antibodies specific against the extracellular segment of the S. salar growth hormone receptor 1 (grh1; GenBank AY462105). Plasma Ghbp levels were measured in S. salar smolts in fresh water and after transfer to seawater (SW; experiments 1 and 2), and in post-smolts kept at different salinities (0, 12, 22 and 34) for 3 months (experiment 3). A transient increase in plasma Ghbp, which lasted for 1 month or less, was noted in smolts after transfer to SW. Concomitantly, plasma GH and gill Na+-K+-ATPase activity increased during smoltification (in experiment 2). No difference in plasma Ghbp was evident between post-smolts kept at different salinities, although the fish kept at salinity 34 had higher plasma GH than the group kept at salinity 22 and higher hepatic ghr1 expression than post-smolts kept at salinity 12. This suggests that plasma Ghbp regulation may respond to salinity changes in the short term. The lack of correlation between Ghbp, plasma GH and hepatic ghr1 expression in the long-term post-smolt experiment indicates that Ghbp levels may be regulated independently of other components of the endocrine GH system in salmonids.
20.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Plasma growth hormone levels during sexual maturation of Atlantic halibut: influence of annual photoperiod cycle 2001 Ingår i: 14th International Congress of Comparative Endocrinology (ICCE), Sorrento, Italy, May 26-30, 2001. ; Book of Abstracts Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract Three groups of sexually mature male and female Atlantic halibut (Hippoglossus hippoglossus), kept in indoor tanks under computer-simulated photoperiod regimes, were serially sampled every month over three annual maturational cycles (Björnsson et al 1998). During the ini¬tial phase of the study, the annual photoperiod cycle was advanced by four months for one group and delayed by four months for another, while the third control group was kept on simulated natural annual pho¬toperiod cycle. Plasma growth hormone levels (GH) were assessed using a newly estab¬lished halibut GH-RIA (Einarsdottir et al 2000, in prep). In the control group, males had much higher GH levels than females and showed strong annual variations, with a rapid rise in GH levels to ca 50 ng/ml towards the beginning of female spawning. At that time, the GH levels of the females were only around 0.5-2 ng/ml, and rarely exceeding 10 ng/ml at other times of the year. The advancement and delay of the annual photoperiod cycle by four months resulted in gradual corresponding shifts in spawning time, reaching four months after three annual cycles (Björnsson et al 1998). As in controls, the over-all difference in GH levels between males and females was seen in the advanced and the delayed groups. However, the shift in annual photoperiod cycle caused a certain disruption of the annual GH cycle in males, but the pattern re-emerged towards the end of the study, shifted in line with the annual photoperiod shift. As the annual spawning period and GH changes are shifted in parallel, the study does not establish a causal relationship between GH levels and sexual maturation in the halibut. However, as in other teleost species such as the salmonids, seasonal changes in plasma GH levels appear to be under photoperiod control. Financial support by EU-FAIR (CT96-1422) and the Nordic Industrial Fund Björnsson et al 1998. Aquaculture 166: 117-140
21.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Purification of pituitary hormones from the Atlantic halibut (Hippoglossus hippoglossus) by continuous-elution electrophoresis and identification of isolated proteins by fingerprinting 2004 Ingår i: 5th International Symposium on Fish Endocrinology, Book of Abstracts. Konferensbidrag (övrigt vetenskapligt/konstnärligt)
22.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Temperature influence on Arctic charr (Salvelinus alpinus L.) antibody response to a cellular antigen. 2000 Ingår i: Polar Biology. - : Springer Science and Business Media LLC. - 0722-4060 .- 1432-2056. ; 23:4, s. 231-235 Tidskriftsartikel (refereegranskat)abstract In order to elucidate the immune responses of Arctic charr in relation to temperature, groups were acclimated to a moderate (9 degrees C) and a cold temperature regime (4 degrees C), as well as subjected to a temperature decrease (from 9 to 4 degrees C) immediately prior to an immunization with sheep red blood cells. The charr kept at 9 degrees C responded with increased primary and secondary antibody titres, as seen by direct haemagglutination, while fish at 4 degrees C, as well as the fish subjected to a temperature reduction, displayed lower and lowest antibody titres, respectively, and only after a second immunization, It is concluded that Arctic charr can respond to a cellular antigen with a humoral immune response typical for other teleosts, but that the immune response is delayed and diminished at low temperatures. This temperature-induced immune suppression is intensified if the fish have not been acclimated to cold water prior to immunization.
23.	Einarsdottir, Ingibjörg, 1951, et al. (författare) The growth hormone – IGF-I system is involved in cranial tissue remodeling in metamorphosing Atlantic halibut larvae 2008 Ingår i: Sixth International Symposium on Fish Endocrinology (6ISFE) Program and Abstract Book. Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract The Atlantic halibut is a large, cold-water flatfish which inhabits the North Atlantic. The pelagic and symmetric larvae undergo metamorphosis as an adaptation to better suit them for a benthic life. Metamorphosis involves many dramatic biological changes including a major reconstruction of the cranium during which the left eye migrates to the right side of the head. The structural driving force of eye migration is assumed to be the growth and migration of the ethmoid plate and the migration of the frontal processes towards the future ocular side, and fibroblast proliferation ventrally to the migrating eye. Metamorphosis is under endocrine control and thyroid hormones are known to be the main regulatory hormones. Data on the role of other growth regulators, including the growth hormone (GH) - insulin-like growth factor I (IGF-I) system, in halibut metamorphosis are sparse. GH content in the head region increases steadily from first feeding throughout the larval phase until post-metamorphosis, while IGF-I content in the body increases until the onset of metamorphosis and decline thereafter. In situ hybridization and immunohistochemistry reveal that mRNA and proteins of the type I GH receptor (GHR), and mRNA of the type II GHR and the IGF-I receptor (IGF-IR) are expressed in fibroblasts in soft tissue between the eyes, frontal bone osteocytes and dorsal cranial chondrocytes during the metamorphic process. In fibroblasts and chondrocytes, the expression appears most intense at the start of metamorphosis (stage 8) and decreases during subsequent developmental stages. IGF-IR expression decreases in frontal bone osteocytes after the completion of metamorphosis (stage 10). The decrease in tissue IGF-I content during metamorphosis, and the localization and change in abundance of GHRs and IGF-IR in head structures undergoing reconstruction, indicate the involvement of the GH-IGF-I system in Atlantic halibut metamorphosis. Acknowledgement: Work financed by the Commission of the European Communities within the project “Arrested development: the molecular and endocrine basis of flatfish metamorphosis” (Q5RS-2002-01192).
