{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T16:54:14Z","timestamp":1790873654871,"version":"4.1.0"},"reference-count":71,"publisher":"Wiley","issue":"2","license":[{"start":{"date-parts":[[2005,5,9]],"date-time":"2005-05-09T00:00:00Z","timestamp":1115596800000},"content-version":"vor","delay-in-days":4650,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["J. Exp. Zool."],"published-print":{"date-parts":[[1992,8,15]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>This review examines the branchial mechanisms utilized by freshwater fish to regulate internal acid\u2010base status and presents a model to explain the underlying basis of the compensatory processes. Rainbow trout, <jats:italic>Oncorhynchus mykiss<\/jats:italic>, and brown bullhead, <jats:italic>Ictalurus nebulosus<\/jats:italic>, were examined under a variety of experimental treatments which induced respiratory and metabolic acid\u2010base disturbances. Acid\u2010base regulation was achieved by appropriate adjustments of Na <jats:sup>+<\/jats:sup> and Cl<jats:sup>\u2212<\/jats:sup> net fluxes across the gills which, in turn, were accomplished by variable contributions of three different branchial mechanisms: 1) differential changes in Na<jats:sup>+<\/jats:sup> and Cl<jats:sup>\u2212<\/jats:sup> diffusive effluxes, 2) changes in internal substrate (H<jats:sup>+<\/jats:sup>, HCO<jats:sub>3<\/jats:sub> <jats:sup>\u2212<\/jats:sup>) availability, and 3) morphological adjustments to the gill epithelium. <jats:italic>Differential<\/jats:italic> diffusive efflux of Na<jats:sup>+<\/jats:sup> over Cl<jats:sup>\u2212<\/jats:sup> was involved only during periods of metabolic alkalosis. The importance of internal substrate availability was demonstrated using a two\u2010substrate model. According to the model, ionic flux rates (J<jats:sub>in<\/jats:sub><jats:sup>Cl<\/jats:sup><jats:sup>\u2212<\/jats:sup>, J<jats:sub>in<\/jats:sub><jats:sup>Na +<\/jats:sup>) are determined not only by the concentration of the external ion (Na<jats:sup>+<\/jats:sup>, Cl<jats:sup>\u2212<\/jats:sup>) but also by the concentration of the internal counterion (H<jats:sup>+<\/jats:sup>, HCO<jats:sub>3<\/jats:sub><jats:sup>\u2212<\/jats:sup>). This system provides for an \u201cautomatic negative feedback\u201d to aid in the compensation of metabolic acid\u2010base disturbances. Morphological alteration of the gill epithelia and the associated regulation of chloride cell (CC) fractional area is an essential third mechanism which is especially important during respiratory acid\u2010base disturbances. Specifically, fish vary the availability of the CC associated Cl<jats:sup>\u2212<\/jats:sup>\/HCO<jats:sub>3<\/jats:sub> <jats:sup>\u2212<\/jats:sup> exchange mechanism by physical covering\/uncovering of CCs by adjacent pavement cells. \u00a9 1992 Wiley\u2010Liss, Inc.<\/jats:p>","DOI":"10.1002\/jez.1402630205","type":"journal-article","created":{"date-parts":[[2005,6,11]],"date-time":"2005-06-11T00:26:16Z","timestamp":1118449576000},"page":"143-159","source":"Crossref","is-referenced-by-count":158,"title":["Mechanisms of ion and acid\u2010base regulation at the gills of freshwater fish"],"prefix":"10.1002","volume":"263","author":[{"given":"G. G.","family":"Goss","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S. F.","family":"Perry","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"C. M.","family":"Wood","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"P.","family":"Laurent","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2005,5,9]]},"reference":[{"key":"e_1_2_1_2_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.142.1.155"},{"key":"e_1_2_1_3_1","series-title":"Comp. Physiol.","first-page":"1","volume-title":"Animal Nutrition and Transport Processes. 