AVOIDMCE ItEACTIONS OF SOME SAUMONI1) AD CENTRARCkI ID FISHES TO LOV CONCENTRATIONS OF DISSOLVED OXYGEN CECIL MARION dITMORE #1 A THESIS submitted to OREGON STATE COLLEGE in partial fulfillment of the requirements for the degree of MASTER OF SC IENCE June 1957 APk'ROVED: Redacted for Privacy Assistant Professor of Fish and Game Management In Charge of Major Redacted for Privacy Head of Lepartxilent of Fiah and Game Management Redacted for Privacy Chairman of h raauate Committee Redacted for Privacy Dean of Graduate School Date thesis is preonted Typed by Lenora Bond i,, i97 AC Ki' OV LDGid;iJT My appreciation is extended to Assistant Professor Charles E. Warren for making it possible for me to conduct this study. The experimental equipment was modified for oxygen avoidance studies bj Assistant Profes8or Charles E. Warren and Dr. Peter Doudoroff of the U. S. Public Health Service. Their encouragement, patience, suggestions, and construe- tive critic lam during the experimental phase of the study and during the preparation of the thesis was invaluable. The inspiration and encouragement offered by Professor H. H. Dimick is sincerely appreciated. The Investigator is indebted to Dr. Max Katz of the U, S. Public nealth Service for lila assistance in recording data and study. for helpful suggestions during the progress of the Thanks are also given to fellow research assistants John Fryer, John McCorrnack, Jack Foster, and Dean Shumway for their assistance. To Orville Greer, Superintendent of the Oregon Fish Commission "MacKenzie River Salmon Hatchery" and to Paul Vroman of the Oregon Game Commission "Alsea Trout Hatchery", gratitude is extended for providing chinook and coho salmon used in the experiments. Without the assistance and inspiration of my wife, this study would have proven more difficult. In addition to all of these, acknowleagment is given for the leading of my Lord and Savior, Jesue Christ. / / LIST OF TABL.: Page Table 1 2 3 Mean Oxygen Avoidance Indices for 15- minutePeriods......... .a.a. 17 Avoidance by Chinook Salmon during June and July, 1956, of Water having Various Concentrations of Dissolved Oxygen . . . . . 19 Avoidance by Chinook Salmon during September to November, 1956, of Water having Various Concentrations of Dissolved . . . . . . . . . . Oxygen . . 20 a 22 . a 25 . a 27 Avoidance by targemouth Black Bass thring August ar. September, 1956, of Water having Various Concentrations of Dissolved Oxygen . 28 . 4 . . . . . . The Effect of Temperature on the Avoidance by Chinook Salmon of Water havIng VLrious Concentrations of Dissolved Oxygen . 5 . . a . a 8 a . and a Avoidance by Coho Saliion during July and October, 1956, of Water having Various Concentrations of Dissolved Oxygen . 7 . Avoidance by Chinook Salmon during June July, 1956, of Water having Various Concentrations of Dissolved Oxygen (Channels A arid D only) 6 . a Avoidance by Bluegill Sunfish during August and September, 1956, of Water having Various Concentrations of Dissolved Oxygen . . . . . 30 Appendix A B C Dissolved Oxygen Concentrations at 12 inches and 3 inches Inside Channels and 3 inches Outside Channels for Channels A and B Dissolved Oxygen Concentrations at 12 inches and 3 inches Inside Channels and 3 inches Outside Channels for Channels C and D 48 a a a . a a 55 . 62 Number of Entries into Channels and Number of Fish Counted at 60-second Intervals . . TABLE OF CONTENTS Page a INTRODuCTIONa . a e a a a a a a 1 a 3 NATERIALS iiFTifJD ANJ) a a a a a a a a a Species, Source, Size and Care of Fish Apparatus . . . . . . . . . . . Experimental ivethod8 Computation of Avoidance Indices . RbULTS . . . . . . . . . ChiriookSalrnon... Coho Sa1ion . Largemouth Black Bass . Bluegill DIbCJSSIL)U . a . . a a . a a . . a a 5 . 5 . . a a a . . . . 7 . . . . . . 10 12 . . . . , . 15 ..a 18 . . . . . a . . a a ,a . . . . . . . . . . . . . . . . . a . . . . . . . . . . . . a a a . a a a a a a a a . a a a a a . Species Differences . Pos1b1e Factors Influencing Avoidance Mechanism of Avoidance . RapidiyLethalLevels a. a.a..... Ecolobical Sinificance . SJMiiARY . . . . . . . . . . . . . , . . a a a a . . . . . a . . . . . . . . . . . . a a a a a . . . a 26 26 29 31 3]. 33 33 36 38 40 BIBLIOGRAPHY .................. 44 APPENDIX 47 , . . . . . . . . . . . a a . a LIST OF FIGURES Pae Figure Apparatus used for avoidance studies . . . 8 AVOIDAiCE ELAUT iU1S uF SU t} iA IL AND C EiNTRkRC HID FIShES TO LOV CONCEThATIONS OF DISSOLVLD OXYGEN INTRODUCT I ON This thesis presents experimental data and. observa- tions on the avoidance reactions of some juvenile, fresh-water fishes to four concentrations of dissolved oxygen. The experiments wore performed at the Oregon State College Fisheries Research Laboratory at Corvallis, Oregon, June to November of 1956. This study was Oregon State College-Public part of the Health Service cooperative research program upon the effects of water pollution on aquatic biology. The primary purpose of this laboratory investigation was to determine whether fish could detect and avoid water of low dissolved oxygen concentrations. The detection and avoidance by fish of lethal dissolved oxygen concentrations could have survival value. On the other hand, detection and avoidance of concentrations of dissolved oxygen normally suitable for a species of fish could mean the 1088 of habitat for a population of that species. The contamination of many of our inland waters with various Industrial, agricultural, and domestic wastes has often rendered these waters unsuitable for desirable fish species. Depletion of dissolved oxygen in water, due to bacterial decomposition of putresciblo organic material and the respiration of various aquatic organisms, is often the 2 cause of sudden mortality of fishes. Information on the oxygen requirements of fresh-water fishes and their reactions in dissolved oxygen Is consequently of considereble Importance. Positive results in avoidance tests, such as these reported here, should not be Interpreted as Indicating that fish will avoid low concentrations of dissolved oxygen in. nature. Laboratory studies are convenient arid ay be Informative, but laboratory conditions are not entirely comparable to conditions in natural waters. In the labora- tory, fishes are necessarily confined In a small 8pace, and the oxygen gradients used are more pronounced than those usually found in nature. their migratory and. to be preBent under Normal responses of the fish to other tendencies can hardly be expected laboratory conditions. This paper bives a review of the few pertinent papers on the avoidance of fish of water havin different dissolved oxygen concentrations, a description of the apparatus used, and an account of experimental results. It also presents discussions of the differences in responses of different species, the possible factors influencin avoidance, the theories of the mechanism of avoidance, and the relation between the minimum tolerable concentrations of dissolved oxygen and the concentrations which were avoided by fish. 3 LITERATURE RVIEV Vhi1e numerous authors have studied the reactions of fish to various environmental conditions, there appear to have been comparatively few studies of the degree to which fish can detect arid avoid waters having low concentrations of dissolved oxygen. Shelford and Allee (20, p. 207) were the first to aply the use of a gradient tank to this Their studies were conducted using a gradient tank with water of different dissolved oxygen concentrations introduced at opposite ends and with a drain in the problem. middle. All the oxygen was renoved from the incoming water by boiling, and. then pure oxygen was added to provide the desired concentration, The results of their experiments ith 16 species of fresh-water fish at tifferent dissolved oxygen concentrations merely indicated that fish show a general tendency to avoid water having a low dissolved oxygen concentration. Several years later, in experimental work with similar gradient tanks, Sholford and. Powers (21, p. 334) observed that Pacific herring, Clupea pallassi, selected the water having the higher dissolved oxygen concentration. J, R. E. Jones (14, p. 403) has used a cylindrical glass tube with waters of different dissolved oxygen concentrations entering at each draining at the center. end of the tube and This differed from previous 4 appar.tus used in avoidance studies by vroduoin a sharp bounthrry between the waters hving different oxygen eon- The desired dieo1ved oxygen concentrations centrations. were obtained by adding boi1ei portions to normal tap water, ater in the ;roper ireOnly a. Single fish uood at a time in the experimental tube. at 0. with the stiok1ebac r Jones' results showed no hesitation of the fish on entering water of low dissolved oxygen coneentrations, bat the fish wore stimu.Latod to swim more rapidly in At this water. ; 0. and with oxygen concentrations below mg./i., the rosionse was very orowot, the fish finding their way out of the water of low oxygen concentration. lie obtained si ilar results with the minnow, Phoxinus, and with brown trout fry. ]. R. . JOneS (ii, . 