24.	Einarsdottir, Ingibjörg, 1951, et al. (författare) The relationship between growth hormone, growth hormone receptor and growth hormone binding proteins in smolting Atlantic salmon 2013 Ingår i: Smolt-conference, Reykjavik, Island, 12-16 August 2013. Konferensbidrag (övrigt vetenskapligt/konstnärligt)
25.	Einarsdottir, Ingibjörg, 1951, et al. (författare) Thyroid and pituitary gland development from hatching through metamorphosis of a teleost flatfish, the Atlantic halibut 2006 Ingår i: Anatomy and Embryology. - : Springer Science and Business Media LLC. - 0340-2061 .- 1432-0568. ; 211:1, s. 47-60 Tidskriftsartikel (refereegranskat)abstract Fish larval development, not least the spectacular process of flatfish metamorphosis, appears to be under complex endocrine control, many aspects of which are still not fully elucidated. In order to obtain data on the functional development of two major endocrine glands, the pituitary and the thyroid, during flatfish metamorphosis, histology, immunohistochemistry and in situ hybridization techniques were applied on larvae of the Atlantic halibut (Hippoglossus hippoglossus), a large, marine flatfish species, from hatching through metamorphosis. The material was obtained from a commercial hatchery. Larval age is defined as day-degrees (D degrees=accumulated daily temperature from hatching). Sporadic thyroid follicles are first detected in larvae at 142 D degrees (27 days post-hatch), prior to the completion of yolk sack absorption. Both the number and activity of the follicles increase markedly after yolk sack absorption and continue to do so during subsequent development. The larval triiodothyronine (T-3) and thyroxine (T-4) content increases, subsequent to yolk absorption, and coincides with the proliferation of thyroid follicles. A second increase of both T-3 and T-4 occurs around the start of metamorphosis and the T-3 content further increases at the metamorphic climax. Overall, the T-3 content is lower than T-4. The pituitary gland can first be distinguished as a separate organ at the yolk sack stage. During subsequent development, the gland becomes more elongated and differentiates into neurohypophysis (NH), pars distalis (PD) and pars intermedia (PI). The first sporadic endocrine pituitary cells are observed at the yolk sack stage, somatotrophs (growth hormone producing cells) and somatolactotrophs (somatolactin producing cells) are first observed at 121 D degrees (23 days post-hatch), and lactotrophs (prolactin producing cells) at 134 D degrees (25 days post-hatch). Scarce thyrotrophs are evident after detection of the first thyroid follicles (142 D degrees), but coincident with a phase in which follicle number and activity increase (260 D degrees). The somatotrophs are clustered in the medium ventral region of the PD, lactotrophs in the anterior part of the PD and somatolactotrophs are scattered in the mid and posterior region of the pituitary. At around 600 D degrees, coinciding with the start of metamorphosis, somatolactotrophs are restricted to the interdigitating tissue of the NH. During larval development, the pituitary endocrine cells become more numerous. The present data on thyroid development support the notion that thyroid hormones may play a significant role in Atlantic halibut metamorphosis. The time of appearance and the subsequent proliferation of pituitary somatotrophs, lactotrophs, somatolactotrophs and thyrotrophs indicate at which stages of larval development and metamorphosis these endocrine cells may start to play active regulatory roles.
26.	Fuentes, EN, et al. (författare) IGF-I/PI3K/Akt and IGF-I/MAPK/ERK pathways in vivo in skeletal muscle are regulated by nutrition and contribute to somatic growth in the fine flounder 2011 Ingår i: American Journal of Physiology - Regulatory, Integrative. - 0363-6119. ; 300:6, s. 1532-1542 Tidskriftsartikel (refereegranskat)abstract The insulin-like growth factor-I (IGF-I) is a key regulator of skeletal muscle growth in vertebrates, promoting mitogenic and anabolic effects through the activation of the MAPK/ERK and the PI3K/Akt signaling pathways. Nutrition also affects skeletal muscle growth, activating intracellular pathways and inducing protein synthesis and accretion. Thus, both hormonal and nutritional signaling regulate muscle mass. In this context, plasma IGF-I levels and the activation of both pathways in response to food were evaluated in the fine flounder using fasting and refeeding trials. The present study describes for the first time in a nonmammalian species that the MAPK/ERK and PI3K/Akt are activated by exogenous circulating IGF-I, as well as showing that the MAPK/ERK pathway activation is modulated by the nutritional status. Also, these results show that there is a time-dependent regulation of IGF-I plasma levels and its signaling pathways in muscle. Together, these results suggest that the nutritionally managed IGF-I could be regulating the activation of the MAPK/ERK and the PI3K/Akt signaling pathways differentially according to the nutritional status, triggering different effects in growth parameters and therefore contributing to somatic growth in fish. This study contributes to the understanding of the nutrient regulation of IGF-I and its signaling pathways in skeletal muscle growth in nonmammalian species, therefore providing insight concerning the events controlling somatic growth in vertebrates.
