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Minimal Dissolved Oxygen Requirements of Aquatic Life With Minimal DissolvedOxygen Requirementsof Aquatic Life with Emphasis on Canadian Species:a Review JoHN C. D,q.vrs Department of the Environmen'r;,::;';;:;:#!Y;:';'ifii'i;Pacific EnvironmentInstitute' Contents 2295 Preface Respiratorydependence in invertebrates 2296 Abstract Oxygen requirementsof some freshwater 2296 R6sum6 in verte brates 2291 lntroduction,Rationale, and Objectives Oxygen requirementsof some intertidal 2297 PhysicalConsiderations marineinvertebrates 2299 Natural O, Regimesin the Aquatic Environ- Oxygenrequirements of benthicmarine or- ment ganismsand communities Oxygen tolerances and requirements of marineplanktonic and pelagic or- 2301 OxygenRequirements of Fish ganisms Bloodoxygen dissociation curves Ventilation, circulation,and respiratoryde- 2318 Summaryof OxygenRequirements of pendence rA.quatic Invertebrates Energeticsand low oxygen 2320 Developmentof Oxygen Criteria for Fish Behavioral responsesto hypoxia Populations Growthin reducedoxygen StatisticalTreatment of Data for Devia- Acclimationto low oxygen tion of Criteria Interactionof low oxygenand toxic agents 2325 Guidelines for Low Oxygen Tolerance of Eggand larval development in low oxygen AquaticInvertebrates 2326 Relationshipof ProposedCriteria to Other For personal use only. 2310 Summaryof OxygenRequirements of Fish Documents 2327 Suggestionsfor Implementationof Criteria 2321 Acknowledgments 2311 Oxygen Requirementsof Aquatic Inverte- 2127Glossary of Terms brates 2328 List of Symbols Aerobicand anaerobic metabolism 2328 References Preface Originally, some of the material in this report was prepared for the National Research Council of Canada Environmental Secretariat under the title "Waterborne Dissolved Oxygen Requirements and Criteria With Particular Emphasis on the Canadian Environ- ment." That document was published n 1975 as part of N.R.C.'s special publication series under the auspicesof the Associate Committee on Scientific Criteria for Environ- mental Quality. The emphasis in the N.R.C. document was on development of oxygen criteria which were defined as cause and efiect interrelations based on a scientific evaluation of environmental harm caused by low oxygen in water. The content of tle N.R.C. report was, of necessity, limited to matters considered the mandate of the N.R.C. environmental secretariat. This report, although similar in many ways to the N.R.C. report, is intended to serve more as a review document to persons interested in oxygen requirements of Canadian aquatic species. Where criteria are discussed, suggestions are made as to how criteria J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by UNIV OF WASHINGTON LIBRARIES on 07/11/18 might be implemented. It must be stressedthat the suggestionsherein are the author's opinions and are not the opinions of N.R.C. whose mandate does not include recom- mendations on criteria implementation. Readers should be very careful to distinguish between the content and positions taken when making reference to these two reports. Printed in Canada (13915) Imprim6 au Canada (J3915) 2295 2296 J. FISH. RES. BOARD CAN.' VOL. 32(t2)' 1975 Abstract Drvrs, J. C. 1975.Minimal dissolvedoxygen requirementsof aquaticlife with emphasison Canadianspecies: a review.J. Fish.Res. Board Can.32:2295-2312. This articlereviews the sensitivity,responses, response thresholds, and minimumoxygen requirementsof marineand freshwaterorganisms with strongemphasis on Canadianspecies. Th-eanalysis attempts to definelow dissolvedoxygen thresholds which produce some physiologi- cal, behavioral,or otherresponse in differentspecies. Oxygenavailability is discussedwith referenceto seasonal,geographical, or spatialvariation in dissolvedoxygen. Factors affectingavailability of dissolvedoxygen include atmospheric exchange,mixing ofwater masses,upwelling, respiration, photosynthesis, ice cover,and physi- cal factorssuch is temperatureand salinity.Dissolved oxygen terminology is summarizedand tablesare includedfor both fresh and saltwaterO, solubility at different temperatures. IncipientO, responsethresholds are used in a statisticalanalysis to developoxygen criteria for safeguardingvarious groups of freshwaterand marine fish. These include mixed freshwater fish populationiincluding or excludingsalmonids, freshwater salmonid populations, salmonid larvie or maturesalmonid eggs, marine anadromous and nonanadromousspecies. Criteria are basedon threshold oxygen levels which influencefish behavior, blood O, saturation, metabolic rate, swimmingability, viabilityand normal development of eggsand larvae, growth, circulatory dynamics,veniilation, gaseous exchange, and sensitivityto toxic stresses.The criteriaprovide three levels of protection for eachfish group and are expressedas percentageoxygen