110; 12, u. 2; nd 13, p. 6l) used similar ap )orstus in stndyin, the avoidare by fish of toxic so.ution, and other workers have used different types of ap aratus fo r experiment a involving v oidance re- actions of fishes to factors other than dissolved oxygen, Doudoroff (3, p. 495) devised a coxnpartmented gradient tank ith numerous inlets and outlets for studying reactions to temperature. Ohidester (o, p. 7) had waters of different salinity flowing down two troughs into a third one so thot the fish swimming up the trough could choose between the salinities, Natural conditions probably have been 5 approximated most nearly by Collins (6, p. 379), who constructed a divided trough in a stream to study factors influencing the migration of anadromous alewives and glut herring, hrett (2, p. 307) used a vertical gradient tank in teats of the responses of fish to vertical temperature gradients. Jones et al. (ID, p. 1403-1414) used an avoidance tank having four channels at one end for studying the avoidance reactions of yearling chinook and coho salmon to pulp mill effluents The inlets and drains to the four channels of the avoidance tank were regulated to give a rather sharp dem&rcatlon between altered and unaltered water at the channel entrance. In this study, the chinook salmon exhibited higher avoidance then did the coho salmon. This appeared to indicate that possible differences exist in the sensitivity or behavior of the two species. The 4-channeled avoidance tank used by Jones et al. was modified for use in the experiments reported here. iETH0L) AND AIIATbRIALS Species, Source, Size and Care of Fish The experimental animals used for thIs investigatIon were juvenile fish of four freshwater species, nae1y, chinook salmon, Oncorhynchus tshawytscha (Walbaum), coho salmon, Oncorhynchus kisutch (Walbaum), largemouth black bass, Mioropterus salinoides (Lacepede), and the bluegill, Lepomis macrochirus Rafinesque. The chinook salmon were obtained periodically from the Oregon Fish Commission's MacKenzie River Salmon Hatchery and the coho salmon were obtained from the Ore&;on Game Commission's Alsea Trout Hatchery. The largemouth black bass and the bluegill were collected from several sloughs and ponds in the vicinity of Corvallis. The fish were kept in 250-gallon, wooden, cylindrical tanks having flowinb water. Water was supplied to both the holding tanks and the experimental tank for avoidance tests from a large, elevated storage tank which was filled with water pumped from Mary's river. Since the source of water was the same for the holding tanks as for the experimental tank, the temperatures for the holding tanks and the experimental tank did not differ appreciably. The fish were fed every morning. During June and July the food for the salmon consisted of ground fish and beef liver with mineral and vitamin supplements. Later ifl the year, dry commercial trout food in pellet form was used as food for salmon and bluegill. Since the bass would not feed on the pellets, small minnows and annelid worms were used. The fish were kept in the holding tanks for periods ranging from several days to two months before being used 7 in experiments. Apparatus The apparatus used in those avoidance studies, which is shown in Figure 1, consisted of a tank 12 inches high, 24 inches wide and 112 inches long. Four channels of equal width and 6 inches long, separated by painted, plate glass partitions were located at one end of the tank. channels were designated A, left to right. 1-3, These C, and D, proceeding from Each channel had an Independent, adjustable water input at the closed end and an independent, adjustable drain at the open end. to cirain across nearly the entire width of the channel at a inches above the tank bottom. level 2 both sides of each drain beneath Each drain was slotted prevented Oblique baffles on water from passinG the drains and helped the fish to find their way readily over the drain. The tank was enclosed In a separate compartment of the laboratory room to avoid unnecessary disturbance of the fish. One end of the compartment was provided with a polarized glass observation port from which the ments of the fish. investigator could observe the moveA two-tube, fixture was suspended from the 96-inch, fluorescent, light ceiling of the white-walled compartment, providing relatively uniform light throughout the tank. ½ RUBBER FLUORESCENT FIXTURE TUBING LIGHT ,,%' /2'/''_ /,_, ,/ / , / ,, / /,/,/,/ /,/ ' , ,1 II I I II ,f ,'1'/," HINGED DOOR } Jl'>;/' )!: / )/ //T7P' POLARIZED GLASS OBSERVATION PORT jCATCH PAN ,FOR DRAINS -T L____ 5c / / ' , : ONE PARTITIONS- /4O.D.PLASTI PIPE - / ,,f/ \ / CHANNELI GLASS .-i. RUBBER CORKS, FLOOR OF TANK I, DIA. GLASS TUBING RUBBER HOSE ____________I ' IIh 1T 1 FIBER jj ADJUSTABLE_CLM (REMOVABLE) 'WH 3-REQQ. OR S ES AVOIDANCE TROUGH GLASS PART4TION END VIEW OF AIN AND II BOARD [L,2X4 coMPARTMENT ENCLOSING AVOIDANCE TROUGH EXTERIOR TYPE PLYWOOD -_ DRAIN BAFFLE - ALUMINUM PARTITION FOR GLASS PARTITIONS EXTERIOR TYPE PLYWOOD FLOOR Figure 1. SLOT FOR DRAI SIDE VIEW OF DRAIN ND CHANNEL Apparatus used for avoidance studies. The dissolved oxygen concentration of the water supplied to two of the channels was controlled by bubblIng nitrogen through the water as it flowed downward in two :;lass columns filled with glass Rasohig rings. The water for the two other channels was aerated as it passed downward through additional columns. The glass columns were 60 inches long and 2 inches in diameter and were connected to the channel inlets with rubber tubing. The flow from each inlet was measured for one minute by attaching a rubber hose to the inlet and holding the outlet of the hose outside the tank at the level of the water In the tank. The flow from each of the four channel Inlets was adjusted to approxImately 1200 ml. per minute. Two inlets at the opposite end of the tank were each adjusted to have a water flow of approximately 600 ml. per minute. The outflow of each of the drains at the channel entrances was adjusted to approximately 1500 ml. per minute. Since the rate of the flow Into a channel was less than the rate of the flow leaving the drain, the difference being water from the tank beyond the channel, a rather sharp boundary between water in the channel and water beyond the channel existed. A painted screen was placed across the body of the tank 26 Inches from the drains in order to confine the fish and to encourage more entries by the fish into the channels. 10 Two lines or rmrkers were used for reference when recording the number of fish going into the channels. The first line was the drain at the entrance to the channel, the second line being a marker on the bottohi of the tank 10 inches from the opposite end of the channel. Experimental Methods Water of reduced dissolved oxygen concentration was introduced into two alternate channels of the tank, these being referred to as the experimental channels. The other two channels received aerated water and served as controls. To eliminate one possible source of bias, the pair of channels receiving the deoxygenatod water was changed every experiment b: changing the connections between the inlets and the glass columns. Oxygen determinations were all done by the Alsterberg (azide) modification of the Winkler method (1, p. 255), a microburette being used for titration. oxygen d.etormiriationw 60 ml. sam11e bottles. Water samples for siphoned slowly from the tank into Samples were taken 3 inches outside each channel and at points 3 inches and 12 inches inside each channel, halfway between the channel sides. The samples were all tahen about an inch below the surface of the water. The oxygen determinations were made just prior to and at the end of each 45-minute experiment. 11 The number of fish used in each experiment varied from 10 to 35, Large numbers of fish were needed when little The number of fish for each experiment movement occurred. was chosen so as to have as many entries as could be water The temperature of the nd reooraed. accurately counted was recorded at the time of placing of the fish in the tank. Observations were not made until the fish began to The time interval from the introduction move about freely. of fish to the beginning of observation averaged about 10 minutes, but varied from 3 to 30 minutes. the species, being 1onest It varied with for chinook salmon and shortest for coho salmon; and it varied also with the temperature and the time of clay. The observations were ordinarily recorded separately during three successive 15-minute periods. xperiment8 8 through 68 were done by two persons, the investigator calling out the observations and the other person recording. Following the 68th experiment, both the observing and the recording vero done by the investigator. The number of fish crossing the urain, the number crossing the second line, and the number present in a channel at 60-second intervals were recorded. Only the first crossing of the second line by any individual fish entering a channel was recorded. 