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Fish Physiology, Toxicology, and Water Quality Proceedings of The Fish Physiology, Toxicology, and Water Quality Proceedings of the Eighth International Symposium Chongqing, China October 12-14, 2004 R E S E A R C H A N D D E V E L O P M E N T EPA/600/R-06/062 June 2006 FISH PHYSIOLOGY, TOXICOLOGY, AND WATER QUALITY Proceedings of the Eighth International Symposium, Chongqing, China October 12-14, 2004 Edited By David Randall and Dung Yuk Man Mandy City University of Hong Kong Published by Ecosystems Research Division Athens, Georgia 30605 U.S. Environmental Protection Agency Office of Research and Development Washington, DC 20460 NOTICE The views expressed in these Proceedings are those of the individual authors and do not necessarily reflect the views and policies of the U.S. Environmental Protection Agency (EPA). Scientists in EPA’s Office of Research and Development have authored or coauthored papers presented herein; these papers have been reviewed in accordance with EPA’s peer and administrative review policies and approved for presentation and publication. Mention of trade names or commercial products does not constitute endorsement or recommendation for use by EPA. ii ABSTRACT Twenty-one participants from Europe, North America and China convened in Chongqing, China, October 12-14, 2005, for the Eighth International Symposium in Fish Physiology, Toxicology and Water Quality. The subject of the meeting was “Hypoxia in vertebrates: Comparisons of terrestrial and aquatic vertebrates”. These Proceedings include 13 papers presented over the three day period and discuss the responses of fish, reptiles and mammals to hypoxia. These papers report on the responses of animals to hypoxia at the behavioural, physiological and molecular levels. Clearly hypoxia has wide ranging effects, the responses are complex and there are many similarities in the responses of all vertebrates to hypoxia. Organisms respond to hypoxia by reducing energy expenditure, in particular inhibiting reproduction, feeding and exercise. That is animals only expend energy that is absolutely necessary for survival. There is extensive reorganization of cellular machinery directed by hypoxia inducible factors (HIFs), proteins that increase during hypoxia and have a marked effect on the expression of many genes. As a result, anaerobic metabolism is up regulated and many aspects of aerobic metabolism are down regulated. Many cells enter cell cycle arrest. Thus organisms stop reproducing and growing during hypoxia and if the effects of hypoxia are severe, development of eggs and larvae is compromised. iii CONTENTS Page NOTICE.................................................................................................................................................ii ABSTRACT..........................................................................................................................................iii ACKNOWLEDGEMENTS ..................................................................................................................vi Response of aquatic vertebrates to hypoxia D.J. Randall, C.Y. Hung and W.L. Poon ................................................................................... 1 Post-genomic approaches to the mechanisms of hypoxia response in the common carp, Cyprinus Carpio E.J. Fraser, A.Y. Gracey and A.R. Cossins ............................................................................11 Isolation and characterization of the VEGF-A gene of grass carp Gallant K.L. Chan, Helen O.L. Mok, Patrick K.S. Ng, Minnie M.L. Wong and Richard Y.K. Kong............................................................................................................19 Molecular studies of the HIF transcription factors of grass carp H.W. Law, Richard Y.C. Kong and Rudolf S.S. Wu.................................................................31 Gene expression profiling of raw 264.7 cell survival and apoptosis under oxidative stress C.C. Fong, Y. Zhang, M.S. Wong, C.H. Tzang, W.P. Lai and M .Yang ..................................39 Extreme adaptations to hypoxia and anoxia in crucian carp Göran E. Nilsson .....................................................................................................................53 Suppressive effects of hypoxia on male rat reproductive function Weigong Liao, Yuqi Gao, Mingchun Cai, Jian Chen, Yi Wu ..................................................59 Endocrine disrupting and teratogenic effects of hypoxia on fish, and their ecological implications Rudolf S.S. Wu, Eva W.H. Shang and B.S. Zhou .....................................................................75 Effects of hypoxia on lipid metabolism in teleosts Guido van den Thillart ............................................................................................................87 