27.	Fuentes, E. N., et al. (författare) Inherent Growth Hormone Resistance in the Skeletal Muscle of the Fine Flounder Is Modulated by Nutritional Status and Is Characterized by High Contents of Truncated GHR, Impairment in the JAK2/STAT5 Signaling Pathway, and Low IGF-I Expression 2012 Ingår i: Endocrinology. - : The Endocrine Society. - 0013-7227 .- 1945-7170. ; 153:1, s. 283-294 Tidskriftsartikel (refereegranskat)abstract A detailed understanding of how the GH and IGF-I regulate muscle growth, especially in early vertebrates, is still lacking. The fine flounder is a flatfish species exhibiting remarkably slow growth, representing an intriguing model for elucidating growth regulatory mechanisms. Key components of the GH system were examined in groups of fish during periods of feeding, fasting, and refeeding. Under feeding conditions, there is an inherent systemic and local (muscle) GH resistance, characterized by higher levels of plasma GH than of IGF-I, skeletal muscle with a greater content of the truncated GH receptor (GHRt) than of full-length GHR (GHRfl), an impaired activation of the Janus kinase 2 (JAK2)-signal transducers and activators of transcription 5 (STAT5) signaling pathway, and low IGF-I expression. Fasting leads to further elevation of plasma GH levels concomitant with suppressed IGF-I levels. The ratio of GHRfl to GHRt in muscle decreases during fasting, causing an inactivation of the JAK2/STAT5 signaling pathway and suppressed IGF-I expression, further impairing growth. When fish are returned to nutritionally favorable conditions, plasma GH levels decrease, and the ratio of GHRfl to GHRt in muscle increases, triggering JAK2/STAT5 reactivation and local IGF-I expression, concomitant with increased growth. The study suggests that systemic IGF-I is supporting basal slow growth in this species, without ruling out that local IGF-I is participating in muscle growth. These results reveal for the first time a unique model of inherent GH resistance in the skeletal muscle of a nonmammalian species and contribute to novel insights of the endocrine and molecular basis of growth regulation in earlier vertebrates. (Endocrinology 153:283-294, 2012)
28.	Fuentes, E. N., et al. (författare) Nutritional status modulates plasma leptin, AMPK and TOR activation, and mitochondrial biogenesis: Implications for cell metabolism and growth in skeletal muscle of the fine flounder 2013 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 186, s. 172-180 Tidskriftsartikel (refereegranskat)abstract Insight of how growth and metabolism in skeletal muscle are related is still lacking in early vertebrates. In this context, molecules involved in these processes, such as leptin, AMP-activated protein kinase (AMPK), target of rapamicyn (TOR), peroxisome proliferator-activated receptor γ coactivator 1α (PGC-1α, and oxidative phosphorylation complexes (OXPHOS), were assessed in the skeletal muscle of a fish species. Periods of fasting followed by a period of refeeding were implemented, using the fine flounder as a model (Paralichthys adspersus). This species exhibits remarkably slow growth and food intake, which is linked to an inherent growth hormone (GH) resistance and high circulating levels of leptin. Leptin increased during fasting concomitantly with AMPK activation, which was inversely correlated with TOR activation. On the other hand, AMPK was directly correlated with an increase in PGC-1α and OXPHOS complexes contents. Dramatic changes in the activation and content of these molecules were observed during short-term refeeding. Leptin, AMPK activation, and PGC-1α/OXPHOS complexes contents decreased radically; whereas, TOR activation increased significantly. During long-term refeeding these molecules returned to basal levels. These results suggest that there is a relation among these components; thus, during fasting periods ATP-consuming biosynthetic pathways are repressed and alternative sources of ATP/energy are promoted, a phenomenon that is reversed during anabolic periods. These results provide novel insight on the control of metabolism and growth in the skeletal muscle of a non-mammalian species, suggesting that both processes in fish muscle are closely related and coordinated by a subset of common molecules.
29.	Fuentes, E. N., et al. (författare) Plasma leptin and growth hormone levels in the fine flounder (Paralichthys adspersus) increase gradually during fasting and decline rapidly after refeeding 2012 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 177:1, s. 120-127 Tidskriftsartikel (refereegranskat)abstract In fish, recent studies have indicated an anorexigenic role of leptin and thus its possible involvement in regulation of energy balance and growth. In the present study, the effects of fasting and refeeding periods on plasma leptin levels were studied in the fine flounder, a flatfish with remarkably slow growth. To further assess the endocrine status of the fish during periods of catabolism and anabolism, plasma growth hormone (GH) levels were also analyzed. Under normal feeding condition, plasma leptin and CH levels remained stable and relatively high in comparison with other teleost species. For the three separate groups of fish, fasted for 2, 3, and 4 weeks, respectively, plasma leptin levels increase gradually, becoming significantly elevated after 3 weeks, and reaching highest levels after 4-week fasting. Plasma GH levels were significantly elevated after 2-week fasting. At the onset of refeeding, following a single meal, leptin levels decline rapidly to lower than initial levels within 2 h, irrespective of the length of fasting. Plasma GH also decline, the decrease being significant after 4, 24 and 2 h for the 2, 3 and 4-week fasted groups, respectively. This study shows that plasma leptin levels in the fine flounder are strongly linked to nutritional status and suggests that leptin secretion is regulated by fast-acting mechanisms. Elevated leptin levels in fasted fish may contribute to a passive survival strategy of species which experience natural food shortage periods by lowering appetite and limiting physical foraging activity. (C) 2012 Elsevier Inc. All rights reserved.