12 some control oxperiiiients were performed in which unaltered water was introduced into all channels. These experiments were conducted for the purpose of observing how the fish behaved without differences in dissolved oxygen concentration in the channels. Inasmuch as the time available for completion of the experiments under COndItiOn8 of temperature suitable for the different species was limited, complete series of control experiments were performed only with largemouth black bass and wIth chinook salmon tested In the fall (September to November). Only three control tests could be performed with coho salmon, and. three were performed with bluegill. The control tests with coho salmon were performed much later than the reaction experiments with this species and may not be reliable. Computation of Avoidance Indices The measure of the degree of avoidance by fish of experimental channels employed in this study is referred to as the "avoidance index" or the "index of avoidance". value, when This or on numbers of fish crossing the second line, wa calculated by the formula: based on numbers of entries into channels Avoidance Index 100 x CM - A)/M where L represents the sum of the entries into all the channels divided by two, and A represents the sum of the 13 entries into the experiruontal channels. It should he noted that the formula relates the number of entries at the first line or the second line of the experimental channels to the corresponding entries at the first line or second line of the control channels. When the "avoidance inUex" was zero, equal numbers of entries into the experimental and the control channels were indicated. indextt A positive "avoidance 1ndiates that the number of entries into the experimental channels was lower than the number of entries Into the control channels. The reater the value of this index, the greater was the difference between control and negative index shows that experimental channel entries. the number of entries into the experimental channels exceeded the number of entries into the control channels. When based on t;ime spent b the fIsh in the channels, the "Index of avoidance" was cof;!puted by the same formula, but in this case i represents the sum of the numbers of fish counted in all channels at 60-second intervals divided b 2, and A is the sum of the number of fish counted at 60-second Intervals In the experimental channels only. Jones et al. (B, p. 1405), in their study of the avoidance reactIon of salmon to pulp mill effluents, used 100 x (E - A)/E, where E represented the number of fish In the control channels, and A represented the nurber of fish counted in the the formula: Per cent Avoidance 14 experimental cbsnn1s. This formula is based on the assumption that a fisn which is repelled by the tater in any channel turns baoa and. does not immediately enter another channel where the repellent stimulus is lacking. This assumption probab1. is not entireLy valid. Fish apeared to be repelled by the water of low dissolved oxygen content in such a manner tt some went back towards the open part of the tank and. others ere diverted into another channel. The formula u.sed. in the present study yields an index value which is lower than the avoidance percentage comuted by the formula of Jones et cii. hen the same data are used. This is so, since in the present study the mean of observed entries into the experimental and the control channels is taken to be the expected number of entries into experimental channels. yIhile under some circumstances it may be more nearly correct to consider the number of observed entries into the control channels as being the expected number of entries into the experimental channels in the absence of a repellent stimulus, the formula of Jones et a.].. has a considerable negative bias, which renders it unsuitable for the present study. For instance, with the formula of Jones et al. if 38 fish entered into the control channels and l30 fish entered into the experimental channels the £oowing result is obtained: 100 x (8 - 180)/38 -373.7ç. Iith the numbers of entries 15 the ame, the "Avoi.iance Index" of this oxperinient would be: 100 x (109 - 180)/109 -65.1. It is apparent that one very extreme negative value could cancel out several positive values. Indeed in a few of the experiments recorded here the application of the formula of Jones at al. would yield extreme negative values such as -300 or study b Jones at al. the salmon never ciexdontrated much "negative avoidance". Ina8-500%. Fortunately, in the much as the "ixciices of avoidance" are intended chiefly for comparative purposes, the formula used in the present study appears to be more satisfactory for general application than that used b Jones at al. RESU IJTS All four species of fish used in these experiments avoided water having some reduced level of dissolved OXOfl. The species differed in the amount and manner oi their avoidance as shown below. The effects of season and water temperature can be shown only for the chinook salmon. In all species, avoidance of a low dissolved oxygen concentration is less evident when the number of fish crossing the first line is considered than by the number of fish crossing the second line or the total the channels at 60-second intervals. numbers of fish in 16 The chinook salmon in the earlier experiments showed some indication of a change in avoidance to the low concentrations ot dissolved oxygen, particularly between With the first 15-minute and the second 15-minute period. the later chinook experiments little more change In avoidance was indicated. "indices of avoidance" for than a hint of a The means of the 15-minute periods are given in Table 1±or all species tested. Tables are given in the text for each species and include the date of the experiment, mean oxygen concentration, water temperature in degrees Centigrade, and millimeters. Calculated based on entries at the first line, of fish in the size "avoidance indices" at the second line, and the number of fish counted in the channels at 60-second intervals are given for each experiment. different "indices of The means of the avoidance" obtained in all tests at each dissolved oxygen concentration with each species are also given. Actual numbers of entries at the first line and second line and the numbers of fish present in the channels at 60-second intervals are given for all species in Appendix Table C. The dissolved oxygen concentrations of samples collected inside the channel at points 3 inches and 12 inches from the drain and at a point 3 inches outside each drain are given In Apendlx Table A and Appendix Table B. The TABLE 1 lEAN OXYGEN AVOLANGE 1 ND IC S FCR 15-MINUTE PhRIQDS Exps. Dis. 02 1st 15 1st line 2nd 15 3rd 15 1st 15 2nd line 2nd 15 3rd 15 1st 15 Time 2nd 15 3rd 15 mm. mlxi. mm. mlxi. mlxi. rain. mm. mm. mlxi. 44.6 35.7 30.0 - 6.5 27.1 20.5 76e5 50.9 75.5 75.0 21.3 - 1.0 67.1 29.8 7.8 1.4 16.6 82.5 45.4 42.8 - 3.1 32.4 18.1 4.8 -10.5 3.2 58.0 47.6 49.1 -10.4 75.4 45.2 2.1 - 5.7 8.4 95.6 77.5 - 4.4 76.1 40.0 4.3 -11.4 12.6 -l0.0 74.1 48.5 10.4 -12.8 3.2 90.4 80.0 8.4 - 2.5 57.8 42.2 1.8 1.9 56.0 57.3 22.8 - 1.3 24.1 17.8 2.3 2.1 5.2 96.2 89.0 60.0 - 3.6 72.4 37.5 17.7 -18.9 9.4 10.7 17.5 4.d 5.5 27.1 27.1 24.4 10.7 34.7 23.2 5.6 6.4 25.2 30.7 13.2 18.2 53.3 44.6 30.8 17.1 60.1 33.6 18.2 19.8 56.1 29.6 11.1 23.8 35.9 62.2 21.0 49.8 65.8 47.8 1.5 10.3 - 5.6 3.0 4.5 2.6 6.6 2.2 12-130 131-140 control - 4.1 16.7 6.4 2.9 - 3.2 2.0 27.7 11.1 5.6 - 6.2 10.2 394 .9 2.9 39.6 14.2 10.9 1.6 10.2 60.6 12.1 11.6 - 1.6 - 1.9 75.3 10.2 6.1 - 2.7 53.0 9.5 14.9 71.2 15.8 14.8 - 4.4 16.1 Bluegill 141-148 149-154 155-160 161-165 29.0 3.4 - 7.0 13.9 21.3 - 4.2 71.9 - 4.6 - 5.5 50.8 26.1 -13.6 11.0 49.7 - 3.0 - 6.5 - 6.4 72.3 23.2 8.9 - 3.7 74.2 mel. mg/i Chinook 1.5 1-8 3.0 9-16 4.5 17-20 21-27 6.0 28-35 1.5 36-43 3.0 4.5 44-51 6.0 52-60 61-68 control - .4 .8 ,9.6 57 .05 15.9 C oho 69-72 73-77 78-81 82-86 1.5 .0 4.5 6.0 bass 90-97 98-109 110-122 1.5 3.0 4.5 6.0 26.4 - 6. 4.0 7.6 - .O -13.8 - .9 .2 3.6 11.3 11.8 12.1 27.9 2.4 o2.5 47.8 21.2 .5 -16.6 - - 5.3 -12.3_ mean dissolved oxyen concentrations recorded in the text tables are the arithmetic means of the four initial and four final dissolved oxy.., en determinations for the samples collected inside the two experimental channels. Chinook Salmon The chinook salfion used in the June and July experi- ments avoided water with concentrations of 1.5, 3.0, and 4.5 mg./i. dissolved oxygen, as shown in Table 2. The chinook salmon of the September, October, and November experiments (Table 3) did not avoid water having 4.5 mg./i. of dissolved oxyon, and neither series avoided water having a dissolved oxygen concentration of 6.0 mg./l. "indices of the The of chinook salmon increased as the dissolved oxyen concentrations decreased. The two :roups of chinook salmon tested showed consistent seasonal differences. The mean "index of avoid- ance" based on entries at the first line with 1.5 mg./l. dissolved oxygen was 61.0 in the experiments performed in June and July (Table 2) and 27.8 in the experiments performed during September, October, and November (Table 3). Similar differences in the "iriex of avoidance" were demonstrated at the second line and with the "avoidance index" based on time in most experiments. This same relationship was apparent at 3.0 mg./l. and at 4.5 mg./l. 19 TABLE 2 AVIDACE BY CijI1OK SALiON DU RiNG JUNE AND JULY, 1956, OF VATER hAVING VARIOUS CONCENTRATIONS OF DISSOLV&i OXYGEN Exp. No. Temp. 0C. Size 16.5 7/23 1.8 1.6 1.4 1.4 2.0 1.5 1.6 1.5 63-78 63-78 63-78 63-78 73-92 73-92 73-92 75-95 6/19 6/19 6/21 6/21 7/18 7/19 7/24 7/24 6.1 2.9 3.0 2.9 .0 6.0 3.0 2.9 16.5 16.5 18.5 19 25.5 23.5 24.5 75-95 63-78 50-83 73-92 78-96 78-96 7/16 7/17 7/25 7/25 4.5 4.5 4.5 4.5 22.5 21.5 22.5 24.5 72 72-84 61-87 70-90 6,/20 6.0 6.3 5.7 6.3 6.1 6.1 6.0 16 70-80 70-80 Date Mean D.O. rng. 1 2 3 4 5 6 '7 8 Mean 9 10 11 12 13 14 15 16 Mean 17 18 19 20 Mean 21 22 23 24 25 26 27 Mean 6/22 6/22 6/25 6/27 7/19 7/20 '7/23 6/20 6/28 6/29 '7/14 7/16 7/26 /1. 