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A Review of Dissolved Gas Supersaturation Literature Don E This article was downloaded by: [US Fish & Wildlife Service] AR-1256 On: 02 February 2012, At: 07:25 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Transactions of the American Fisheries Society Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/utaf20 A Review of Dissolved Gas Supersaturation Literature Don E. Weitkamp a & Max Katz b a Parametrix, Incorporated, 13020 Northup Way, Suite 8, Bellevue, Washington, 98005, USA b Environmental Information Services Incorporated, 3455 72nd Place Southeast, Washington, 98040, USA Available online: 09 Jan 2011 To cite this article: Don E. Weitkamp & Max Katz (1980): A Review of Dissolved Gas Supersaturation Literature, Transactions of the American Fisheries Society, 109:6, 659-702 To link to this article: http://dx.doi.org/10.1577/1548-8659(1980)109<659:ARODGS>2.0.CO;2 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and­ conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Transactionsof the American Fisheries Society 109:659-702 @ Copyrightby the AmericanFisheries Society 1980 A Review of Dissolved Gas Supersaturation Literature DoN E. WEITK^Me Parametrix,Incorporated, 13020 NorthupWay, Suite 8 Bellevue,Washington 98005 MAX KATZ EnvironmentalInfo•ation ServicesIncorporated, 3455 72ndPlace Southeast MercerIsland, Washington98040 Abstract Dissolvedgas supersaturation is a conditionthat resultsfrown natural and human-causedpro- cesses.Supersaturation can result in gas bubble diseasewhich has been describedin a wide varietyof fishesand invertebrates.In recentyears dissolved gas supersaturation resulting from dams and thermal dischargeshas produced mortalitiesof fish in severalcases. This review discussesmost of the available literature dealing with dissolvedgas supersaturationand the recordedcases of gasbubble disease. Gas bubble disease is a condition that affects conditions. Much remains to be learned about aquatic animals residing in fresh or marine the physiologicalaspects of gasbubble disease. watersthat are supersaturatedwith atmospher- In order to understandgas bubble disease ic gases.Supersaturation, and the gas bubble and its cause,it is necessaryto be familiar with diseasethat may result in aquaticorganisms, the physicallaws governing dissolved gases and are not recent discoveriesnor are they only the factors that determine the level of super- causedby human activities.However, only in saturation.Boyer (1974), Woelke et al. (1974), recent years has supersaturation become a and Harvey (1975) discussedthe solubilitiesof problemof sufficientmagnitude to draw wide- dissolvedgases in water as they relate to gas spreadattention and concern. bubble disease. The majority of research dealing with dis- Harvey (1975) provided an excellentdiscus- solvedgas supersaturation has been stimulated sionof this subjectfor thosenot familiar with by a problem of considerablemagnitude that the physicallaws describingthe solubilitiesof wasobserved in the ColumbiaRiver systembe- gasesin a liquid. The solubilityof atmospheric ginning in the 1960's. More recently, interest gasesin water is determinedby the water'sdis- hasbeen further stimulatedby the discoveryof solved solids content, characteristics of the var- deleteriouseffects of supersaturationresulting ious gases,the total pressure, and the water from thermal effluents. temperature. Although total dissolvedsolids This reviewis an attemptto providea greater canaffect solubility, this is not a significantvari- Downloaded by [US Fish & Wildlife Service] at 07:25 02 February 2012 disseminationof the availableexisting knowl- able in most fresh waters but must be consid- edge regarding dissolvedgas supersaturation ered as a significantvariable in marine waters. and the resultinggas bubbledisease. The re- The atmosphericgases of importanceare ni- view discussesthe causesof supersaturation, trogen, oxygen, and argon. These gasesare the organismsaffected by supersaturation,fac- present in air at partial pressuresof approxi- tors affecting susceptibilityof aquatic organ- mately78% nitrogen,21% oxygen,and 1% ar- ismsto gas bubbledisease, and variousother gon. Nitrogen and argon are normallyconsid- related topics.The k•owledge of this subjectis ered together becauseboth are biologically considerableas evidenced by the length of this inert