30.	Fuentes, E. N., et al. (författare) The TORC1/P70S6K and TORC1/4EBP1 signaling pathways have a stronger contribution on skeletal muscle growth than MAPK/ERK in an early vertebrate: Differential involvement of the IGF system and atrogenes 2015 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 210, s. 96-106 Tidskriftsartikel (refereegranskat)abstract Knowledge about the underlying mechanisms, particularly the signaling pathways that account for muscle growth in vivo in early vertebrates is still scarce. Fish (Paralichthys adspersus) were fasted for 3 weeks to induce a catabolic period of strong muscle atrophy. Subsequently, fish were refed for 2 weeks to induce compensatory muscle hypertrophy. During refeeding, the fish were treated daily with either rapamycin (TORC blocker), P098059 (MEK blocker), or PBS (V; vehicle), or were untreated (C; control). Rapamycin and P098059 differentially impaired muscle cellularity in vivo, growth performance, and the expression of growth-related genes, and the inhibition of TORC1 had a greater impact on fish muscle growth than the inhibition of MAPK. Blocking TORC1 inhibited the phosphorylation of P70S6K and 4EBP1, two downstream components activated by TORC1, thus affecting protein contents in muscle. Concomitantly, the gene expression in muscle of igf-1, 2 and igfbp-4, 5 was down-regulated while the expression of atrogin-1, murf-1, and igfbp-2, 3 was up-regulated. Muscle hypertrophy was abolished and muscle atrophy was promoted, which finally affected body weight. TORC2 complex was not affected by rapamycin. On the other hand, the PD98059 treatment triggered ERK inactivation, a downstream component activated by MEK. mRNA contents of igf-1 in muscle were down-regulated, and muscle hypertrophy was partially impaired. The present study provides the first direct data on the in vivo contribution of TORC1/P70S6K, TORC1/4EBP1, and MAPK/ERK signaling pathways in the skeletal muscle of an earlier vertebrate, and highlights the transcendental role of TORC1 in growth from the cellular to organism level. (C) 2014 Elsevier Inc. All rights reserved.
31.	Gong, Ningping, 1979, et al. (författare) Alternative Splice Variants of the Rainbow Trout Leptin Receptor Encode Multiple Circulating Leptin-Binding Proteins 2013 Ingår i: Endocrinology. - : The Endocrine Society. - 0013-7227 .- 1945-7170. ; 154:7, s. 2331-2340 Tidskriftsartikel (refereegranskat)abstract In mammals, leptin (Lep) binding proteins (LepBPs) derived from Lep receptor (LepR) gene or protein bind most of the circulating Lep, but to date, information on LepBPs in nonmammalian vertebrate classes is lacking. This study details the characterization of multiple LepBPs in rainbow trout (Oncorhynchus mykiss), an early poikilothermic vertebrate, and presents the complete coding sequences for 3 of them. Size-exclusion chromatography and cross-linking assay identified plasma proteins bound to Lep ranging from 70 to 100 kDa. LepBPs were isolated from plasma by affinity chromatography, and their binding specificity was assessed by a competitive binding assay. A RIA for LepBPs indicates that plasma LepBP levels decline after fasting for 3 weeks. Immunoblotting of LepBPs using antibodies against different LepR epitopes shows that the LepBPs are indeed LepR isoforms. The alternatively spliced LepR transcripts (LepR(S1-3)) that include only the extracellular segment transcribe the 90-kDa LepBP1, the 80-kDa LepBP2, and the 70-kDa LepBP3, respectively. LepR(S1) generally has lower expression than the long-form LepR in most tissues. LepR(S2) is primarily expressed in adipose tissue, whereas LepR(S3) is expressed abundantly in brain and spleen, and moderately in liver and gills. The mRNA levels of hepatic LepR(S3) increase after 2 weeks of fasting. This study demonstrates a mechanism in fish for the generation of LepBPs that differs from that seen in mammals and indicates that the physiologic action of Lep in these poikilothermic vertebrates can be modulated, both centrally and peripherally, by the differentiated, tissue-specific expression of multiple LepBPs.
32.	Gräns, Albin, 1979, et al. (författare) Aerobic scope fails to explain the detrimental effects on growth resulting from warming and elevated CO2 in Atlantic halibut 2014 Ingår i: Journal of Experimental Biology. - : The Company of Biologists. - 0022-0949 .- 1477-9145. ; 217:5, s. 711-717 Tidskriftsartikel (refereegranskat)abstract As a consequence of increasing atmospheric CO2, the world's oceans are becoming warmer and more acidic. Whilst the ecological effects of these changes are poorly understood, it has been suggested that fish performance including growth will be reduced mainly as a result of limitations in oxygen transport capacity. Contrary to the predictions given by the oxygen- and capacity-limited thermal tolerance hypothesis, we show that aerobic scope and cardiac performance of Atlantic halibut (Hippoglossus hippoglossus) increase following 14–16 weeks exposure to elevated temperatures and even more so in combination with CO2-acidified seawater. However, the increase does not translate into improved growth, demonstrating that oxygen uptake is not the limiting factor for growth performance at high temperatures. Instead, long-term exposure to CO2-acidified seawater reduces growth at temperatures that are frequently encountered by this species in nature, indicating that elevated atmospheric CO2 levels may have serious implications on fish populations in the future.