17 16.5 18.5 25.5 23.5 22.5 25.5 22 16.5 19 18 18.5 19.5 21.5 mm, 75-95 '75-95 63 63 72 72 72 Avoidance Indices Line 1 Line 2 Time 75.1 54.3 41.3 66.8 66.4 53.0 68.1 63.2 61.0 9.3 52.7 30.2 63.8 46.7 68.1 26.4 76.2 46.7 91.6 85.0 65.9 94.9 94.3 96.1 93.2 100.0 90.1 77.1 87.8 79.8 91.3 7.9 86.4 96.8 98.8 89.5 15.1 80.4 31.5 85.3 74.8 93.8 62.6 97.5 67.6 45.0 72.5 56.3 69.7 83.5 25.6 26.2 26.5 36.5 38.9 44.9 33.3 38.4 10.8 53.1 32.6 22.6 32.7 35.2 22.1 -65.1 8.9 32.2 27.6 10.3 -50.5 - 5.2 -12.2 32 -13.7 2o.9 - 1.9 '79.1 7.8 95.4 71.9 -63.? -14.8 -16.2 -12.9 3o.2 43.2 -16.2 -38.4 31.2 28.8 - 4.7 - 1.9 20 TABLE 3 AVOIDANCE BI CHThu)K SALVk)N LURING SEPTEMBR, OCT03ER A1D NOVEMBER, 1956, OF WATER EAVING VARIOUS CJNCETRATI0NS OF DISSOLVED OXYGEN Mean D.O. mg./i. Exp. Date 28 29 30 31 32 33 34 35 9/29 9/29 10/1 10/2 11/11 li/li 11/16 11/16 1.7 1.7 1.5 1.5 1.9 1.5 1.6 1.6 10/3 10/4 10/5 10/6 11/5 11/7 11/7 11/8 3.0 3.1 3.0 3.0 3.0 2.8 3.0 2.8 10/6 10/9 10/9 10/11 10/29 11/2 11/3 11/3 4.6 4.6 4.5 4.5 4.5 4.6 4.5 4.5 Temp. 1? 17.5 18 15.5 10 10 8.5 8.5 Size Avoidance Indices Line 1 Line 2 Time 93 93 78-97 78-91 95-105 85-100 90-105 90-105 Mean 36 37 38 39 40 41 42 43 16.5 13.5 15.5 15.5 10.5 11.5 11.5 1]. 44 45 46 47 48 49 50 51 1'? 14.5 15 13 10 10 9 10 56 57 58 59 60 Mean 10/12 10/15 10/16 10/16 10/23 10/23 10/24 10/26 10/29 6.0 6.0 5.9 6.2 6.0 5.8 6.1 6.1 6.3 14 14 13.5 13.5 11 11 10 9 10 49.2 32.9 85.5 69.5 72.1 27.8 10.3 22 30.4 18.5 17.1 22.2 9.9 13.5 63.4 91-97 91-97 91-99 91-99 85-95 80-100 95-105 85-90 -15.8 -17.1 -15.6 - 3.1 24.5 62.1 Mean 52 53 54 55 843 91-97 91-97 91-97 91-97 85-90 90-105 90-105 90-105 Mean 91-99 91-99 91-99 85-100 80-90 80-100 90-110 90-110 90-110 -14 1.8 10.2 217.8 - .03 '74.5 64 67.1 60.3 oO.1 26 53.8 59.2 27.2 5'7.6 29 36.3 40.7 40.2 43.3 53.4 2'7.7 44 15.8 39.8 - 2 9.4 9.6 3.4 11.2 7.4 -40.1 - 5 - 6.2 44.9 2.8 12.3 -27.5 61.8 89.3 73.4 1714 62.4 42 80 93.4 81.6 26 37.6 19 30.7 31.8 12.6 31.9 33 27.8 - 6.9 - 1.2 3 8.2 50.2 2 .8 10.5 11.1 5.4 20.9 5.8 -46.4 - .8 - 8.7 4.3 .8 10 -32.1 - 5.1 -16.8 14.6 10.8 18.9 6.3 -57.4 2.3 - 2 .8 .8 - 7 -50.2 - 8.3 21 TABLE 3 (continued.) Exp. M an Date DO Teo. Size 00. imi. Avoidance Indices Line 1 £ine 2 Time ;T 1.8 8.8 61 62 63 64 6b 66 11/2 11/17 11/17 11/17 11/17 11/19 I.O 10 10 10 7.b 7.5 67 11/19 11/20 9.8 9.i 7.5 6.b 68 Mean 10 9 9 12.2 19.2 5.2 1.8 90-100 85-105 O-1O5 95-110 95-110 db-100 - 3.1 14.8 85-100 90-10 3 15 34.3 2b.i 2. - 2.9 - i.4 37.9 -14 12.3 .4 24.5 25.8 -1t3.2 .8 3.2 27.d -21.2 5.2 dissolved oxygen. Water temperature, abe, and size of fish may be factors contributing to the seasonal differences in avoidance. There was some evidence sugesting temperature to be an important factor influencing the avoidance by chinook salmon of some dissolved oxygen concentrations, but there was little evidence available on the effect of size and age of the fish. Temperature would be expected to tor influencing avoidance since it is in e an important fac- known that an increase metabolic rate results from an increase in The temperature effect was particularly temperature. noticeable in tests at 1.5 mg./l. and at 3.0 mg./1. of dissolved oxygen CTable 4). At other concentrations, differences due to tempera- ture were not demonstrated. Though differences may have 22 TAE rE 4 f1j T: : ViIAUC:F c ;mTuoK SALMON OF VATER HAVING VARIOUS 00NCEThATIONS OF DISSOLVED OXYGEN No. of xps. Appiox. D. 0. 1-4 5-8 4 4 9-12 13-16 4 1.5 1.5 6.0 3.0 Exp. o. 28-31 4 4 4 3 37 3. 40-43 4 1.5 1.5 3.0 3.1 3.0 32-35 36,38,39 Avoidance Ldices Temp. °C. Line 1 16.5-19 22.5-26 17 22 -19 -26 15.5-18 8.5-10 15.5-16.5 13.o -11.5 II Line 2 Time 5.4 34.4 o2.7 39.0 54.4 95.9 53.1 02.2 4.0 95.0 60.8 83.0 28.3 27.3 29.4 10.3 16.9 84.8 59.3 58.8 26.0 29.0 '74.0 60.3 57.1 30.1 36.2 been present at other concentrations, lack of temperature differences between experiments or low amounts of avoidance made a demonstration of the presence of temperature effects i:apossible. Data presented in Table 4 suest that at low concentrations of dissolved oxreri the "index of avoid- ance" increases with increasin' water temperatures. This may account for some of the apparently seasonal differences In avoidance, and it may also account for some of the variability in the avoidance determinations at particular concentrations durin; the same season. During the same season the "index cf avoluanee" based on time for experiment No. 37, at a temperature of 13.50 C. was 30.1 while the mean "Index of avoidance" for experiments No. 36, 38, and 39 at a temperature of 15.6° to 16.5° C. was 57.1, 23 as recorded. in able 4. Exoeriments No. 40 to 43 have "indices of avoidance" similar in magnitude to experiment No. 3? at about the same temDerature. Comparing the means of the avoidanoe indioes in Table 4, it is seen that little difference is evident at a particular dissolved oxygen concentration where the range of water teiperaturo is approximately the came. 3everal deaths of fish in low dissolved oxygen channels may have been due to increased metEbolio demand at teIDeratures from 23 to In addition to the influence on the avoidance of concentrations of low dissolved oxygen, high temperatures of the water appeared to reduce the activity of the chinook salmon in the tanks. 260 0. Schooling behavior of the chinook salmon was observed to a greater extent than usual in experiments No. 1 to 4, 9 to 12, and also experiments 21 and 23. The term schooling is used here to designate a tendency of the fish to swim in groups, the individuals of which are strongly influenced by each otherts movements. lhen a single fish moving into a channel at the same time that other fish were moving out of the channel turned and left the channel with the other fish, this fish is considered to have behaved in a schooling manner. This was observed in the experiments listed above. The combined effect of schooling and an apparent 24 preference for channels A and D by chinook sa1on seemed to affect the reactions in several ways. One observed effect was the turning of an entire group of fish away from a channel whiCh the group was about to eater, parently due to the hesitation or the turning at the entrance of the channel of the leading fish in the school. Another effect was the aeparatica of several of the £1SI from the rest of the group movin the channels. into a channel by the partitions between This splitting of groups of fish apparently resulted in an inordinately large number of entries into the low oxygen channel adjacent to the control channel located next to the edge of the tank. Consequently, the number of entries into this low-oxygen channel often exceeded the number of entries into the control channel which was removed from the edge of the tank. This effect is evident on examination of the data of experiments No. 1 to 4, 9 to 12, 21 and 22 in Appendix Table C. Due to the separation of the school of fish at the entrances to the channels in the experiments listed above, the use of only tue data for channels A and D appeared to give a better eotinito of the avois.ance than the the data from all four channels ise of Table $ shows that higher indices result from using data only from channels A arid. D than from using data from all four channels (Table 2), particularly in experiments where schooling was evident. 25 TABLE 5 AVOIDANCE4 BY CHINOOK SALMON DURING JUNE AND JULY, 1956, OF VATER HAVING VARIOUS CONCLNTRATIONS OF DISSOLVED OXYGEN (CHANNELS A AND B ONLY) Exp. No. 1 2 3 4 5 6 7 8 Date Mean D.0. 1.6 1.6 1.1 1.2 Size 00. mm. 16.5 17 16.5 18.5 25.5 23.5 22.5 25.5 63-78 63-78 63-78 63-78 73-92 73-92 73-92 75-95 83.7 72.7 59.2 87.5 87.7 70.1 91 56.5 76 96.2 98.5 75.4 98.7 93.2 96.3 100 100 94.8 79.8 94.1 94.3 93.8 93.1 13.8 63.1 26.8 53.8 78.6 73.1 74.2 81.3 92.8 55.9 15.7 85.6 32.8 98.4 73.4 95.2 53.8 98.7 69.2 55.2 49.6 59.4 21.8 46.5 6/22 6/22 6/25 6/2? 7/19 7/20 7/23 7/23 1.8 1.5 1.6 1.5 6/19 6/19 6/21 6/21 7/18 7/19 7/24 7/24 3.1 2.9 6.0 2.9 3.0 3.0 3.0 2.9 16.5 16.b 18.5 24.5 75-95 75-95 75-95 63-78 50-83 73-92 78-96 78-96 7/16 7/17 7/25 7/25 4.5 4.5 4.5 4.5 22.5 21.5 22.5 24.5 72 72-84 61-87 70-90 28.9 43.1 43.3 41.9 39.3 37.5 52.9 54.9 38.2 45.4 6/20 6/20 6/28 6/29 7/14 7/16 7/26 6.2 6.2 16 70-80 70-80 63 63 72 72 72 15.8 -70.6 -18.8 - 7.1 37.6 -19.2 37.1 - 3.6 23.8 -77.1 -23.7 - 7.4 41.3 -21.4 Mean 9 10 11 12 13 14 15 16 Mean 17 18 19 20 19 25.5 23.5 22 Mean 21 22 23 24 25 26 27 Mean Avoidance Indices Line 1 Line 2 Time Temp. 5.6 6.3 6.2 6.0 6.0 16.5 19 18 18.5 19.5 21.5 87.1 50.9 89 23.6 45.2 - 2.8 94 98.3 97.3 93.1 94 70.2 95.8 77.6 50 -76.? 4.5 -18.9 52.? 