2. Transport, Respiration and Excretion: Comparative and Environmental Aspects","author":"Avella M.","year":"1990"},{"key":"e_1_2_1_4_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.1402410202"},{"key":"e_1_2_1_5_1","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 hemoglobin oxygen\u2010carrying capacity","volume":"123","author":"Boutilier R. G.","year":"1986","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_6_1","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1242\/jeb.64.3.711","article-title":"Branchial ion uptake in Arctic grayling: Resting values and the effects of acid\u2010base disturbance","volume":"64","author":"Cameron J. N.","year":"1976","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_7_1","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1242\/jeb.86.1.171","article-title":"Body fluid pools, kidney function, and acid\u2010base regulation in the freshwater catfish Ictalurus punctatus","volume":"86","author":"Cameron J. N.","year":"1980","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_8_1","doi-asserted-by":"publisher","DOI":"10.1086\/physzool.62.4.30157933"},{"key":"e_1_2_1_9_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.133.1.183"},{"key":"e_1_2_1_10_1","doi-asserted-by":"publisher","DOI":"10.1139\/z89-432"},{"key":"e_1_2_1_11_1","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1242\/jeb.63.3.587","article-title":"The branchial chloride pump in the goldfish Carassius auratus: Relationships between Cl\u2212 \/HCO3 \n                  \u2212 and Cl\u2212 \/Cl\u2212 exchanges and the effect of thiocyanate","volume":"63","author":"De Renzis G.","year":"1975","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_12_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.59.2.339"},{"key":"e_1_2_1_13_1","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1242\/jeb.141.1.441","article-title":"Adrenaline and branchial nerve stimulation inhibit 45Ca+ + influx into the gills of rainbow trout, Salmo gairdneri","volume":"141","author":"Donald J. A.","year":"1989","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_14_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00706593"},{"key":"e_1_2_1_15_1","first-page":"149","volume-title":"Gills. Society for Experimental Biology Seminar Series","author":"Evans D. H.","year":"1982"},{"key":"e_1_2_1_16_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.1402390104"},{"key":"e_1_2_1_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/0006-3002(57)90529-2"},{"key":"e_1_2_1_18_1","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1016\/S0021-9258(18)85922-1","article-title":"Carbonic acid and bicarbonate in urine","volume":"51","author":"Gamble J. L.","year":"1922","journal-title":"J. Biol. Chem."},{"key":"e_1_2_1_19_1","doi-asserted-by":"publisher","DOI":"10.1085\/jgp.47.6.1195"},{"key":"e_1_2_1_20_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1977.sp012089"},{"key":"e_1_2_1_21_1","first-page":"R260","article-title":"Ion exchanges through respiratory and chloride cells in freshwater\u2010and seawater\u2010adapted teleosteans","volume":"7","author":"Girard J. P.","year":"1980","journal-title":"Am. J. Physiol."},{"key":"e_1_2_1_22_1","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1242\/jeb.152.1.521","article-title":"Na+ and Cl\u2212 uptake kinetics, diffusive effluxes, and acidic equivalent fluxes across the gills of rainbow trout. II. Responses to environmental hyperoxia","volume":"152","author":"Goss G. G.","year":"1990","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_23_1","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1242\/jeb.152.1.549","article-title":"Na+ and Cl\u2212 uptake kinetics, diffusive effluxes, and acidic equivalent fluxes across the gills of rainbow trout. II. Responses to bicarbonate loading","volume":"152","author":"Goss G. G.","year":"1990","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_24_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00260755"},{"key":"e_1_2_1_25_1","doi-asserted-by":"crossref","unstructured":"Goss G. G. P. L.Laurent andS. F.Perry(1992)Evidence for a morphological component in acid\u2010base regulation during environmental hypercapnia in the brown bullhead (Ictalurus nebulosus). Cell Tissue Res. in press.","DOI":"10.1007\/BF00319161"},{"key":"e_1_2_1_26_1","first-page":"309","volume-title":"Acid\u2010Base Regulation in Animals","author":"Heisler N.","year":"1986"},{"key":"e_1_2_1_27_1","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1242\/jeb.85.1.99","article-title":"The role of physico\u2010chemical buffering and of bicarbonate transfer processes in intracel\u2010lular pH regulation in response to changes of temperature in the larger spotted dogfish (Scyliorhinus stellaris)","volume":"85","author":"Heisler N.","year":"1980","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_28_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1979.tb03380.x"},{"key":"e_1_2_1_29_1","first-page":"R984","article-title":"Basis for apparent saturation kinetics of Na+ influx in freshwater hyperregulators","volume":"254","author":"Kirschner L. B.","year":"1988","journal-title":"Am. J. Physiol."