33.	Hildahl, Jon, 1976, et al. (författare) CHARACTERIZATION OF THE GROWTH HORMONE – INSULIN-LIKE GROWTH FACTOR I SYSTEM DURING ATLANTIC HALIBUT METAMORPHOSIS 2006 Ingår i: 7th International Congress on the Biology of Fish, Book of Abstracts. Konferensbidrag (övrigt vetenskapligt/konstnärligt)abstract The GH-IGF-I system is critical for proper growth and development in fish and thus is likely to play a role in the control of flatfish metamorphosis.. In order to characterize the role of GH and IGF-I in Atlantic halibut development, hormonal content (GH and IGF-I RIA) and receptor gene expression (GHR and IGF-IR QPCR) was assessed during normal and abnormal metamorphosis GH larval head content increases throughout normal metamorphosis in parallel with larval weight. The full-length GH receptor (hhGHRfl) mRNA and IGF-I protein levels change in parallel and in a stage-specific manner, with an increase at the onset of metamorphosis followed by down-regulation. Two isoforms of the IGF-I receptor are regulated differently; isoform one (hhIGF-IR1) mRNA increases from pre-metamorphosis to the post-metamorphic juvenile stage, while isoform 2 (hhIGF-IR2) mRNA is highest at early pre-metamorphosis and decreases prior to the onset of metamorphosis. In larvae experiencing arrested development, mRNA levels for hhGHRfl and hhIGF-IR1 and IGF-I larval content are depressed relative to normal individuals, whereas GH content, hhIGF-IR2 mRNA levels and weight are not changed. Increased hhGHRfl mRNA and IGF-I protein levels at the onset of metamorphosis suggest that tissue sensitivity to GH increases during metamorphosis while tissue sensitivity to IGF-I does not change. The data provide new evidence suggesting that GHR and IGF-IR levels may be of critical importance for metamorphic success in Atlantic halibut. Funded by the EU and FORMAS.
34.	Hildahl, Jon, 1976, et al. (författare) Cloning of Atlantic halibut growth hormone receptor genes and quantitative gene expression during metamorphosis 2007 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 151:2, s. 143-152 Tidskriftsartikel (refereegranskat)abstract To gain insight into the possible regulatory role of the growth hormone (GH)-insulin-like growth factor I (IGF-I) system in flatfish metamorphosis, body GHR gene expression as well as IGF-I protein content was quantified in larval Atlantic halibut throughout metamorphosis (developmental stages 5–10). The cDNA of the full-length GH receptor (hhGHR) was cloned from adult liver and characterized. The hhGHR shows common features of a GHR, including a (Y/F)GEFS motif in the extracellular domain, a single transmembrane region, and an intracellular domain containing a Box 1 and Box 2. Additionally, a truncated GHR (hhGHRtr), similar to turbot and Japanese flounder GHRtr, was cloned and sequenced. These sequences are highly similar to the full-length and truncated GHRs in turbot (89%/86%) and Japanese flounder (93%/91%) with lower identity with other fish type I GHR (less-than-or-equals, slant81%) and type II GHRs (less-than-or-equals, slant58%). A quantitative real-time RT-PCR assay was used to measure hhGHR and hhGHRtr mRNA content in normally and abnormally metamorphosed individuals at six developmental stages, from early pre-metamorphosis to post-metamorphosis, when the fish is considered a juvenile. The level of hhGHR gene expression was highest at pre-metamorphic stage 6 and at stage 8 at the onset of metamorphosis, and then decreased during metamorphic climax and post-metamorphosis. Expression of hhGHRtr reached highest levels at stage 6 and then decreased to post-metamorphosis. The ratio of expression between the full-length and the truncated GHR (hhGHR:hhGHRtr) varied among stages and was highest at the onset of metamorphosis and at metamorphic climax. A radioimmunoassay was used to measure halibut IGF-I body content throughout metamorphosis. IGF-I increases from early metamorphosis to the onset of metamorphosis and then decreases towards post-metamorphosis. In comparison between normally and abnormally metamorphosing larvae, IGF-I content, hhGHR and hhGHRtr mRNA levels were reduced in the abnormal fish. These data indicate that the GH-IGF-I system either has a regulatory role in metamorphosis, or is being affected as a consequence of the abnormal metamorphosis.
35.	Hildahl, Jon, 1976, et al. (författare) Identification of two isoforms of Atlantic halibut insulin-like growth factor-I receptor genes and quantitative gene expression during metamorphosis 2007 Ingår i: Comparative Biochemistry and Physiology Part B: Biochemistry and Molecular Biology. - : Elsevier BV. - 1096-4959. ; 147:3, s. 395-401 Tidskriftsartikel (refereegranskat)abstract Insulin-like growth factor-I (IGF-I) is an important regulator of growth and development in vertebrates. Both the endocrine and paracrine actions of IGF-I are mediated through ligand-binding to a membrane-bound IGF-I receptor (IGF-IR). The characterization of this receptor and subsequent expression studies thus help elucidate the endocrine regulation of developmental processes. As other flatfish species, the Atlantic halibut (Hippoglossus hippoglossus) undergoes a dramatic larval metamorphosis. This process is largely under endocrine control, and data indicate that IGF-I could be a key regulator. IGF-I content increases up to late pre-metamorphosis and decreases during metamorphosis. The IGF-IR has, however, not been studied during flatfish metamorphosis. To examine IGF-IR gene expression, two IGF-IR mRNA were cloned and sequenced. These partial sequences share high identity (≥ 95%) and similarity (≥ 97%) with other fish IGF-IR and lower identity (≥ 77%) and similarity (≥ 83.5%) with Japanese flounder insulin receptors. The expression of mRNA for both IGF-IR was analyzed by quantitative real-time RT-PCR during six larval developmental stages from pre- to post-metamorphosis. IGF-IR1 and IGF-IR2 mRNA are differentially expressed during metamorphosis, but if this indicates an isoform-specific regulation of developmental processes by circulating and/or locally-secreted IGF-I is unclear. Both IGF-IR genes are down-regulated in halibut larvae experiencing arrested metamorphosis, suggesting the IGF-I system is critical for metamorphic success in halibut.