60.9 40.6 1.2 26 Coho Salmon All except the control experiments with coho salmon were conducted during the first two weeks of July. The coho salmon avoided concentrations of 1.5, 3.0, 4.5 and 6.0 mg./l. dissolved oxygen in some experiments. These experimental results are presented in Table 6. At 1.5 mg./l. the magnitude of the "index of avoidance" for the coho salmon was smaller than that for the chinook salmon and about the same as that for the largemouth black bass. The "avoidance index" was highly variable between experiments. The coho salmon exhibited more activity and greater speed in swimming than did the other species. It was observed that the coho salmon exhibited an erratic behavior when stimulated by a low dissolved oxygen concentration. An examination of Appendix Table C reveals the coho salmon usually made more entries into channels In a single experiment than did the other species. Largemouth Black Bass Generally, the largemouth black bass avoided channels having a concentration of 1.5 mg./1. of dissolved oxygen, as shown in Table 7. There appeared to be a slight avoidance of wster of 3.0 and 4.5 mg./l. dissolved oxygen, also. Among the bass tested in the younger there was more activity and schooling and snaller individuals than in the older 27 TABLE 6 AVOIDANCE BY COHO SALFON DURING JULY AND OCTOBER, 1956, OF WATER HAVING VARIOUS CONOENTR1TIONS OF DISSOLVED OXYGEN Exp. No. Date Mean Temp. L).O. °C. Size mm, Avoidanee Indicea Line 1 Line 2 Time in ./i. 69 70 71 72 Mean 74 75 76 77 7/7 7/9 7/9 7/10 1.7 1.7 1.6 1.5 18 18.5 21 18.5 7/6 7/6 7/10 7/11 7/11 3.1 3.1 3.0 3.3 3.0 17 18.5 20.5 18 7/5 7/5 7/12 7/12 4.5 '70 70 70 74 79 80 81 4.6 4.6 4.6 84 85 86 7/2 7/3 7/3 7/13 7/13 5.6 6.0 6.0 6.2 5.9 87 89 Mean 6.9 6.8 6.8 53.2 56.6 35.4 35.8 55.2 10.5 50.9 44.3 18.5 18.5 18.5 20.5 66-77 66-77 10.3 - 3.5 18.4 22.7 12 19.3 - 1.3 26.1 37.5 20.4 28.2 -13.2 35.3 18 17 18.5 18 20.5 63 63 37.1 - 9.5 2.2 4.6 6.2 35.4 20.7 7.7 8.8 17.3 18 23.6 - 4.7 35 27.5 16.1 -34.3 18.2 24.7 -64.9 3.1 2.7 18.5 76 76 63 76 73-80 8.1 10/17 10/17 10/17 56 59 43.4 10.6 37.3 3.7 39.6 32.6 Mean 88 34 79.5 77.4 66.1 28.4 7.4 25.2 1.6 28.8 22.4 Mean 82 83 29.1 68.8 68-74 70 73-76 64 63-67 Mean 78 - 4.1 39.8 22.4 42.5 25.1 13.5 13.5 13.5 85-105 80-100 80-90 12.6 9.2 -29.2 9.7 35 21.3 A6 42.3 4'i.6 28 TABLE 7 AVOIDANCE BY LARGEMOUTH BLACK BASS DURING AUGUST AND SEPTEMBER, 1956, OF WATER HAVING VARIOUS CONCENTRATIONS OF DISSOLVED OXYGEN Exp. No. Date 90 91 92 93 94 95 96 97 Mean Temp. D.O. . /i. 00. urn. 8/3 8/4 8/6 8/6 8/20 8/21 9/7 9/8 1.6 1.6 19.5 18.5 18.5 50-60 48-66 50-60 50-60 8/3 8/7 8/8 8/8 8/16 8/16 8/22 8/23 8/29 8/29 9/21 9/24 3.0 3.0 3.0 2.9 1.5 15 1.5 1.4 1.5 1.5 22 22 22 20 Size 80 60-75 78-91 17.5 91 19 50-60 50-60 70 70 80 80 66-74 85 Mean 98 99 100 101 102 103 104 105 106 107 108 109 Mean 110 111 112 113 114 115 116 117 118 119 120 121 122 Mean 8/8 8/9 8/9 8/9 8/30 8/30 8/30 8/31 9/19 9/19 9/20 9/20 9/20 3.0 3.0 3.0 2.9 3.0 6.0 3.1 18.5 19 21.5 20.5 21.5 21.5 21 18.5 20 17.5 3.1 20 4.5 4.7 4.5 4.6 4.5 4.6 4.6 4.6 4.3 4.6 4.? 4.8 4.7 22.5 20 20.5 22.5 18 19.5 20.5 18 18 20 17.5 18 19 91 91 85-91 74-91 30-40 80 80 80 91 85 72-Ba 91 85-91 85-91 85-91 85-91 65-70 Avoidance Indices Line 1 Line 2 Time 12.8 14.7 49.4 11.9 12.8 33.1 - 6.4 60.2 23.6 43.6 65.2 72.1 32.7 25.8 60.3 61.9 28.5 48.8 82.5 76.8 58.4 77.2 61.2 68.2 74.8 57.1 69.5 14.7 28.1 - 2.0 45.1 - 8.1 - 3 -20.2 - 2.3 0.5 4.4 5.4 18.9 6.8 16.9 42.9 - 9.7 22.0 - 3.0 14.2 -33.3 8.7 6.4 16.8 13.9 16.7 9.4 24.1 26.9 12.7 38.4 2.6 23.7 - 7.1 13 7.? -36.9 1.3 26.9 11.1 5.4 13.5 12.4 21.3 37.0 - 3.1 - 7.2 40.8 - 6.5 9.4 15.3 -24.6 0.6 13.9 9.7 69.9 16.4 18.0 7.0 .4 6.2 8.4 - 3.0 - 2.2 36.7 6.0 6.4 11.5 -26.8 10.1 5.7 5.0 .4 8.9 18.3 15.5 16.4 19.1 -30.2 1.8 1.3 10.2 29 TABLE r xp. No. Date iean D.O. Temp. 0G. (continued) Size mm. Avoidance Indices Line 1 tine 2 Time rng./1. 126 124 125 126 127 128 129 130 Mean 131 132 133 134 135 136 137 138 139 140 Mean 9/13 9/14 9/15 9/15 9/15 9/17 9/17 9/17 8/15 8/15 8/29 8/31 8/31 9/12 9/12 9/13 9/13 9/13 5.9 6.1 6.0 6.4 6.0 6.1 6.1 6.0 7.9 8.3 8.5 7.5 7.8 7.4 7.8 7.9 7.7 8.1 and larger fish. 20 17.5 17 18 20 17.5 19.5 20 78-91 78-91 85-104 91 91 E2-91 91 91 19.5 60-80 20 80 88 45-91 78-91 91 91 74-91 78-91 7C-91 20.5 20 21.5 17 17.5 16.5 1'l.S 13.5 10.4 -20.9 30.9 14.8 9.2 -19.1 -20.7 -11.5 - 0.89 8.2 .6 - 7.7 3.4 - 4.0 - 2.9 26.6 1.4 - 2.7 8.8 3.2 14.7 -23.1 16.]. 14.4 1.3 - 8.7 -25.6 - 9.8 - 2.6 26.6 - 3.? 10.9 11.6 23.2 -25.5 - 9.2 -16.5 2.2 - 4.7 11.7 - 3.0 7.6 - 3.9 3.7 52.2 1.7 -12.3 1.2 -19.2 31.]. 27.1 4.4 6.4 9.1 7.4 .2 - 1.1 - 5.4 1.8 5.2 Schooling was only observed in experi- merits No. 110 and 13]. with this species. Bluegill Table SiOwC that bluegill avoIded experimental channels at concentrations of' 1.5 mg./1. and 3.0 mg./1. dissolved oxygen on the basis of the time criterion and. avoided only 1.5 mg./1. dissolved oxygen on the basis of entries at the first and second lines. It appears that In 30 TABLE 8 AV0IDACE BY BLUEGILL SUiFISH DURING AUGUST AND SEPTEER, 1956, OF VATER hAY ING VARI0U CONCENTRATI8 OF DISSOLY&) OXYGEN Exp. Late No. Mean Tebip. D.(), °C. Size nm. Avoidance Indices Line 1 Line 2 TIme rng./1. 141 8/10 142 143 144 145 a/u. s/li 3/11 8/17 3/17 8/20 8/21 146 147 148 1.6 1.6 1.5 1.5 1.5 1.4 1.5 1.4 20.5 1.5 23.5 20 20 21.5 22 24 60-80 60-30 60-80 oO-80 75-90 75-90 70-90 75-90 Mean 149 10 151 152 153 154 Mean 8/12 8/13 s/is 3/14 3/16 8/2 3.4 3.0 3.0 3.0 6.0 10.5 19.5 20.5 20.5 19.5 .0 20.5 155 156 157 158 159 160 Mean 8/2 8/2 9/19 9/20 9/26 9/27 4.5 4.6 4.6 4.7 4.6 4.5 19.5 17.5 19 19 19 161 7/2o 8/i 9/18 9/18 9/18 5.9 21.5 19 17 162 163 164 165 Mean 166 167 168 Mean 8/15 8/15 8/15 6.0 6.1 6.1 7.9 8.3 8.3 17 18 22 19 20 21 40-50 70-(35) 75-(36) 72-(35) 7b-90 75-90 14.9 21.6 - 1.8 40.8 37.4 -10.1 40.5 3.9 18.4 37.2 4.1 3.9 81.1 75.4 25 76.3 42.6 51.8 24.3 18.4 2.0 -23.0 - .3 .5 -- -28.6 42.9 12 - 6.8 18.4 6.3 82.2 66.6 74.9 78.D 54 45.2 33.3 49 60.5 72.2 5.2 49 22.9 - 1.7 25.3 28.9 62-82 63-89 64-70 63-70 65-70 65-70 3.4 6.b -11.9 25.5 -13.4 -26.8 - 2.8 60-80 60-80 60-66 60-70 60-70 15.3 - 6.1 2.9 - 5.9 15.5 4.3 -40 2.4 - 4.9 25.1 .7 - 5.4 13.1 - 5.8 11.4 5.2 38.1 - 9.8 - 7.7 10.2 20.2 -64.9 60-30 60-80 75-80 -33.3 20 - 7.7 19.9 - 9.2 25.2 - 5.9 16.8 - 25.8 12.6 - 2.5 23.6 -32.1 -43.6 - 2.7 - 7.8 - 7.4 -32.3 9.4 11 .6 -15.1 31 bluegill, the "index of avoidance" based on the time criterion 18 greater than that based on entries. This is true of other species also, but differences were more pronounced in the bluegill. Individual bluegill in some experiments defended their po8ltions in the tank, thereby influencing the numbers of entries by other bluegill into channels. DISCUSSION Species Differences All species of fishes used in these experiments avoided at least the lowest tested concentration of dissolved oxygen. The species varied in the amount of avoidance and also in their manner of reaction. Avoidance of water with low dissolved oxygen concentrations was evidenced not only by the fewer entries into the channels with low dissolved oxygen than into control channels, but also by the shorter length of time spent by fishes In the experimental channels. In experiments performed at about the same date, chinook salmon exhibited considerably more avoidance than did the echo salmon at all concentrations of aissolved oxygen except 6.0 mg./l., at which concentration only the coho salmon showed avoidance. Certain behavioral 32 observations suggest possible re&sons for The juvenile these differences. coho salmon appeared to be more sensitive or irritable in water containing 6.0 mg./l. of dissolved oxygen than did juvenile chinook salmon. High indices of avoidance in the chinook salmon may in part have been due to a tendency for individuals of this species to their direction imediately upon sensin concentrations. reverse disagreeable Frequently the chinook 8almon did not enter the low-oxygen channels after a few initial entries. However, the coho salmon often became erratic upon entering low concentrations and appeared to be too disturbed to act in such a seemingly rational manner. often seen The coho salmon were to make a "start" upon entering a low dissolved oxygen channel, then dart this way and that until they found their way out of the channel. swimuing speed Also the greater of the coho salmon often would not enable them to turn from the channel entrance very quickly upon being disturbed by the low-oxygen water. Possibly as yet undemoristrated differences in the species to low concentrations of dissolved oxygen could tolerance of the two explain some of the differences in their responses. The coho salmon were the most active fish, followed by the chinook salmon at temperatures below 210 C., then by bass, and then bluegill. The magnitude of the "avoidance index" was about the same with the largomouth black bass as 33 with the echo salmon. The bluegill, and to some extent the largemouth bass, differed froui the salmon by continuing to even though enter water of low dissolved oxygen frequently, substantial avoidance was indicated by the time spent in the channels. Possible Factors Influencing Avoidance Season, water temperature, schooling behavior, size, age, swimrnin; speed, time of day, learning and experimental error may contribute to variations in results. Due to the time arid nurnoor of experiments, seasonal differences were only apparent in the chinook salmon. that a rise in the water temperature It has been shown resulted in an increase of the avoidance by chinook salmon of water of low dissolved oxygen, and that a lowering of the water temperature resulted in a decrease in the avoidance. This temperature-avoidance relationship was apparent not only in comparing experiments of the same season but also in comparing experiments of different seasons. Difference in schooling, swimming speed and activity which could have influenced avoidance have been considered above. Mechanism of Avoidance Jones (14, p. 414), in his studies of the stickleback, Gasterosteus aculeatus, the minnow, Phoxinus phoxinus, and 34 the brown