},{"key":"e_1_2_1_30_1","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1007\/BF00339641","article-title":"The active absorption of ions in some freshwater animals","volume":"25","author":"Krogh A.","year":"1938","journal-title":"Z. Verg. Physiol."},{"key":"e_1_2_1_31_1","volume-title":"Osmotic Regulation in Aquatic Animals","author":"Krogh A.","year":"1939"},{"key":"e_1_2_1_32_1","first-page":"R147","article-title":"Morphology of gill epithelia in fish","volume":"238","author":"Laurent P.","year":"1980","journal-title":"Am. J. Physiol."},{"key":"e_1_2_1_33_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF01740769"},{"key":"e_1_2_1_34_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00216356"},{"key":"e_1_2_1_35_1","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(88)90562-4"},{"key":"e_1_2_1_36_1","first-page":"1085","article-title":"Les \u00e9changes de sodium chez le poisson Carrassius auratus L. Action d'un inhibiteur de l'anhydrase carbonique","volume":"48","author":"Maetz J.","year":"1956","journal-title":"J. Physiol. (Paris)"},{"key":"e_1_2_1_37_1","first-page":"255","article-title":"Branchial sodium exchange and ammonia excretion in the goldfish Carassius auratus","volume":"58","author":"Maetz J.","year":"1972","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_38_1","doi-asserted-by":"publisher","DOI":"10.1085\/jgp.47.6.1209"},{"key":"e_1_2_1_39_1","first-page":"273","article-title":"Branchial and renal acid and ion fluxes in the rainbow trout, Salmo gairdneri","volume":"98","author":"McDonald D. G.","year":"1981","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_40_1","doi-asserted-by":"publisher","DOI":"10.1086\/physzool.59.3.30156103"},{"key":"e_1_2_1_41_1","doi-asserted-by":"publisher","DOI":"10.1139\/z88-396"},{"key":"e_1_2_1_42_1","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1242\/jeb.102.1.141","article-title":"The interaction of environmental calcium and low pH on the physiology of the rainbow trout, Salmo gairdneri. 2. Branchial ionoregulatory mechanisms","volume":"102","author":"McDonald D. G.","year":"1983","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_43_1","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1242\/jeb.147.1.471","article-title":"The role of \u03b2\u2010adrenoreceptors in the recovery from exhaustive exercise in freshwater adapted rainbow trout","volume":"147","author":"McDonald D. G.","year":"1989","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_44_1","doi-asserted-by":"publisher","DOI":"10.1139\/z89-428"},{"key":"e_1_2_1_45_1","first-page":"86","volume-title":"Methods of Enzymatic Analysis","author":"Michal G.","year":"1985"},{"key":"e_1_2_1_46_1","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1242\/jeb.123.1.93","article-title":"Tissue intracellular acid\u2010base status and the fate of lactate after exhaustive exercise in the rainbow trout","volume":"123","author":"Milligan C. L.","year":"1986","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_47_1","first-page":"335","article-title":"Mise en \u00e9vidence d'un \u00e9change Na + \/NH4\n                  + dans la branchie de la truite adapt\u00e9e \u00e0 l'eau de mer: Contr\u01d2le adr\u00e9nergique","volume":"286","author":"Payan J. P.","year":"1978","journal-title":"C.R. Acad. Sci. Paris Ser. D."