{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T01:48:43Z","timestamp":1789696123945,"version":"4.0.1"},"reference-count":57,"publisher":"The Company of Biologists","issue":"1","license":[{"start":{"date-parts":[[1994,9,1]],"date-time":"1994-09-01T00:00:00Z","timestamp":778377600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[1994,9,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>This study tests the hypothesis that the increase in blood and associated respiratory acidosis after exhaustive exercise play an important role in stimulating ventilation during post-exercise recovery in fish. Injection of bovine carbonic anhydrase (10 mg kg\u22121) into the bloodstream of rainbow trout caused a persistent 40% increase in the HCO3\u2212 dehydration capacity of the blood. The treatment was designed to increase CO2 excretion and therefore to reduce build-up and acidosis after exercise. Aerobic and anaerobic swimming performance were not affected by carbonic anhydrase, and there were only very minor effects on arterial blood acid\u2013base status in resting fish. However, carbonic anhydrase attenuated post-exercise increases in and decreases in pHa by about 50% without altering arterial O2 variables, red cell swelling or the intracellular pH of the brain or muscle tissues. The effects on arterial pH (pHa) resulted largely from alleviation of the increase in . In accordance with the original hypothesis, normal post-exercise hyperventilation was greatly attenuated, through reductions in both ventilatory stroke volume and frequency, and excess post-exercise O2 consumption was reduced. Post-exercise increases in plasma levels of adrenaline and noradrenaline were also reduced by the carbonic anhydrase treatment. Overall, there was a strong correlation between increases in relative ventilation and decreases in pHa after exhaustive exercise. The results provide functional significance for the phenomenon of elevation and associated respiratory acidosis after exercise and are consistent with other recent studies indicating an important secondary drive to ventilation in fish based on arterial acid\u2013base status, in addition to the primary drive based on arterial O2 levels.<\/jats:p>","DOI":"10.1242\/jeb.194.1.225","type":"journal-article","created":{"date-parts":[[2021,4,23]],"date-time":"2021-04-23T23:25:26Z","timestamp":1619220326000},"page":"225-253","source":"Crossref","is-referenced-by-count":82,"title":["Carbonic Anhydrase Injection Provides Evidence for the Role of Blood Acid\u2013Base Status in Stimulating Ventilation after Exhaustive Exercise in Rainbow Trout"],"prefix":"10.1242","volume":"194","author":[{"given":"Chris M.","family":"Wood","sequence":"first","affiliation":[{"name":"McMaster University Department of Biology , , 1280 Main Street West, Hamilton, Ontario, Canada L8S 4K1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"R. S.","family":"Munger","sequence":"additional","affiliation":[{"name":"McMaster University Department of Biology , , 1280 Main Street West, Hamilton, Ontario, Canada L8S 4K1"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","reference":[{"key":"2023051512351850100_JEXBIO_194_1_225C1","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1242\/jeb.151.1.57","article-title":"A possible role for catecholamines in the ventilatory responses associated with internal acidosis or external hypoxia in rainbow trout Oncorhynchus mykiss","volume":"151","author":"Aota","year":"1990","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C2","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1111\/j.1095-8649.1973.tb04510.x","article-title":"Routine haematological methods for use with fish blood","volume":"5","author":"Blaxhall","year":"1973","journal-title":"J. Fish. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C3","first-page":"403","article-title":"Physico-chemical parameters for use in fish respiratory physiology","volume-title":"Fish Physiology","author":"Boutilier","year":"1984"},{"key":"2023051512351850100_JEXBIO_194_1_225C4","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1242\/jeb.123.1.145","article-title":"The promotion of catecholamine release in rainbow trout, Salmo gairdneri, by acute acidosis: interactions between red cell pH and haemoglobin oxygen-carrying capacity","volume":"123","author":"Boutilier","year":"1986","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C5","doi-asserted-by":"crossref","first-page":"381","DOI":"10.1016\/0034-5687(78)90125-1","article-title":"Some response characteristics of CO2 electrodes","volume":"32","author":"Boutilier","year":"1978","journal-title":"Respir. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C6","doi-asserted-by":"crossref","first-page":"1183","DOI":"10.1139\/f64-103","article-title":"The respiratory metabolism and swimming performance of young sockeye salmon","volume":"21","author":"Brett","year":"1964","journal-title":"J. Fish. Res. Bd Can."},{"key":"2023051512351850100_JEXBIO_194_1_225C7","doi-asserted-by":"crossref","first-page":"632","DOI":"10.1152\/jappl.1971.31.4.632","article-title":"Rapid method of determination of total carbon dioxide in small blood samples","volume":"31","author":"Cameron","year":"1971","journal-title":"J. appl. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C8","doi-asserted-by":"crossref","first-page":"1627","DOI":"10.1139\/f70-183","article-title":"Evaluation of opercular catheterization as a method for sampling water expired by fish","volume":"27","author":"Davis","year":"1970","journal-title":"J. Fish. Res. Bd Can"},{"key":"2023051512351850100_JEXBIO_194_1_225C9","first-page":"117","article-title":"Problems of control of breathing in fishes","volume-title":"Comparative Physiology: Locomotion, Respiration, Transport and Blood","author":"Dejours","year":"1973"},{"key":"2023051512351850100_JEXBIO_194_1_225C10","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1016\/0034-5687(90)90088-G","article-title":"Control of ventilation in the hypercapnic skate, Raja ocellata. I. Blood and extradural fluid","volume":"80","author":"Graham","year":"1990","journal-title":"Respir. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C11","doi-asserted-by":"crossref","first-page":"1518","DOI":"10.1139\/z81-206","article-title":"Toxicity of environmental acid to the rainbow trout: interactions of water hardness, acid type and exercise","volume":"59","author":"Graham","year":"1981","journal-title":"Can. J. Zool."},{"key":"2023051512351850100_JEXBIO_194_1_225C12","first-page":"215","article-title":"Acid\u2013base regulation","volume-title":"Physiology of Elasmobranch Fishes","author":"Heisler","year":"1989"},{"key":"2023051512351850100_JEXBIO_194_1_225C13","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/0034-5687(88)90018-7","article-title":"Regulation of ventilation and acid\u2013base status in the elasmobranch Scyliorhinus stellaris during hyperoxia-induced hypercapnia","volume":"71","author":"Heisler","year":"1988","journal-title":"Respir. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C14","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1242\/jeb.46.2.297","article-title":"Changes in blood pressure in the rainbow trout during hypoxia","volume":"46","author":"Holeton","year":"1967","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C15","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1242\/jeb.175.1.115","article-title":"Propranolol impairs the hyperventilatory response to acute hypercapnia in rainbow trout","volume":"175","author":"Kinkead","year":"1993","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C16","doi-asserted-by":"crossref","first-page":"1087","DOI":"10.1086\/physzool.64.4.30157958","article-title":"The role of circulating catecholamines in the ventilatory and hypertensive responses to hypoxia in the Atlantic cod, Gadus morhua","volume":"64","author":"Kinkead","year":"1991","journal-title":"Physiol. Zool."},{"key":"2023051512351850100_JEXBIO_194_1_225C17","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1007\/BF01075676","article-title":"An investigation of the role of circulating catecholamines in the control of ventilation during acute moderate hypoxia in rainbow trout (Oncorhynchus mykiss)","volume":"160B","author":"Kinkead","year":"1990","journal-title":"J. comp. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C18","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/0034-5687(91)90020-J","article-title":"The effects of catecholamines on ventilation in rainbow trout during hypoxia or hypercapnia","volume":"84","author":"Kinkead","year":"1991","journal-title":"Respir. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C19","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1016\/0034-5687(85)90091-X","article-title":"A mathematical model for gas exchange in the fish gill based on non-linear blood gas equilibrium curves","volume":"62","author":"Malte","year":"1985","journal-title":"Respir. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C20","doi-asserted-by":"crossref","first-page":"2765","DOI":"10.1139\/f69-271","article-title":"Rapid cold-branding of salmon and trout with liquid nitrogen","volume":"26","author":"Mighell","year":"1969","journal-title":"J. Fish. Res. Bd Can."},{"key":"2023051512351850100_JEXBIO_194_1_225C21","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1242\/jeb.135.1.119","article-title":"In vivo lactate kinetics at rest and during recovery from exhaustive exercise in coho salmon (Oncorhynchus kisutch) and starry flounder (Platichthys stellatus)","volume":"135","author":"Milligan","year":"1988","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C22","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1242\/jeb.99.1.397","article-title":"Disturbances in haematology, fluid volume distribution and circulatory function associated with low environmental pH in the rainbow trout, Salmo gairdneri","volume":"99","author":"Milligan","year":"1982","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C23","first-page":"R668","article-title":"Intracellular pH transients in rainbow trout tissues measured by dimethadione distribution","volume":"248","author":"Milligan","year":"1985","journal-title":"Am. J. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C24","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1242\/jeb.123.1.93","article-title":"Intracellular and extracellular acid\u2013base status and H+ exchange with the environment after exhaustive exercise in the rainbow trout","volume":"123","author":"Milligan","year":"1986","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C25","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1242\/jeb.123.1.123","article-title":"Tissue intracellular acid\u2013base status and the fate of lactate after exhaustive exercise in the rainbow trout","volume":"123","author":"Milligan","year":"1986","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C26","doi-asserted-by":"crossref","first-page":"C728","DOI":"10.1152\/ajpcell.1989.256.4.C728","article-title":"Na+\u2013H+ exchange and pH regulation in red blood cells: role of uncatalyzed H2CO3 dehydration","volume":"256","author":"Motais","year":"1989","journal-title":"Am. J. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C27","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/BF02265152","article-title":"Extracellular fluid volume measurements in tissues of the rainbow trout (Oncorhynchus mykiss) in vivo and their effects on intracellular pH and ion calculations","volume":"9","author":"Munger","year":"1991","journal-title":"Fish Physiol. Biochem."},{"key":"2023051512351850100_JEXBIO_194_1_225C28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1242\/jeb.105.1.1","article-title":"Cardiac output and regional blood flow in gills and muscles after exhaustive exercise in rainbow trout (Salmo gairdneri)","volume":"105","author":"Neumann","year":"1983","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C29","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1242\/jeb.154.1.257","article-title":"Adrenergic control of red cell pH in salmonid fish: roles of the sodium\/proton exchange, Jacobs\u2013Stewart cycle and membrane potential","volume":"154","author":"Nikinmaa","year":"1990","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C30","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1242\/jeb.180.1.315","article-title":"Pre-and postbranchial carbon dioxide content of rainbow trout (Oncorhynchus mykiss) blood after catecholamine injection","volume":"180","author":"Nikinmaa","year":"1993","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C31","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1139\/z86-083","article-title":"Carbon dioxide excretion in fishes","volume":"64","author":"Perry","year":"1986","journal-title":"Can. J. Zool."},{"key":"2023051512351850100_JEXBIO_194_1_225C32","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1242\/jeb.180.1.39","article-title":"An evaluation of factors limiting carbon dioxide excretion by trout red blood cells in vitro","volume":"180","author":"Perry","year":"1993","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C33","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1007\/BF00042917","article-title":"Are circulating catecholamines involved in the control of breathing by fishes?","volume":"2","author":"Perry","year":"1992","journal-title":"Rev. Fish Biol. Fisheries"},{"key":"2023051512351850100_JEXBIO_194_1_225C34","first-page":"39","article-title":"The role of carbonic anhydrase in carbon dioxide excretion, acid\u2013base balance and ionic regulation in aquatic gill breathers. In Animal Nutrition and Transport Processes 2. Transport, Respiration and Excretion Comparative and Environmental Aspects","volume-title":"Comp. Physiol.","author":"Perry","year":"1990"},{"key":"2023051512351850100_JEXBIO_194_1_225C35","doi-asserted-by":"crossref","first-page":"2961","DOI":"10.1139\/z89-419","article-title":"Control and coordination of gas transfer in fishes","volume":"67","author":"Perry","year":"1989","journal-title":"Can. J. Zool."},{"key":"2023051512351850100_JEXBIO_194_1_225C36","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1242\/jeb.157.1.367","article-title":"Adrenergic inhibition of carbon dioxide excretion by trout red blood cells in vitro is mediated by activation of Na+\/H+exchange","volume":"157","author":"Perry","year":"1991","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C37","doi-asserted-by":"crossref","first-page":"2956","DOI":"10.1139\/z89-418","article-title":"Factors affecting gas transfer in respiratory organs of vertebrates","volume":"67","author":"Piiper","year":"1989","journal-title":"Can. J. Zool."},{"key":"2023051512351850100_JEXBIO_194_1_225C38","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1242\/jeb.152.1.353","article-title":"Effects of catecholamines on gas exchange and ventilation in rainbow trout (Salmo gairdneri)","volume":"152","author":"Playle","year":"1990","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C39","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1242\/jeb.100.1.275","article-title":"The control of respiration and circulation in fish during exercise and hypoxia","volume":"100","author":"Randall","year":"1982","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C40","first-page":"255","article-title":"Catecholamines","volume-title":"Fish Physiology","author":"Randall","year":"1992"},{"key":"2023051512351850100_JEXBIO_194_1_225C41","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1007\/BF00157582","article-title":"Evidence of a role for catecholamines in the control of breathing in fish","volume":"1","author":"Randall","year":"1991","journal-title":"Rev. Fish Biol. Fisheries"},{"key":"2023051512351850100_JEXBIO_194_1_225C42","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1242\/jeb.167.1.155","article-title":"Gas exchange, metabolite status and excess post-exercise oxygen consumption after repetitive bouts of exhaustive exercise in juvenile rainbow trout","volume":"167","author":"Scarabello","year":"1992","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C43","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1152\/jappl.1956.9.2.197","article-title":"Variations of serum carbonic acid pK1 with pH and temperature","volume":"9","author":"Severinghaus","year":"1956","journal-title":"J. appl. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C44","first-page":"857","article-title":"Control of breathing in ectothermic vertebrates","volume-title":"Handbook of Physiology, section 3, The Respiratory System,","author":"Shelton","year":"1986"},{"key":"2023051512351850100_JEXBIO_194_1_225C45","first-page":"11","article-title":"Improved method of dorsal aorta catheterization: haematological effects followed for three weeks in rainbow trout (Salmo gairdneri)","volume":"1","author":"Soivio","year":"1972","journal-title":"Finnish Fish. Res."},{"key":"2023051512351850100_JEXBIO_194_1_225C46","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1242\/jeb.131.1.427","article-title":"Effect of burst swimming and adrenaline infusion on O2 consumption and CO2 excretion in rainbow trout, Salmo gairdneri","volume":"131","author":"Steffensen","year":"1987","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C47","first-page":"R395","article-title":"Correlation between catecholamine release and degree of acidotic stress in trout","volume":"255","author":"Tang","year":"1988","journal-title":"Am. J. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C48","first-page":"160","article-title":"Control and consequences of adrenergic activation of red blood cell Na+\/H+ exchange on blood oxygen and carbon dioxide transport","volume":"263","author":"Thomas","year":"1992","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C49","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1152\/jappl.1967.23.3.410","article-title":"Method for oxygen content and dissociation curves on microliter blood samples","volume":"23","author":"Tucker","year":"1967","journal-title":"J. appl. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C50","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1242\/jeb.160.1.285","article-title":"Acid\u2013base and ion balance, metabolism and their interactions, after exhaustive exercise in fish","volume":"160","author":"Wood","year":"1991","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C51","doi-asserted-by":"crossref","DOI":"10.1016\/0034-5687(94)90079-5","article-title":"HCO3-dehydration by the blood of rainbow trout following exhaustive exercise","volume-title":"Respir. Physiol. (in press)","author":"Wood","year":"1994"},{"key":"2023051512351850100_JEXBIO_194_1_225C52","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1242\/jeb.96.1.221","article-title":"The influence of experimental anaemia on blood acid\u2013base regulation in vivo and in vitro in the starry flounder (Platichthys stellatus) and the rainbow trout (Salmo gairdneri)","volume":"96","author":"Wood","year":"1982","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C53","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1007\/978-3-642-70610-3_1","article-title":"Respiratory, circulatory and metabolic adjustments to exercise in fish","volume-title":"Circulation, Respiration, Metabolism","author":"Wood","year":"1985"},{"key":"2023051512351850100_JEXBIO_194_1_225C54","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1242\/jeb.157.1.349","article-title":"A new in vitro assay for carbon dioxide excretion by trout red blood cells: effects of catecholamines","volume":"157","author":"Wood","year":"1991","journal-title":"J. exp. Biol."},{"key":"2023051512351850100_JEXBIO_194_1_225C55","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1007\/BF00004447","article-title":"The conversion of plasma HCO3-to CO2 by rainbow trout red cells in vitro: adrenergic inhibition and the influence of oxygenation status","volume":"12","author":"Wood","year":"1994","journal-title":"Fish. Physiol. Biochem."},{"key":"2023051512351850100_JEXBIO_194_1_225C56","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/0034-5687(90)90089-H","article-title":"Control of ventilation in the hypercapnic skate Raja ocellata. II. Cerebrospinal fluid and intracellular pH in the brain and other tissues","volume":"80","author":"Wood","year":"1990","journal-title":"Respir. Physiol."},{"key":"2023051512351850100_JEXBIO_194_1_225C57","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1242\/jeb.136.1.149","article-title":"The distribution of ammonia and H+ between tissue compartments in lemon sole (Parophrys vetulus) at rest, during hypercapnia and following exercise","volume":"136","author":"Wright","year":"1988","journal-title":"J. exp. Biol."}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/194\/1\/225\/2855577\/jexbio_194_1_225.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/194\/1\/225\/2855577\/jexbio_194_1_225.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,5,15]],"date-time":"2023-05-15T13:25:48Z","timestamp":1684157148000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/194\/1\/225\/6848\/Carbonic-Anhydrase-Injection-Provides-Evidence-for"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[1994,9,1]]},"references-count":57,"journal-issue":{"issue":"1","published-print":{"date-parts":[[1994,9,1]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.194.1.225","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"value":"0022-0949","type":"print"},{"value":"1477-9145","type":"electronic"}],"subject":[],"published":{"date-parts":[[1994,9,1]]}}}