trout, Slmo trutta, concludes that the basis for the selection of the high-oxygen zone is sirply "the removal of the stimulus to swimming." Stimulation of activity at low oxygen concentrations was also noted by Hoar, Black, and Black ( 9 , p. 97-99) and Vella (24, p. 323). Stimulation of activity, as suggested by Jones, appears to be a logical explanatIon for avoidance in the bluegill and in the lar2emouth black bass, avoidance at the first but the exceedingly high line, in particular, and also at the second line, in the chinook salmon introduces doubt as to whether stimulation was the only mechanism. Actually, in the present study, fish were more difficult to count during control experiments at low oxygen concentrations because of more or more varied movement at the higher concentrations. It is particularly noticeable that the chinook salmon of the June and July experiments appeared to avoid low dissolved oxygen concentrations at the first line to a greater degree than did the later group of chinook salmon or the other species. An almost complete avoidance of the low dissolved oxygen channels, foilowin; several trial entries, could be explained in several ways, namely: 1) the fish could learn that particular channels were undesirable, 2) the fish could learn that an increasingly uncomfortable situation followed continuing movement into 35 a channel which was slightly uncomfortble at the entrance, and 3) the fish could actually sense a low dissolved oxygen concentration close to the channel entrance. The magnitude of avoidance at the first line may be influenced by interaction among the fish. hesitate at the entrance of the A fish niiht channel or Inside the channel and other fish comIng towards the channel might notice the hesitation or turning of the first fish and turn because of it. This reaction aparently could have contributed to the production of a positive index of avoidance at the first line in some of the experiments with all species used. Evidence at' a considerable speed of recognition of water of low dissolved oxygen has been presented. connection it Is interesting to note that In this Powers and Clark (18, p. 104-107) speak of receptors located in the regions of the gills and innervated by the ninth cranial nerves as more Important fishes than system. In in controlling the normal breathing of the cephalic or central portion of the nervous a later publication, Powers and Clark (19, p. 110) reported that fishes with lateral line sectioned just distal to the gills do nerves not respond to the carbon dioxide tension of the water, even though branches of the ninth arid tenth cranial nerves leading to the gills are intact. Prior to this time it was considered that only ö6 the CO2 tension of the blood activated the changes in gill movements of fish. Further investiation should put more light on the presence or absence of external receptors of stimuli controlling respiration in fishes. Rapidly Lethal Levels Evidence that fish do detect and avoid some low dissolved oxygen concentrations has already been presented in this paper. The lower limits of tolerance for certain of these species of fishes can be found in the literature. There appears to be a great possibility that fishes may be able to detect and avoid concentrations of dissolved oxygen above their minimum tolerance where some barrier does not block the escape of fish from water of those concentrations. These .tudies have shown that fishes more strongly avoid low oxygen conceLtr8tions at hi than at lower water temperatures. the minimum oxygen oxygen water temperatures Likewise, a survey of tolerance limit (or minimum lethal level) in fisheries publications shows higher minimum levels at high water temperatures and lower minimum levels at lower water ternoeratures. juvenile chinook salmon avoided concentrations of dissolved oxygen as high The present study has shown that as 4.5 mg./l. at temperatures ranging iron l6 to 260 C. and avoided aissolved oxygen concentrations of 1.5 ar. 37 3.0 mg./1. at temperatures onerally lower (6.50 - 190 0.). Chapman (4, p. 197) found that a dissolved oxygen concentration below 2.5 rag./l. proved lethal to the fingerling chinook salmon In a very short time. Townsend and Earnest (23, p. 346) in their table of the published minimum oxygen tolerances of salmonid fish list 2.3 to 2.7 mg./l. as the minimum levels for aault chinoeks at 210 to 230 C, The coho salmon avoided the reduced oxygen channels at all concentrations. Davison (7, p. 15) found that at temperatures of 12° or 160 0,, coho salu4on were able to survive at concentrations of dissolved oxygen above 1.4 mg./1., while at 240 C., concentrations of more than 2.2 mg./l. were required. Similar lower oxygen tolerance limits for echo salmon are given for comparable temperatures by Townsend, Erichson, and Earnest (22, p. 1-47). Very few reliable tolerance studies have been conducted with centrarchids, Preliminary tolerance tests with 76 niu. largernouth black bass by Katz (unpublished data) appeared to indicate that their minimum tolerance limit was below 1.0 mg./1. at 20° C. The mean lethal oxygen limit of a related species, smailmouth black bass, Uicropterus dolomieu , was reported by Burdick et al. (3, p. 84-87) as 0.63 and 0.73 mg./1. dissolved oxygen at 52° F. (11.10 C.) and 1.16 and 1.05 mg./1. at 80° F. (26.67° C.) in two groups of experiments. If the 38 largemouth black iJass WOi.lc avolu in natural water 1.5 rrig./l. dissolved oxyen concentration as it dIO. in the avoidance tank, fish of this species night sometimes be able to move away frtxn areas of lethal concentrations of dissolved oxyen. Ecological Significance It Is unlikely that laborat-ory conditions can be xnsde to approxiniate natural conditicns. A ooderi tank, painted whIte, with vertical sides, shallow water, fluorescent lIghtIng and somevthat controlled variables presents a much different environment than does a streambed. The sharp transition from water of high dissolved oxygen to water of low dissolved oxygen encountered in the experimental tank would hardly over be encountered In nature. Probably these conditions are most nearly apiroachod at the confluence of a well aerated, unpolluted stream with a polluted stream with little or no dissolved oxygen. Normally in rAature there would be a dradual transi- tion from high corcentrations of dissolved oxygen to lower concentrations, the transition area being possibly many feet or oven miles in a stream instead of a few inches as in the avoidance tank. Small differences in dissolved oxygen concentration may not produce as large an avoidance react ton as is produced by the sharp boundary of the 39 In nature a dissolved oxygen gradient may avoidance tank. be vertical, as in the thermally stratified lake in which Moore tested the tolerance of fish over 24 hour periods (16, p. 319). It may appeer reasonable, perhaps, that the fish tested in these experiments would detect and avoid in the avoidance tank any dissolved oxygen concentration which they would detect and avoid in the native habitat. It cannot be assumed, however, that the fish would avoid ifl nature every dissolved oxygen concentration avoided in the experimental tank. Young salLilon ordinarily occur in highly aerated fresh- water streams. The sa1non fingerlings in these experiments appeared to choose water with a high level of dissolved oxygen similar to that found in their habitats in nature. The bass and the bluegill generally inhabit ponds, lakes, and sluggish streams. The latter two species appeared to be relatively insensitive to low concentrations of dissolved oxygen which occur in their native habitat more frequently than in the habitats of salmon. Under natural conditions recuced dissolved oxygen concentrations are always associated with elevated carbon dioxide tensions. Inasmuch as helford and Powers (21, p. 334) and Powers (17, p. 20) have reported that some fishes, notabli salmon and herring, are very sensitive and 40 react strongly to differences of carbon dioxide tension, it might be supposed that water with reduced diesolved oxygen concentration may be avoided in nature in part because of its high 002 tension. in the experiments reported here the 002 tensions were not increased, but on the contrary were probably reduced somewhat, in reducing the dissolved oxygen concentration by mns of nitrogen. The reactions considered here are believed to be reactions to dissolved oxygen only. ork needs to be done to determine if increased carbon dioxide rather than reduced oxygen may not be the controlling factor in avoidance of these related conditions in natural waters. StJLMRY 1. Studies of the avoidance reactions of some fresh-water fishes were conducted at the isheriee Research Laboratory, Oregon 8tate College, £ron June through November, 1956. 2. The apparatus used was a tank 112 inches long end 24 inches wide with four, oual, parallel, 36-ineh long channels, two of which v,ere supplied with water at some controlled ,Low dissoived oxygen concentration and two of which were supplied with well sorated water. The dissolved oxygen concentration was reduced by 41 bubbling nitrogen through the flowing water. ll four species of fish used in the experiments, 3. namely, chinook salmon, colic salmon, largomou.th biao bass, and the bluegill avoided some low concentrations of dissolved oxygen. 