},{"key":"e_1_2_1_48_1","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1242\/jeb.116.1.411","article-title":"Kinetics of branchial calcium uptake in the rainbow trout: Effects of acclimation to various external calcium levels","volume":"116","author":"Perry S. F.","year":"1985","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_49_1","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1242\/jeb.147.1.147","article-title":"Adaptational responses of rainbow trout to lowered external NaCl concentration: Contribution of the branchial chloride cell","volume":"147","author":"Perry S. F.","year":"1989","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_50_1","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1242\/jeb.92.1.289","article-title":"Branchial ion uptake and acid\u2010base regulation in the rainbow trout Salmo gairdneri","volume":"92","author":"Perry S. F.","year":"1981","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_51_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF02464406"},{"key":"e_1_2_1_52_1","doi-asserted-by":"publisher","DOI":"10.1139\/z87-142"},{"key":"e_1_2_1_53_1","doi-asserted-by":"publisher","DOI":"10.1139\/z87-143"},{"key":"e_1_2_1_54_1","doi-asserted-by":"publisher","DOI":"10.1016\/0034-5687(89)90122-9"},{"key":"e_1_2_1_55_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00694377"},{"key":"e_1_2_1_56_1","doi-asserted-by":"publisher","DOI":"10.1038\/318368a0"},{"key":"e_1_2_1_57_1","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1242\/jeb.36.1.126","article-title":"The absorption of sodium ions by the crayfish Astacus pallipes (Lereboullet). I. The effect of external and internal sodium concentration","volume":"36","author":"Shaw J.","year":"1959","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_58_1","first-page":"122","volume-title":"Proton Passage Across Cell Membranes","author":"Steinmetz P. R.","year":"1987"},{"key":"e_1_2_1_59_1","doi-asserted-by":"publisher","DOI":"10.1016\/0034-5687(78)90079-8"},{"key":"e_1_2_1_60_1","doi-asserted-by":"publisher","DOI":"10.1139\/y83-207"},{"key":"e_1_2_1_61_1","first-page":"R395","article-title":"Correlation between release of catecholamines and disturbance of blood pH in rainbow trout (Salmo gairdneri) following exhaustive exercise","volume":"255","author":"Tang Y.","year":"1988","journal-title":"Am. J. Physiol."},{"key":"e_1_2_1_62_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-83130-0"},{"key":"e_1_2_1_63_1","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1242\/jeb.128.1.255","article-title":"The effects of prolonged epinephrine infusion on the physiology of the rainbow trout, Salmo gairdneri. III: Branchial solute fluxes","volume":"128","author":"Vermette M. G.","year":"1988","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_64_1","doi-asserted-by":"publisher","DOI":"10.1016\/0034-5687(84)90020-3"},{"key":"e_1_2_1_65_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.136.1.461"},{"key":"e_1_2_1_66_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00694238"},{"key":"e_1_2_1_67_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.1402050214"},{"key":"e_1_2_1_68_1","doi-asserted-by":"publisher","DOI":"10.1016\/0034-5687(80)90125-5"},{"key":"e_1_2_1_69_1","series-title":"Comp. Physiol.","first-page":"119","volume-title":"Animal Nutrition and Transport Processes. 2. Transport, Respiration and Excretion: Comparative and Environmental Aspects","author":"Wood C. M.","year":"1990"},{"key":"e_1_2_1_70_1","doi-asserted-by":"publisher","DOI":"10.1016\/0034-5687(84)90021-5"},{"key":"e_1_2_1_71_1","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1242\/jeb.126.1.499","article-title":"Downstream pH changes in water flowing over the gills of rainbow trout","volume":"126","author":"Wright P.","year":"1986","journal-title":"J. Exp. Biol."},{"key":"e_1_2_1_72_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00693000"}],"container-title":["Journal of Experimental Zoology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Fjez.1402630205","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jez.1402630205","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,10,23]],"date-time":"2023-10-23T13:03:33Z","timestamp":1698066213000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/jez.1402630205"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[1992,8,15]]},"references-count":71,"journal-issue":{"issue":"2","published-print":{"date-parts":[[1992,8,15]]}},"alternative-id":["10.1002\/jez.1402630205"],"URL":"https:\/\/doi.org\/10.1002\/jez.1402630205","archive":["Portico"],"relation":{},"ISSN":["0022-104X","1097-010X"],"issn-type":[{"value":"0022-104X","type":"print"},{"value":"1097-010X","type":"electronic"}],"subject":[],"published":{"date-parts":[[1992,8,15]]}}}