gaseswhile oxygenis a biologicallyactive review. Many important questionsremain to be gaso answered.This is particularlytrue regarding The solubilityof each gas is determinedby the applicationof laboratoryresults to condi- the massof the individual gas and its partial tionsfaced by aquaticorganisms under natural pressurein the atmosphere.Oxygen (21%) has 659 660 WEITKAMP AND KATZ onlyone-fourth the partialpressure of nitrogen marine fishes.Bubbles frequently occurred be- (79%) in the atmosphere,but is twiceas soluble hind the cornea and in the loose connective tis- asnitrogen. Therefore, in water,oxygen (35%) suesof the eye, producinga severeexophthal- is one-halfas plentiful as nitrogen (65%) (Har- mia or "pop-eye" condition. Bubbles were vey 1975). found lessfrequently in the gills,lining of the The major environmentalfactors that affect mouth, or along the lateral line of exposedfish. solubilityare pressureand temperature.Ac- Thesebubbles gradually increased in sizeas the cordingto Henry'sLaw, the massof a gasdis- length of exposure to supersaturation in- solvedin a liquid at a constanttemperature is creased.Fish with these signsalso showed loss proportionalto the pressureexerted on the sol- of equilibrium.The time to death varied from vent. Thus, as the pressureon a givenvolume severalhours to severalweeks following the ap- of water increases,the capacityof that volume pearanceof detectablebubbles. of water to hold dissolvedgas also increases. Marsh and Gorham (1905) further described Pressureis increasedin water by hydrostatic internal signsand lesionsof the disease.Free head. Hydrostaticpressure increases rapidly gas bubbles(or emboli) within blood vessels with depth, greatlyincreasing the capacityof were observed. The amount varied from a few deeperwater to hold dissolvedgas as compared scatteredbubbles to complete occlusionand to shallow water. distention of the vessels. The walls of the au- The capacityof water to hold dissolvedgas ricle and ventriclewere often emphysematous. is inverselyrelated to temperature.As the tem- In somefish, the auricle wasfilled with gaseven perature of a volume of water increases,the though it continuedto beat. The main vessels volumeof dissolvedgas it will hold at equilib- of the gills containedgas bubbles.Gas in the rium decreases.Thus, increasingwater tem- gill filamentswas described as the mostconstant peratureswill producesupersaturation in water and significantlesion of gas bubble disease. that is initially saturated. Death of the fish was attributed to stasis of the This brief reviewof the factorsaffecting sol- blood causedby emboli. ubilityonly discussesa few of the majorvari- In addition to observations of fish, Gorham ables. Those interested in a more detailed re- (1901) also reported signs of the diseasein view of the subject should refer to Harvey squid, bivalve mollusks, scallops, hydroids, (1975) or to textsdescribing the gaslaws in de- squid egg-sacs,and green algae. No detailed tail. discussionof the diseasein theseorganisms was History of Gas BubbleDisease given by Gorham. Gorham(1901) experimentallyproduced gas Early Observations bubble disease in fish held in closed containers There are severalnineteenth century records by reducingthe pressure.He alsowas able to of what appearsto have been gasbubble dis- causethe signsto disappearby subjectingfish ease.Hoppe-Seyler (1857), Bert (1873), and to a pressurecomparable to that exerted by a Regnard(1884) recorded external signs in fish water depth of 4.9 m. He concludedthat gas that apparentlyrepresent gas bubbledisease. bubbledisease was caused by a reductionin the The first completedescription of gas bubble pressureto whichfish normally were subjected. Downloaded by [US Fish & Wildlife Service] at 07:25 02 February 2012 diseaseand its causewas provided in a seriesof Marsh and Gorham (1905) later corrected this papersresulting from an air supersaturationmistaken conclusion. problemat the United StatesBureau of Fish- Gorham (1901) reported that whereasfish in eries station at Woods Hole, Massachusetts shallowaquaria developedgas bubble disease, (Gorham 1898, 1901;Marsh 1903, 1910;Marsh fishheld in ponds24 m deepwith waterfrom and Gorham 1905). This seriesof initial papers the same source remained free of it. This was dealtwith mortality,signs, and experimentsto the first indicationof the
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