4. Groups of experiments for each species were conducted with water having dissolved oxygen concentrations of 5. 1.5, 3.0, 4.5, and 6.0 mg./l. Fish, during the 45-minute test periods, were recorded as tbr crossed the drain at the entrance to the channel and as they crossed a marker near the back of the channel. The number of fish present in the channel at 60-secand intervals was also recorded. separate "index of avoidance was calculated using each of the three counts. 6. it 7. The "index of avoidance" w2.s computed from the formula: 100 x "Index of voidanoet' CM - whore M represents the sum of the entries in all the channels divided by 2, and represents the awn of the entries in the experimental channels. hen the index it was based upon time, M represents the awn of the number of fish counted in all channels at 60-second intervals divided by 2 and represents the total number of fish in the experimental channels at 60-seood intervals. 8. In all species the "index of avoidiice was less at the 42 firit line than voidanoe' The 'index of t the a000rid iinO aio leas than the t the firat line index of avoithuee detorined bi the amount Of tiiie fiah rornainod in the channel$, 9. Chinook. 2alLiOfl in the June and July exrcrimenta nd 4.b rn./1. ivoided concentretiona of 1.t5, 3.0, diao1ved oxygen uid did not avoiä Q.) 10. Chinook ealmon in C./l. etebor, October, and November eporirnenti avoided coneentratton of i.L x 0 rng./i. diiao1ved oxygen, avoided 4.b onui slight1', :.nd aid. riot avoid d.0 mg./1, II. The chinook teted during the June arid Juii eries of experitonts gave higher t1indieea o± avoidanco' than aid the late eriea of chinook salaion or other a ' eel e s. 12. ;ater te;perature arpeared to be an iaortant factor oontributin to diUerencea in the magnitude of the ndicee of avoidance" between aeaaori 1$. The magnitude of voidaneo varied directly ter tcmerature in chinook 14. Coho ith the a1rron. aimon avo..ded vatcr ol' concentrations of l.i3, .0, 4.b, and lb. and within a .o m./l. diesolvod oxy;en. Coho aa1on bearne err tie imon enterin; the 1ocat dio1vod oçgcn channola. 43 16. Laraernouth b1ck baes voided 1.b m./1. dissolved. oxygen and appeared to avoid slightly eoucentrtions of 17. .O nd 4.b uig./l. Both on the basis of entries and on the basis of time bluegill voided concentrations of 1.5 mg./l., and on the basis of time only, they avoided 3.0 mg./l. dissolved oxygen. 13. some indiction of learning s noted when the '9.ndices of cvoidance" were oomared for sucocsive 15-minute periods with the chinook salmon of Juzie and July exporinents; little if any learning ws demonstrated in other experiments, r-;i ;L . wrion public holth odi for th induj3tricl , 3rett, J. . ' 1on, netbocition. :t nthrd tor, Hogo, nd x;in&tion of ow i(tJi ccl. eifio er:critro toierncc of zoung norhirwhu3. Jouruzi I oi the ;enuB fi'te ro:orch uord of Cnd .. 4, ork, ' : 25-; 7. LethJ. OXOfl ooncentrtion8 i. et i. . milriouth b&i. now ora fish br trout me journ1 1:35-7. J.nuar 19,4. nd 3urUiok, decreaec ox;i,fiect of jlbcrt Mo Leod, imrman, on ocke;e ,nu ohnook o1mon. 1 en aoiety rnotionB of the iericn ficherio t: 5, Obido3ter, . . Jtudie on fish migration. Ii. ihe influence of aa1initj on the diooraJ. of fiheu. oicn natu.rdl8t ó. otor inf1uing ralu B. oi1ins, ol nirit in ax 4ro.ou3 £iabou. U.. orvioe. p, 12herj onw offoct of 1oi eonobert centratiuis of d i >olved o::;7gen u'on juv4iniie 1ron. .tor' o the;if3. 11ver Cov1li3, Davison, roon ot&'te eol1e, lcJL4, :J, aiiiugton, overwient orb tins oUoe, ibi. (u,. fish tnd wildiifo oalletin no. 7, vol. 7. the orinttion ioudorofb, eter. t..erture M numb. lenvos. ec3tloflk3 01 mririe flEhOs to r:4ionti. Biologbc;1 buLLetin 7cl-&. l)3. J. 7. , Black, nd . 0. B1ck. icne . , :. 13peet of the oIoi.ogy of Itoh. oronto, University of oronto :rec, li. lii. (Univor$it of ¶oronto boioioai 3erios no. ublication of the Ont'bo iithorieu borori, rio. 71,, eioh 9. 4b Jones, Benjain ., et al. voidance reactions of salrLlonid fishes to øuip mill effluents, ;ewage nd industrial vaetes 2;140-14l. November lib6. 11. Jones, J. H. irichsen. The reactions of Pyosteus Dungitius L. to toxic solutions. Journal of 10. experimental hioogy 24:110-122. 1947. 12. 13. further study of the reactions of fish to toxic solutions. Journal of experimental biology 25:22-4. 1943. The reactions of the minnow, Phoxinus phoxinus L., to solutinne of phenol, orthodresol and paraorcsol. Journal of exDerimental. biology 28:261-270. 14. 15. 16. The reactions of fish to 10 Journal of exoerimental bio1oy 29:403-41. 3e.ptember 1952. iatz, fan. Unab1isbed dstc on oxyen tolersnee of oxygen concentrations. largemoath black bass. Corvallis, Oregon state ublic health service fisheries col.Lége-U.$. research laboratory. 1956. Field atadies of the oxygen requirements of certain froh-water fishes. ;oore, 7a1ter G. Ecology 29:19-29. 17. 1951. 1942. .xperiments and observations in the dwin B. behavior of marine fishes toward the hydrogen- powers, ion ooicentration of sea-water in relation to ub1icstheir migratory movements and. habitat, tion of the Fuget sound biological station (b7) :1-22. 1921. 18. 19. thin B. and iobert T. Clark Jr. Control of norma.l breathing of fihe by receptors located in regions of the gills and innervated by the iXth and th eranici nerves. meriean joial of physiology l:lC4-10?. December 1942. Powers, Further evidence on chemical factorsfeeting the migratory movements of fishes, eseoia11y salmon. Ecology 24:19-113. 20. Shelford, Victor E. and C. Allee. Reactions of fishes to gradients of dissolved atmospheric . gases. Journal of experimental zoology 14: 207-263. 1913. B. and Edwin I. Powers. An experiienta1 study of the movemnents of herring and other marine fishes. Biological bulletin 28:315-334. 1915. 21. Shelf ora, \'lctor 22. Townsend, Lawrence 1)., Arne Erichsen, and Don Earnest. Progress report on field investigations and research. Washington state pollution committee, Olympia, Vashinton. p. 1-47. 1938. 23. Townsend, Lawrence D. and Don Earnest. The effects of low dissolved oxygen and other extreme conditions on salmonoid fish. Proceedings of the sixth Pacific science congress 3:345-351. 24. 1940. wel1s, Morris M. The resistance of fishes to different concentrations and combinations of oxygen and carbon dioxide. Biological bulletin 25:323-347. 1913. / 47 APPENDIX TABLE A DISSOLVED OXYGEN COCENTRT1ON AT 12 lNCiii,S A1Li 3 INChES INSIDE CkiAiNEI OUTSIDE CHANNELS (for channels A and B) Channel A Experiment Number 12 in. AI4D 3 INC1S Channel B S in. 6 in.O. 12 in. 9.8-9.6 1.4-1.4 9.8-9.1 1.8-0.9 8.2-8.3 1.5-1.5 8.6-8.8 1.6-1.4 10.0-9.7 2.8-2.8 10.2-10.0 9.2-9.2 6.8-5.8 9.2-9.2 6.4-5.6 6.3-6.9 4.3-6.0 2.1-2.0 9.1-9.2 1.6-2.0 8.7-9.0 2.1-1.8 8.4-8.6 1.5-1.5 8.6-8.6 3.1-3.1 10.2-10.0 3.2-3.1 3 in. S in.O. 1.6-1.9 9.6-9.4 1.6-1.9 8.7-8.7 2.1-2.9 8.4-8.6 1.7-1.6 8.4-8.6 6.0-3.4 10.1-10.0 3.0-3.0 5.3-5.1 5.9-8.3 5.5-7.9 6.4-6.6 7.0-6.3 6.5-6.0 5.8-5.8 5.5-5.4 6.4-5.8 8.2-8.0 5.2-5.5 Chinook Salmon 1 2 3 4 5 6 7 8 9 10 11 '10 A. 17. ad A. 14 15 16 17 18 19 20 21 22 23 24 25 9.4-9.6 2.0-1.6 9.6-9.6 1.1-0.8 8.6-8. 1.5-1.5 8.6-8,6 1.5-1.4 9.8-9.8 2.9-3.1 10.1-9.7 00 .. U Ca ) . '.1 7., 00 0r7 r)cU . 0 ') a.) 7'1j. 8.3-8.3 .0-3.1 8.5-8.4 4.6-4.4 9.1-9.2 6.9-8.7 4.5-4.5 6.3-6.2 9.9-9.5 9.1-8.8 6.4-6.2 9.4-9.3 8.4-8.1 3.0-3.1 8.7-8.3 4.5-4.3 9.1-9.1 8.9-8.7 4.6-4.4 6.2-6.1 9.8-9.8 9.3-9.1 6.6-6.4 9.2-9.1 a.) Ca . '7.6-7.7 7.0-5.6 8.1-8.4 '1.8-4.8 9.9-8.5 7 J .z ad 7 A t . ad £ iz; a., V7 A. I 7.9-7.1 4.0-6.3 7.6-7.1 7.8-7.2 8.5-7.8 8.0-7.5 6.1-6.6 6.4-7.5 9.2-8.4 10.0-8.6 7.4-7.6 8.5-7.5 0 0 0 .. . a., C)" A. a) a.. ., '-. . a, 2.8-2.9 8.9-8.9 2.8-2.8 8.7-9.2 4.6-4.6 4.4-4.6 8.5-8.4 10.1-10.4 6.2-6.3 5.8-5.4 9.8-9.5 6.0-6.1 0U 0s J 'Z aJ ad * ad C) a_ A 3.0-2.9 8.6-9.0 3.0-6.0 8.9-9.2 4.6-4.5 4.3-4.6 8.5-8.5 10.4-10.1 6.4-6.5 5.8-5.8 9.8-10.0 6.0-6.2 4D7 i tZ a, C 5.0-6.2 7.5-6.7 7.2-6.2 7.8-7.5 7.0-6.4 7.5-6.6 6.0-6.6 8.3-7.7 8.6-8.o 6.8-6.4 8.2-8.2 7.6-7.1 TABLE A (continued) Experiment Number 26 27 28 29 30 31 32 34 35 36 37 39 40 4]. 42 43 44 45 46 4? 48 49 50 51 52 53 54 12 In. 6.0-6.0 6.0-6.0 1.6-1.8 9.5-9.4 1.5-1.6 9.6-10.0 11.4-10.8 1.6-1.7 11.4-12.2 1.6-1.7 3.0-3.1 10.4-10.8 3.0-2.9 10.2-10.2 11.8-12.2 3.0-3.2 11.1-11.6 2.9-3.1 4.6-4.6 4.6-4.4 4.ô4.6 11.2-10.8 11.4-11.4 4.2-4.7 12.2-11.6 4.5-4.5 6.0-6.0 10.4-10.8 5.9-5.8 Channel A 3 in. 6.1-6.0 5.9-6.0 1.8-1.8 9.7-8.7 1.6-1.5 10.0-10.5 11.7-10.0 1.7-1.5 11.4-11.0 1.6-1.7 3.0-3.0 10.6-10.6 3.0-6.0 10.3-10.0 11.8-11.8 2.9-2.8 11.1-11.1 2.9-3.1 4.5-4.6 4.6-4.5 4.6-4.6 11.0-10.7 11.4-11.0 4.7-4.7 12.2-11.8 4.5-4.5 t,.9-6.l 10.6-10.2 5.9-6.0 Channel B 3 in,0. 12 in. 3m. 3 In.0. 6.6-6.8 6.2-7.4 6.0-6.3 7.5-7.1 6.5-7.1 7.8-9.2 8.0-7.7 7.8-6.8 r,8..79 7.2-6.8 5.6-5.6 9.1-7.6 8.1-8.1 8.1-8.0 9.3-9.2 8.4-8.0 8.4-8.6 8.0-8.0 8.1-8.4 9.0-c.6 6.7-8.6 7.5-7.4 9.3-9.5 9.1-9.1 8.8-8.5 10.2-10.1 2.0-1.7 9.8-9.7 1.6-1.4 1.2-1.4 10.8-11.2 1.5-1.6 11.7-11.1 9.3-8.9 2.9-3.2 10.1-10.4 3.0-6.1 9.1-9.1 8.8-8.5 7.4-7.7 7.4-7.3 9.2-6.0 7.6-7.1 6.0-5.9 8.2-7.1 7.6-7.6 8.5-7.8 7.1-7.1 7.7-7.1 9.4-.0 12.0-1.0 10.0-9.6 9.4-8.9 7.5-7.? 8.0-8.0 7.4-6.9 4.5-4.4 11.9-11.? 10.6-10.9 .0-2.9 11.0-11.2 2.9-6.2 11.4-11.3 9.8-10.2 10.4-10.6 10.6-10.5 4.5-4.6 4.4-4.5 i.0.i-10.2 1.6-1.6 9.6-9.8 1.4-1.5 1.2-1.4 10.8-11.2 1.4-1.5 11.3-11.1 9.4-9.2 3.0-6.2 10.4-10.6 6.0-6.0 2.8-2.9 10.9-11.2 2.9-2.8 li.2-11.3 10.1-10.2 l0.7-iU.6 10.7-10.4 4.4-4.6 4.4-4.4 12.0-12.0 4.4-4.5 11.7-11.? 