36.	Hildahl, Jon, 1976, et al. (författare) Involvement of growth hormone-insulin-like growth factor I system in cranial remodeling during halibut metamorphosis as indicated by tissue- and stage-specific receptor gene expression and the presence of growth hormone receptor protein. 2008 Ingår i: Cell and tissue research. - : Springer Science and Business Media LLC. - 0302-766X .- 1432-0878. ; 332:2, s. 211-25 Tidskriftsartikel (refereegranskat)abstract The role of growth hormone (GH) and insulin-like growth factor-I (IGF-I) in the tissue remodeling associated with the transition of a symmetrical larva to an asymmetrical juvenile during flatfish metamorphosis is unknown. In order to investigate the potential role of these hormones in the remodeling of cranial bone and soft tissue that accompanies eye migration during metamorphosis of Atlantic halibut (Hippoglossus hippoglossus) larvae, tissue-specific gene expression was monitored by in situ hybridization for Atlantic halibut type I growth hormone receptor (hhGHR), type II hhGHR, and insulin-like growth factor-I receptor (hhIGF-IR). Polyclonal antibody generated against the extracellular domain of type I hhGHR was used for the immunohistochemical localization of type I GHR protein. Type I hhGHR, type II hhGHR, and hhIGF-IR mRNA were expressed in fibroblasts, frontal bone osteocytes, and dorsal chondrocytes at the onset of metamorphosis (stage 8), during metamorphic climax (stage 9), and in fully metamorphosed juveniles (stage 10). Type I GHR protein showed similar expression patterns to those of type I hhGHR mRNA, except in chondrocytes in which little GHR protein was detected. The localization of GHR and IGF-IR transcripts and GHR protein in cranial structures that undergo remodeling is intriguing and suggests that, in addition to thyroid hormones, the GH-IGF-I system is involved in morphological transformations during metamorphosis in Atlantic halibut.
37.	Johansson, Marcus, 1982, et al. (författare) Energy stores, lipid mobilization and leptin endocrinology of rainbow trout 2016 Ingår i: Journal of Comparative Physiology B-Biochemical Systemic and Environmental Physiology. - : Springer Science and Business Media LLC. - 0174-1578. ; 186:6, s. 759-773 Tidskriftsartikel (refereegranskat)abstract The physiological role of leptin in fish is not fully elucidated. In the present study, the involvement of the leptin system in lipid deposition and mobilization in rainbow trout during feeding and 1, 2 and 4 weeks of fasting was investigated in two lines of rainbow trout with different muscle and visceral adiposity: a fat line (FL) with high total energy reserves, high muscle adiposity, but low visceral adiposity and a lean line (LL) with lower total energy reserves and lower muscle adiposity, but higher visceral adiposity. During 4 weeks of fasting, muscle lipids decreased by 63 % in the FL fish, while no such energy mobilization from muscle occurred in the LL fish. On the other hand, lipid stores in liver and visceral adipose tissue was utilized to a similar extent by the two fish lines during fasting. Under normal feeding conditions, plasma leptin levels were higher in the LL than the FL fish, suggesting a possible contribution of visceral adipocytes to plasma leptin levels. Plasma leptin-binding protein levels did not differ between the lines and were not affected by fasting. After 4 weeks of fasting, the long leptin receptor and the leptin-binding protein isoforms 1 and 3 muscle expression increased in the LL fish, as well as hepatic expression of leptin A1 and the two binding protein isoforms. These responses were not seen in the FL fish. The data suggest that the Lep system in rainbow trout is involved in regulation of energy stores and their mobilization.
38.	Jönsson, Elisabeth, 1968, et al. (författare) Circulating levels of cholecystokinin and gastrin-releasing peptide in rainbow trout fed different diets. 2006 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 148:2, s. 187-194 Tidskriftsartikel (refereegranskat)abstract Cholecystokinin (CCK) and gastrin-releasing peptide (GRP) are gastrointestinal peptides thought to be important regulators of intake and digestion of food in vertebrates. In this study, pre- and postprandial plasma levels of CCK and GRP were measured in rainbow trout (Oncorhynchus mykiss) by the establishment of homologous radioimmunoassays, and the hormonal levels assessed in relation to dietary lipid:protein ratio and food intake. Fish were acclimated to either a high protein/low lipid diet (HP/LL diet; 14.1% lipids) or a normal protein/high lipid diet (NP/HL diet; 31.4% lipids). On three consecutive sampling days, radio-dense lead-glass beads were included in the diets for assessment of feed intake. Fish were terminally sampled for blood and stomach contents prior to feeding at time 0, and at 0.3, 1, 2, 4, 6, and 24 h after feeding. There was a postprandial elevation of plasma CCK levels, which was most evident after 4 and 6 h. Fish fed the NP/HL diet had higher plasma CCK levels compared with those fed the HP/LL diet. Plasma CCK levels were not affected by the amount of food ingested. GRP levels in plasma were not influenced by sampling time, diet, or feed intake. The results indicate that the endocrine release of gastrointestinal CCK is increased during feeding and may be further influenced by the dietary lipid:protein ratio in rainbow trout. Plasma GRP levels, on the other hand, appear not to be influenced by feeding or diet composition.