saturatlon for a rangeofseasonal temperature maxima. Oxygentolerances and responsesofaquatic invertebratesto low oxygenare reviewedfor freshwaterand marine species according to habitat.No inverlebratecriteria are proposed owing to the capacityfor many invertebratespecies to adopt anaerobicmetabolism during low Ot stress. It is suggestedthat the criteriaproposed for fish specieswill providea reasonablesafeguard to_ most ini-ertebratespecies. It appearslikely, however,that a changein oxygenregime to one of increasedO, scarcitywill probablyinfluence invertebrate community structure. It is suggestedthat iriteria for protectionof aquaticlife be implementedby groups of experienced-individuals.The group shouldconsider the naturaloxygen regimefor a specific waler body and its natural variability, the aquatic life therein and its value, importance,relative O, sensitivity,and the possibilityof interactionswith toxicants and other factors that may compoundthe stressproduced by low O, on aquaticlife. Each water body and its aquaticlife shouldbe consideredas a uniquesituation and criteria application should not encompassdiverse areas,habitats, or biologicalassociations as if they wereidentical. For personal use only. R6sum6 Davrs, J. C. 1975.Minimal dissolvedoxygen requirementsof aquaticlife with emphasison Canadianspecies: a review.J. Fish.Res. Board Can. 3;2:2295-2332' Danscet article,I'auteur passe en revuela sensibilit6,les r6ponses, les seuilsde r6ponseet les exigencesminimales en oxygEned'organismes marins et dulgaquicoles,avec accentpar- ticuliei sur les espdcescanadiennes. Il y d6finit les seuilsminimums d'oxygbne dissous qui d6clenchentdes r6ponses physiologiques, de comporlementou autreschez diff6rentes espdces' I1analyse |a diiponibiiit6d'oxygdne en relation avec la variationg6ographique ou spatialede I'oxygbne.Les factiurs affectantla disponibilit6de I'oxygbnedissous sont, entre autres,les 6changesavec I'atmosphbre,le m6langedes massesd'eau, la remont6ed'eaux profondes,la respirition, la photosynthdse,la couverturede glace et des facteurs physiquestels que la temp6ratureet la salinit6.Il donneun 16sum6de la terminologiede I'oxygdne dissous et destables de solubilit6 de I'oxygbne, tant en eau douce qu'en eau sal6e,ir differentestemp€ratures. Il utiliseles seuils de d6butde r6ponse)r I'oxygbne dans une analyse visant ir mettreau point des critdresde sauvegardedes divers gloupesde poissonsde mer et d'eau douce.Ces groupes comprennentdes populationsmixtes de poissonsd'eau douce, incluant ou excluant les salmonid6s,des populationsde salmonid6sd'eau douce, des larves ou des ceufsmfirs de salmonid6s,et desespbces marines anadromes et nonanadromes. Ces critbres sont fond6s sur les niveaux de seuil d'oxygdne affectant le comportementdes poissons,la saturationdu sangen O, , le taux m6tabolique,I'aptitude i la nage,la survivanceet le ddveloppementnormal des ceufset deslarves, la croissance,la dynamiquede la circulation,les 6changes gazeux et la sensibilit6aux stresstoxiques. Les critbresfournissent trois paliers de protectionpour chaquegroupe de poissonsetiont exprim6sen pourcentagede saturationd'oxygbne sur une gamme de maximums J. Fish. Res. Bd. Can. Downloaded from www.nrcresearchpress.com by UNIV OF WASHINGTON LIBRARIES on 07/11/18 saisonniersde temp6rature. L'auteur passLen revuela tol6ranceir I'oxygbneet lesr6ponses d'invert6br6s aquatiques ir de bas niveaux d'oxygbne, pour des espbcesdulEaquicoles et marines, selon l'habitat' Il ne proposeaucun critdre pour les invert6br6sir causede I'aptitude de plusieursespbces ir adopterun m6tabolismean6robie au moment d'un stresscaus6 par un basniveau d'Or. Les critEressugg6r6s pour lesespdces de poissonfourniraient une sauvegarderaisonnable pour la plupartdes espbces DAVIS:REVIEW OF OXYGENREQUIREMENTS oF AQUATICLIFE 2297 d'invert6br6s.Il est probable,cependant, qu'un changementde r6gimed'O, en directiond'une plusgrande raret6 d'Or, affectela strxcturedes communaut6s d'invert6br6s. L'auteur suggEreque les critEresde protectionde la vie aquatiquesoient appliqu6s par des groupesd'individus exp6riment6s. Ces groupes devraient 6tudier le r6gimenaturel d'O, d'un plan d'eau particulier et sa variabilit6 naturelle, la sensibilit6relative ir I'O2 et la possibilit6 d'interactionsavec les toxiques et autresfacteurs qui peuvents'ajouter au stress caus6 par un bas niveaud'O, sur la vie aquatique.Chaque plan d'eau et la vie aquatiquequ'il contientdevraient Otreconsid6r6s comme situation unique, et I'applicationdes critEres ne devraitpas englober des r6gions,des associations biologiques et deshabitats diff6rents comme s'ils 6taientidentiques. ReceivedJuly 21, 1975 Regule 2l I ,tlllet197 5 AcceptedJuly 30,1975 Accept6le 30juillet 1975 Introduction, Rationale, and Obiectives be exhibited by the test species,and
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