The influence of oxygen shortage on the development of the cardiovascular system in Zebrafish Bernd Pelster .........................................................................................................................107 A biocomputational tool to measure cardiac rhythms in zebrafish S.H. Cheng, Eric P.K. Chan and Alex C.C. Lin ....................................................................121 iv Hypoxia in reptiles Tobias Wang and Nini Skovgaard.........................................................................................127 Bioaccumulation and enantioselective biotransformation of fipronil by rainbow trout (Oncorhynchus mykiss) Brad J. Konwick, Aaron T. Fisk, Jimmy K. Avants, J. MacArthur Long and Arthur W. Garrison ...............................................................................................................139 v ACKNOWLEDGEMENTS The 8th Symposium was organized by Professor Gao Yuki of the Institute of High Altitude Medicine, Third Military University, Chongqing, China and Professor David Randall of the Department of Biology and Chemistry, City University of Hong Kong, with the able help of Ms. Mandy Dung Yuk Man, also from Biology and Chemistry, City University of Hong Kong. The meeting was managed in Chongqing by the Third Military University, who provided excellent meeting rooms for the Symposium. Professor Gao and his associates were excellent hosts and showed us both the University and Chongqing. Funding for the symposium was provided by the U.S. Environmental Protection Agency and we were advised throughout by Dr. George Bailey and Dr. Rosemarie Russo of the Ecosystems Research Division, U.S. Environmental Protection Agency, Athens, Georgia. Ms. Dung Yuk Man assisted in the preparation of this document. Finally I attribute the success of this symposium to the many people who came to Chongqing to present and discuss their work. David Randall Department of Biology and Chemistry, City University of Hong Kong vi RESPONSE OF AQUATIC VERTEBRATES TO HYPOXIA D.J.Randall1, C.Y.Hung and W.L.Poon ABSTRACT The major effect of hypoxia on the individual is to reduce exercise capacity. Fish respond to hypoxia by inhibiting feeding and reproduction, and moving to lower temperatures, all of which lower energy expenditure. Oxygen delivery is augmented by increasing both gill ventilation and hemoglobin content and oxygen affinity. Aerobic metabolism is down-regulated whereas anaerobic metabolism is up-regulated. Steroid metabolism is reduced. Liver cells go into cell cycle arrest. Genes associated with cell growth and aerobic metabolism are down regulated but genes associated with anaerobic metabolism are up-regulated. Uncoupling proteins 2 & 3 are up-regulated and may play a role in reducing mitochondrial activity during hypoxia. Much of the response to hypoxia occurs in the first few hours and days. There are many similarities between fish and mammalian responses to hypoxia, presumably because they evolved in common aquatic vertebrate ancestors. Cardiovascular responses have been somewhat modified in terrestrial vertebrates and organic phosphate levels in the red blood cell decrease rather than increase in fish exposed to hypoxia. The relative role of temperature change, starvation, adenosine production, and HIF-1 expression in metabolic depression is not clear in any vertebrate. INTRODUCTION Hypoxia is a common feature of the aquatic environment. Oxygen levels in the water depend on photosynthetic activity in the water, exchange with the atmosphere, usage by aquatic organisms and oxidation reactions in the water. Photosynthesis requires light and, therefore, is restricted to surface waters and during daylight. Nocturnal hypoxia is common in tropical lakes and lagoons. Gas diffusion in water is slow and so hypoxia occurs at depth in unmixed waters. There is a minimum oxygen layer in oceans, blow the photosynthetic zone where a large number of organisms live on food dropping from above. Hypoxia is also common beneath the ice on frozen lakes and ponds. Although hypoxia is a natural and frequent event in the aquatic environment, it has been exacerbated by human actions. Eutrophication of waterways generally leads to hypoxia, and many regions of the world where there are large human populations have increasing levels of hypoxia. The movement of vertebrates onto land is relatively recent and much of vertebrate evolution occurred in water at lower oxygen levels than at present. It is possible that terrestrial vertebrates, living in an oxygen rich atmosphere, have lost some of their capacity to survive hypoxia. EFFECTS OF INCREASING HYPOXIA ON AQUATIC VERTBRATES Increasing aquatic hypoxia has led to changes in species composition, some leave, some die, what is left are those more tolerant of
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