39.	Jönsson, Elisabeth, 1968, et al. (författare) Plasma ghrelin levels in rainbow trout in response to fasting, feeding and food composition, and effects of ghrelin on voluntary food intake 2007 Ingår i: Comparative Biochemistry and Physiology - Part A: Molecular & Integrative Physiology. - : Elsevier BV. - 1095-6433. ; 147:4, s. 1116-1124 Tidskriftsartikel (refereegranskat)abstract Ghrelin, a peptide hormone which stimulates growth hormone (GH) release, appetite and adiposity in mammals, was recently identified in fish. In this study, the roles of ghrelin in regulating food intake and the growth hormone (GH)–insulin-like growth factor I (IGF-I) system of rainbow trout (Oncorhynchus mykiss) were investigated in three experiments: 1) Pre- and postprandial plasma levels of ghrelin were measured in relation to dietary composition and food intake through dietary inclusion of radio-dense lead-glass beads, 2) the effect of a single intraperitoneal (i.p.) injection with rainbow trout ghrelin on short-term voluntary food intake was examined and 3) the effect of one to three weeks fasting on circulating ghrelin levels and the correlation with plasma GH and IGF-I levels, growth and lipid content in the liver and muscle was studied. There was no postprandial change in plasma ghrelin levels. Fish fed a normal-protein/high-lipid (31.4%) diet tended to have higher plasma ghrelin levels than those fed a high-protein/low-lipid (14.1%) diet. Plasma ghrelin levels decreased during fasting and correlated positively with specific growth rates, condition factor, liver and muscle lipid content, and negatively with plasma GH and IGF-I levels. An i.p. ghrelin injection did not affect food intake during 12-hours post-injection. It is concluded that ghrelin release in rainbow trout may be influenced by long-term energy status, and possibly by diet composition. Further, in rainbow trout, ghrelin seems to be linked to growth and metabolism, but does not seem to stimulate short-term appetite through a peripheral action.
40.	Jönsson, Elisabeth, 1968, et al. (författare) Pre- And Postprandial Levels Of Cholesystokinin (CCk) In Rainbow Trout Fed Two Lipid Diets Measured By A Homologous Radioimmunoassay 2004 Ingår i: 5th International Symposium on Fish Endocrinology. Castellon, Spain, September 5-9 2004.. Konferensbidrag (övrigt vetenskapligt/konstnärligt)
41.	Kling, Peter, 1968, et al. (författare) The role of GH in lipid homeostasis, energy utilization and partitioning in rainbow trout: interactions with ghrelin, leptin and insulin-like growth factor I 2012 Ingår i: General and Comparative Endocrinology. - : Elsevier BV. - 0016-6480. ; 175:1, s. 153-162 Tidskriftsartikel (refereegranskat)abstract The growth-promoting effects of in vivo growth hormone (GH) treatment were studied in relation to size and lipid content of energy stores including liver, mesentery, white muscle and belly flap in rainbow trout. In order to elucidate endocrine interactions and links to regulation of growth, adiposity and energy metabolism, plasma levels of GH, insulin-like growth factor I (IGF-I), leptin (Lep) and ghrelin, were assessed and correlated to growth and energy status. In addition tissue-specific expression of lepa1 mRNA was examined. Juvenile rainbow trout were implanted with sustained-release bovine GH implants and terminally sub-sampled at 1, 3 and 6 weeks. GH increased specific growth rate, reduced condition factor (CF) and increased feed conversion efficiency resulting in a redistribution of energy stores. Thus, GH decreased mesenteric (MSI) and liver somatic index (LSI). Lipid content of the belly flap increased following GH-treatment while liver and muscle lipid content decreased. Independent of GH substantial growth was accompanied by an increase in muscle lipids and a decrease in belly flap lipids. The data suggest that the belly flap may function as an energy buffering tissue during episodes of feeding and lean growth. Liver and muscle lipids were positively correlated to body weight, indicating a size-dependent change in adiposity. Hepatic lepa1 mRNA positively correlated to MSI and CF and its expression decreased following GH treatment, coinciding with decreased hepatic lipid content. Plasma Lep was positively correlated to MSI and belly flap lipid content, suggesting that Lep may communicate energy status. In summary, the observed GH tissue-specific effects on lipid metabolism in rainbow trout highlight the complex physiology of the energy reserves and their endocrine control.
42.	Power, Deborah M, et al. (författare) The molecular and endocrine basis of flatfish metamorphosis 2008 Ingår i: Reviews in Fisheries Science. - : Informa UK Limited. - 1064-1262 .- 1547-6553. ; 16:S1, s. 93-109 Tidskriftsartikel (refereegranskat)abstract A significant component of aquaculture is the production of good quality larvae, and, in the case of flatfish, this is tied up with the change from a symmetric larva to an asymmetric juvenile. Despite the pioneering work carried out on the metamorphosis of the Japanese flounder (Paralichthys olivaceus) and summer flounder (Paralichthys dentatus), the underlying molecular basis of flatfish metamorphosis is still relatively poorly characterized. It is a thyroid hormone (TH) driven process, and the role of other hormones in the regulation of the process along with the interplay of abiotic factors are still relatively poorly characterized as is the extent of tissue and organ remodeling, which underlie the profound structural and functional modifications that accompany the larval/juvenile transition. The isolation of genes for hormones, receptors, binding proteins, and other accessory factors has provided powerful tools with which to pursue this question. The application of molecular methodologies such as candidate gene approaches and microarray analysis coupled to functional genomics has started to contribute to understanding the complexity of tissue and organ modifications that accompany flatfish metamorphosis. A better understanding of the biology of normal metamorphosis is essential to identify factors contributing to abnormal metamorphosis.
43.	Power, DM, et al. (författare) The Ontogeny and Tissue Distribution of Ghrelin and its receptor during Metamorphosis of the Atlantic Halibut (Hippoglossus hippoglossus). 2008 Ingår i: 6th International Symposium on Fish Endocrinology, calgary, Canada. 2008.. Konferensbidrag (övrigt vetenskapligt/konstnärligt)
44.	Power, DM, et al. (författare) Thyroid hormones in growth and development of fish 2001 Ingår i: COMPARATIVE BIOCHEMISTRY AND PHYSIOLOGY. - 1532-0456. ; 130:4, s. 447-459 Tidskriftsartikel (refereegranskat)abstract The thyroid hormones (THs), thyroxine (T-4) and triiodothyronine (T-3) are products of the thyroid gland in all vertebrates. Their role in early development and metamorphosis is well established in mammals and amphibians, respectively, and recently several studies in fish have highlighted the importance of THs during flatfish metamorphosis. THs are present in high quantities in fish eggs and are presumably of maternal origin. During embryogenesis the concentration of T-4 and T-3 in the eggs decrease until endogenous production starts. Thyroid hormone receptors (TR) have been isolated from several teleosts and in common with tetrapods two receptor isoforms have been identified, TR alpha and TR beta. Both the receptors are expressed in early embryos and larvae of the Japanese flounder (Paralichthys olivaceus), zebrafish (Danio rerio) and seabream (Sparus aurata) although a different temporal pattern is apparent. The role of THs and TRs in fish embryogenesis, larval development and during metamorphosis will be discussed. (C) 2001 Elsevier Science Inc. All rights reserved.
45.	Sweeney, GE, et al. (författare) Endocrine Control as Determinant of Larval Quality in Fish Aquaculture 2001 Rapport (övrigt vetenskapligt/konstnärligt)
46.	Thorstad, E. B., et al. (författare) Reduced marine survival of hatchery-reared Atlantic salmon post-smolts exposed to aluminium and moderate acidification in freshwater 2013 Ingår i: Estuarine Coastal and Shelf Science. - : Elsevier BV. - 0272-7714. ; 124, s. 34-43 Tidskriftsartikel (refereegranskat)abstract Short-term Al-exposure and moderate acidification increased initial marine mortality in migrating post-smolts, and can thereby reduce viability of Atlantic salmon stocks. The delayed impact of short-term aluminium (Al) exposure on hatchery-reared Atlantic salmon smolt in moderately acidified freshwater (pH 5.88–5.98) was investigated during the first 37 km of the marine migration. Smolts were tagged with acoustic tags and exposed to low (28.3 ± 4.6 μg l−1 labile Al, 90 h) or high (48.5 ± 6.4 μg l−1 labile Al, 90 or 48 h) Al concentrations within the hatchery. Thereafter their movements, together with a control group, were monitored throughout the marine fjord. Al-exposure resulted in increased gill-Al and compromised hypoosmoregulatory capacity, as shown by elevated mortality in laboratory seawater challenge tests and reduced Na+, K+-ATPase activity levels. Further, Al-exposure resulted in decreased plasma concentrations of growth hormone (GH), while the insulin-like growth factor (IGF-I) was unaffected. There was a significant mortality in the 90 h high-Al group during exposure, and those surviving until release died during the first 3.6 km of the marine migration. Physiological stress and mortality were not only a result of the Al-concentrations, but also dependent on exposure duration, as shown by results from the 48 h high-Al group. Elevated mortality was not recorded in freshwater or after entering the sea for this group, which highly contrasts to the 100% mortality in the 90 h high-Al group, despite both groups having similarly high gill-Al levels. The low-Al group showed a 20% higher mortality compared to the control group during the first 10 km of the marine migration, but during the next 28 km, mortality rates did not differ. Hence, post-smolts surviving the first 10 km subsequently showed no differences in mortality compared to controls. At least one third of the mortality in both the low-Al and control groups were due to predation by marine fishes, indicating that the proximate cause for elevated mortality due to Al-exposure may have been predation. Migration speeds over 3.6, 9.6 or 37.1 km from the release site was not affected by Al-exposure.
47.	Valenzuela, C. A., et al. (författare) Chronic stress inhibits growth and induces proteolytic mechanisms through two different nonoverlapping pathways in the skeletal muscle of a teleost fish 2018 Ingår i: American Journal of Physiology-Regulatory Integrative and Comparative Physiology. - : American Physiological Society. - 0363-6119 .- 1522-1490. ; 314:1 Tidskriftsartikel (refereegranskat)abstract Chronic stress detrimentally affects animal health and homeostasis, with somatic growth, and thus skeletal muscle, being particularly affected. A detailed understanding of the underlying endocrine and molecular mechanisms of how chronic stress affects skeletal muscle growth remains lacking. To address this issue, the present study assessed primary (plasma cortisol), secondary (key components of the GH/IGF system, muscular proteolytic pathways, and apoptosis), and tertiary (growth performance) stress responses in fine flounder (Paralichthys adspersus) exposed to crowding chronic stress. Levels of plasma cortisol, glucocorticoid receptor 2 (gr2), and its target genes (klf15 and redd1) mRNA increased significantly only at 4 wk of crowding (P < 0.05). The components of the GH/IGF system, including ligands, receptors, and their signaling pathways, were significantly downregulated at 7 wk of crowding (P < 0.05). Interestingly, chronic stress upregulated the ubiquitin-proteasome pathway and the intrinsic apoptosis pathways at 4wk (P < 0.01), whereas autophagy was only significantly activated at 7 wk (P < 0.05), and meanwhile the ubiquitin-proteasome and the apoptosis pathways returned to control levels. Overall growth was inhibited in fish in the 7-wk chronic stress trial (P < 0.05). In conclusion, chronic stress directly affects muscle growth and downregulates the GH/IGF system, an action through which muscular catabolic mechanisms are promoted by two different and nonoverlapping proteolytic pathways. These findings provide new information on molecular mechanisms involved in the negative effects that chronic stress has on muscle anabolic/catabolic signaling balance.

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