10.9-11.0 b.9-b.E3 b.0-.9 10.b-10.7 10.4-lo.8 r,.2_7.4 5.5-7.6 8.3-.3 8.0-7.3 9.5-9.6 8.4-8.4 8.5-8.4 rl.5.7,5 8.2-8.7 9.2-8.2 6.3-9.4 7.5-7.3 9.7-9.6 9.4-9.4 9.7-8.8 1.9-7.9 8.0-7.8 7.6-7.1 TABLE A (continued) Channel A Experiment Number 55 56 57 58 59 60 61 62 63 64 65 66 67 68 12 n. 10.7-10.8 6.0-6.1 9.-9.9 5.9-6.0 12.0-11.5 6.3-6.6 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 Channel B 3 in. 3 in.0. 12 in. 3 in. 3 in.0. 10.4-10.5 6.0-6.1 9.3-9.3 6.0-6.0 12.0-11.5 6.0-6.2 9.8-9.8 8.6-9.0 9.0-9.0 7.1-7.9 7.5-7.2 7.3-7.1 8.3-7.8 9.0-9.2 8.4-8.2 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 6.0-6.3 9.1-9.1 5.9-5.7 11.5-11.5 6.1-6.1 11.8-11.8 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.? 9.7-9.8 6.0-6.3 9.3-9.1 6.1-5,? 11.3-11.0 6.2-6.1 11.4-11.9 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10,2 10.2-9.8 9.8-9.7 9.7-9.8 7.0-7.6 7.7-7.7 7.2-6.8 8.3-8.0 9.0-9.2 9.0-8.8 9.3-9,8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 6.7-7.1 7.9-7.5 5.2-6.3 9.0-7.8 9.2-7.o 5.6-7.3 8.2-8.0 7.3-7.6 5.3-5.9 6.0-'.6 8.8-8.3 8.0-7.4 7.8-6.9 10.2-9.7 ).7-i.9 9.0-9.1 1.5-1.6 3.1-3.3 8.7-8.9 ô.1-2.9 .4-10.1 8.4-9.2 9.0-9.1 4.6-4.8 4.6-4.6 9.4-8.9 10.0-9.6 98-10.2 10.2-10.2 10.2-.8 9.8-9.7 9.7-9.8 Silvar Sa1ion 69 70 71 72 73 1.5-1.6 9.2-8.8 1.6-1.5 9.4-9.3 9.o-9.? 1.5-1.? 8.c-9.0 1.6-1.5 9.4-9.2 0.6-9.7 3.v3.0 75 9.4-9.1 76 77 78 79 80 81 3.1-3.0 4.5-4.4 9.1-9.1 9.2-9.4 4.6-4.5 9.5-9.2 3.1-3.6 3.1-3.0 4.2-4.5 9.2-9.1 9.4-9.3 4.6-4.6 1.-1.8 8.1-9.4 1.4-1.8 3.1-6.3 9.1-8.9 3.0-6.0 9.7-10.1 8.4-9.5 9.0-9.2 4.5-4.b 4.6-4.6 9.3-9.0 7.6-7.9 7.4-5.5 5.9-5.? 6.6-6.4 5.5-6.5 r/7rIo 8.4-/.8 7.2-7.9 7.7-7.2 3.b-7.9 8.2-7.4 6.8-7.0 7.7-7.5 0 TABLE A (continued) Channel A Experiment Number 62 84 85 86 87 88 89 3 In. 3 in.0. 12 in. 9.7-8.9 9.8-9.6 6.2-5.9 9.1-9.2 6.0-5.3 7.0-6.8 6.8-6.8 6.8-6.8 9.0-i .5 8.8-8.5 0.6-8.4 7.5-7.9 7.6-7.7 7.3-7.5 9.0-6.8 6.8-6.8 6.8-6.8 5.6-5.6 4.4-5.7 8.4-7.9 3.4-6.1 8.0-7.6 8.4-7.2 Largemoutli Base 90 1.8-1.6 91 9.1-9.2 92 93 94 95 96 1.5-1.o 8.8-9.0 .0-8.1 1.4-1.2 l.b-1.5 crI c 98 9.7-9.2 3.1-2.9 8.8-8.8 2.'-2.8 8.9-8.9 3.0-3.2 9.0-8.7 3.0-2.9 9.3-8.9 3.0-3.0 3.1-3.3 99 100 101 102 103 104 105 106 107 108 Channel B 12 In. z c J S J 9.6-9.8 6.0-6.0 9.6-9.3 6.0-5.9 7.0-6.8 6.8-6.6 6.8-6.8 1.8-1.6 9.b-9.3 1.5-1.6 8.8-8.5 8.9-8.6 1.4-1.3 1.6-1.5 17 17 S ¶.J 9.8-8.5 3.1-6.0 8.9-8.9 2.8-2.8 9.2-8.7 8.8-8.9 3.0-3.1 9.5-9.0 3.0-3.0 6.1-ô.3 33_5,7 5.4-5.6 ci )S 17 4 5 3 In. 3 In.0, 5.9-5.8 6.2-6.0 9.2-9.3 6.4-6.3 9.5-9.4 7.0-6.8 6.8-6.8 6.8-6.8 7.7-8.2 6.9-7.6 8.2-7.9 6.4-7.3 7.6-7.8 7.0-6.8 6.8-6.8 6.8-6.8 9.5-9.3 1.7-1.5 9.0-9.1 1.5-1.5 1.4-1.3 9.0-8.6 7.7-7.6 9.4-9.4 1.7-1.4 9.2-9.2 1.5-1.5 1.4-1.4 9.0-8.4 5.2-7.0 6.7-7.0 8.3-6.1 6.7-6.5 4.6-6.6 7.3-6.3 4.8-6.8 i. (L, I A r Ju I 9.2-9.2 6.2-6.3 9.4-9.2 7.0-6.8 6.8-6.8 6.8-6.8 r 17 9.3-8.0 3.8-5.6 8.3-8.1 6.4-6.0 8.7-7.2 6.0-6.5 8.4-7.8 6.7-6.0 8.-8.0 5.8-5.5 5.4-6.8 3.0-3.0 8.9-9.1 3.1-3.0 8.5-8.6 3.0-3.0 8.9-9.0 2.8-3.1 8.8-8.8 3.0-3.1 9.4-9.4 10.0-10.0 8.0-'7.6 L . ti 2.7-2.8 9.2-9.3 3.0-2.9 8.7-8.6 3.0-3.0 9.1-8.6 2.9-3.1 8.8-9.1 3.0-3.1 9.3-9.3 10.0-10.0 A Z. 4.1-7.0 6.9-5.7 6.0-6.6 6.2-7.0 8.0-7.1 6.2-7.6 7.5-7.6 7.6-7.6 6.4-6.6 5.7-6.6 6.8-7.9 Cr1 TABLE A (continued) 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 12 in. 3 in. 3 in.0. 12 in. 3 in. 3 in.0. 9.6-9.5 8.4-8.6 4.5-4.7 8.9-8.8 4.6-4.5 9.3-8.9 3.0-3.0 4.6-4.5 9.2-9.2 10.0-9.8 4.6-4.5 10.0-10.2 4.8-5.2 9.6-9.8 9.4-9.4 8.8-8.7 4.8-4.6 8.9-8.8 4.7-4.5 9.5-9.0 3.0-3.0 4.6-4.6 9..-9.4 10.0-9.8 8.4-3.6 7.9-7.4 5.6-6.7 8.7-7.8 5.8-6.5 8.6-8.0 5.8-5.5 5.6-6.7 9.0-8.0 10.1-9.8 4.6-5.2 9.6-9.6 9.5-9.4 6.0-6.1 10.0-9.9 8.8-8.6 7.2-8.4 3.1-3.0 4.4-4.5 8.8-9.1 4.5-4.4 8.8-8.9 3.0-3.1 9.4-9.4 9.1-9.2 4.6-4.6 4.6-4.6 9.8-9.7 4.5-4.5 10.0-10.0 4.6-5.1 5.9-6.2 9.8-9.8 6.1-6.0 3.2-3.2 4.5-4.4 9.0-9.1 4.7-4.5 3.6-8,7 3.0-3.1 9.3-9.3 9.1-9.1 4.6-4.7 4.4-4.5 9.8-9.4 4.6-4.5 10.0-9.6 4.6-5.0 5.9-5.8 9.9-10.0 6.0-6.0 8.0-7.8 7.0-7.8 6.7-7.3 8.3-7.7 b.9-6,7 6.4-6.6 5.7-6.6 7.7-6.8 6.7-6.7 8.0-8.5 8.4-8.8 6.2-7.0 7.9-8.6 6.9-9.6 7.6-7.6 8.4-8.0 .2-8.9 6.1-6.2 10.0-10.0 £ a. 127 128 129 130 131 132 133 134 135 136 13? Channel B Channel A Experiment Number I J J. 1 SJ . a. 9.6-9.4 6.2-6.0 9.6-9.4 6.2-6.1 7.9-8.0 8.4-8.6 8.5-8.5 7.3-7.6 77..79 7.2-7.2 7.6-7.9 4547 'J L .J J I L. 9.8-9.3 6.2-6.2 9.2-9.2 6.1-6.1 7.9-8.0 8.4-8.6 8.5-8.5 r,.3_17,6 I7,7...79 7.2-8.t, r/5_rf 6 9.6-.6 7.8-8.1 6.8-7.5 8.4-8.2 7 I 1 - rI a. ( S i 9.2-8.4 7.6-7.6 8.2-8.5 3.4-8.3 7.9-8.0 8.4-8.3 8.5-8.5 7.3-7.6 77,..79 '7.2-7.2 r/272 7.6-7.9 7.6-7.9 t J (T 7 S J 6.2-5.9 10.4-10.2 6.1-6.0 9.2-9.6 7.9-8.0 8.4-8.3 8.5-8.5 7.3-7.6 r,7_79 7.3-7.3 7.6-7.9 r rj I - (.1 £ ¼) 6.9-.6 r 'D ¼) r / 1 r,,9_8.O 6.8-7.8 7.8-7.9 8.5-8.1 8.9-8.4 7.9-8.0 8.4-d.o 8.5-8.5 7.3-7.6 7.7-7.9 7,3_773 7.6-7.9 8.5-8.5 7.3-7.6 7.7-7.9 7.3-7.3 7.6-7.9 6.2-5.8 10.0-10.2 6.2-6.0 9.6-9.4 t TAbLE A (continued) '-==---------- 12 in. 138 7.3-7.3 3 In. rj 3 in.0. 17 I S J I 7.7-7.7 7.8-7.8 140 B1ue4l1 E.unfih 141 1.5-1.9 1 L 5 142 143 144 145 8.-9.0 9.1-8.9 1.5-1.5 8.8-9.0 9.2-9.9 1.5-1. 1 1'6 147 148 149 150 151 152 153 154 155 156 157 158 159 loO 161 162 .Lbó --- Channel A Experinent Number 19 --------- 1.4-1. c . 1 -1 C) C . o 9.1-8.9 9.1-9.0 1.b-1. ai 5 aU ) 1.4-1. a 3.0-3.0 Li S 9 3.2-2.8 3.0-p 9 9.1-8.9 9.1-3.0 3.0-2.8 J. U 4.5-4.6 9.6-9.5 9.6-9.8 9.8-10.0 4.4-4.6 5.8-5.8 9.5-9.4 9.8-9.8 9.6-9.6 9.8-10.0 5.8-6.0 9.5-9.0 lu.0-9.8 rj -z I r - I Li 12 in. ry i . A __________ Channel L 3 in. A r i . - r; - , . . 3 in.0. r A f . ri - , 7.7-7.7 .8-7.8 7.7-7.7 7.8-7.3 7.7-7.7 7.3-7.8 7.7-7.7 7.8-7.8 C Js 0L10 Qr; L' I aJ. U .JU. 5.5-6.6 o.8-7.b aL5 I 1.5-1.5 C a . i. - . a 7.8-7.6 8.8-8.0 o.0-4.4 R J. 0 C S 0 3.3-5.7 U A R US r CO 7.6-6.1 4.6-5.3 4.5-6.3 A A . 'i I C . a 6.1-6.4 9,4_r/,9 0 - c . I J A A 1 S 2 ) a . (1 1.5-1.4 1.6-1.4 9.0-9.1 1 1 J S U i.0-O.6 9.0-8.4 a 0 a ). a a, a a _ .J ' 8.8-8.8 9.0-8.9 p a .a oa v.4-9.2 4.6-4. 8.6-8.7 4.-4.6 9.6--9.6 4.6-5.1 4.6-4.6 10.5-10.0 8,9-9.0 t.8-5.o 5.9-6.1 8.7-8.4 3.6-8.2 6.2-6.9 9.3-7.7 7.6-8.3 a L . L 1.4-1.5 1 1.6-1.4 1.6-1.4 8.9-8.4 I 01 C SLsi) . i 9 0 . 6.1-3.1 9.-8.&3 9.0-8.9 a a. rt -b 8 9.4-9.ô 4.5-4.8 4.5-4.7 4.6-5.0 4.6-4.6 10.5-10.4 9.0-9.0 b.8b.9 6.0-6.2 C . 0 b. - 6.1-6.5 6.2-7.4 3.1-7.0 U S U C 7.3-6.6 ai. c r; :.a ' . 17_.4Z 5.4-6.1 r/36] 7.1-6.9 .0 i 8.8-7.3 5.8-9.5 8.5-7.4 C.9-9.6 8.4-8.3 7.8-8.6 8.2-7.7 6.0-7.9 7.8-7.8 TABLE A (continued) 12 ifl. 164 165 166 167 .J .J L Channel h Channel A Experiinent 6.1-6.1 9.6-9.2 3.0-7.9 8.2-d.4 QjU %JZ - CL g) 3 ln. 3 in.O. 12 in. 6.0-6.0 9.5-.3 7.6-/.3 8.9-3.2 ---- --- 9.8-9.6 6.0-6.1 --- ---- --- - - - - - in. 9.8-9.8 6.2-6.0 3 in.0. 7.8-8.4 8.8-8.4 --- - c-I TABLE B DISSOLVED OXYGEi CONCEiTRATIuNS AT 12 INCHES AND 3 INCHES INSIDE CHANNELS AND 3 INCHES OUTSIDE CHANNELS (f or channels C arid D) Channel C Experiment Number 12 in. ins 3 in.O. 12 in. 7.0-7.9 9.3-7.8 7.2-6.6 6.2-7.6 6.7-6.4 5.4-6.0 5.5-5.5 6.5-6.1 8.0-7.9 8.6-6.8 8.1-7.8 1.5-1.5 9.3-9.4 Channel D 3 in. 3 th.O. Chinook Salmon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 9.8-9.7 1.9-1.7 9.9-9.0 1.2-1.5 8.2-8.1 1.5-1.5 9.1-9.1 1.6-1.5 10.5-9.0 2.8-3.2 10.2-10.2 2.9-2.9 3.0-3.0 8.3-8.7 3.1-3.1 8.7-8.7 4.7-4.0 9.0-9.1 9.0-9.0 4.5-4.6 5.5-5.9 9.9-10.0 9.3-9.1 6.4-6.3 9.2-9.4 9.6-9.7 1.6-1.4 9.9-9.6 2.1-1.7 8.2-8.1 1.5-1.5 9.1-8.8 1.7-1.5 10.0-10.0 3.2-6.1 10.2-9.8 2.9-2.9 3.6-3.0 8.7-8.9 3.0-3.1 8.7-8.7 4.9-4.2 8.9-9.1 9.0-9.0 4.4-4.7 5.5-6.0 10.1-10.1 9.4-9.4 6.4-6.4 9.3-9.3 64-5.9 7.4-6.4 7.2-6.6 5.9-6.2 7.5-6.7 7.7-7.4 i.O-7.5 7.9-7.6 6.2-4.8 8.1-8.2 8.6-8.4 6.8-7.7 8.4-8.6 '7.7-7.3 .9-1.0 8.7-9.0 1.7-1.7 8.6-8.6 1.6-1.5 8.6-8.6 3.0-3.2 9.8-9.8 2.9-3.0 9.3-9.4 8.0-8.4 5.0-3.0 8.9-8.9 3.0-2.8 8.9-9.0 4,54.45 4.3-4.7 8.9-8.5 10.0-9.9 6.3-6.2 5.8-5.4 9.6-10.0 6.2-6.3 1.8-1.6 9.4-9.8 .9-1.5 8.7-8.7 1.9-1.9 8.6-8.6 1.7-1.6 8.5-8.4 3.0-3.3 9.6-9.8 3.0-2.9 9.6-9.2 8.5-8.4 2.9-3.1 8.5-9.0 3.0-2.8 8.8-8.7 4.7-4.5 4.3-4.6 8.5-8.2 10.0-9.8 6.4-6.1 5.8-5.4 9.4-9.7 6.1-6.2 4.2-4.2 8,2-8.0 6.2-6.7 8.4-7.9 5.8-5.6 4.0-4.0 6.6-6.1 7.9-7.9 8.6-8.1 5.4-5.6 6.5-6.4 7.4-6.1 4.2-6.1 5.8-6.3 7.2-6.7 7.9-7.6 7.1-6.8 5.7-6.4 6.9-8.1 8.7-8.2 8.7-8.5 6.4-6.5 9.4-8.6 7.1-7.3 TABLE B (continued) Experiment Number 12 in. 42 43 6.1-6.1 6.2-6.0 1.6-1.8 9.9-10.1 1.5-1.5 9.6-10.1 11.5-10.3 1.4-1.6 11.4-11.4 1.6-1.4 3.0-2.9 10.8-10.6 3.2-3.0 10.2-10.1 11.7-12.1 2.8-2.5 11.1-11.6 2.9-2.7 44 4.-4.5 45 46 4.6-4.6 4.5-4. 11.2-11.2 11.4-11.6 4.6-4.5 12.2-12.0 4.4-4.5 6.0-6.2 10.7-10.6 5.9-5.8 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 47 48 49 50 5]. 52 53 54 Channel C 3 in. 6.2-6.2 6.0-6.0 1.6-1.8 9.9-10.2 1.5-1.5 9.8-9.9 11.6-10.4 1.4-1.5 11.2-11.3 1.8-1.7 3.0-2.8 10.8-10.4 3.2-.0 10.0-9.9 11.7-11.9 2.9-2.4 11.0-11.4 2.9-2.? 4.6-4.4 4.6-4.6 4.5-4.4 11.0-11.2 11.5-11.4 4.8-4.5 11.6-11.7 4.5-4.4 5.8-6.1 10.6-10.6 5.9-5.8 Channel D 3 in.0. 12 in. 3 in. 3 in.0. 7.6-6.9 7.4-7.3 6.3-6.5 7.6-7.4 7.0-7.3 7.8-7.8 7.7-8.4 8.4-7.2 7.3-7.3 7.3-6.8 7.8-7.6 6.2-7.6 8.6-7.0 8.0-7.2 9.9-9.6 8.2-8.2 8.3-8.6 8.0-7.8 8.3-8.7 8.9-7.8 8.6-8.4 7.9-7.6 9.2-9. 9.7-9.2 9.7-9.4 9.7-9.2 7.9-7.6 8.0-7.9 7.4-6.9 8.8-9.1 8.8-8.6 10.5-10.2 1.8-1.6 10.1-9.6 1.6-1.5 1.7-1.5 10.2-11.0 1.7-1.7 11.0-11.2 9.7-9.2 3.6-3.2 10.2-10.2 .1-3.0 3.0-3.0 11.5-11.1 3.0-3.0 11.2-11.4 10.2-9.8 10.7-10.8 10.6-10.4 4.4-4.5 4.8-4.5 12.0-11.8 4.5-4.5 11.9-12.0 10.9-10.9 6.1-6.1 10.6-10.8 9.1-8.6 8.8-8.6 10.5-10.0 1.8-1.6 9.8-9.7 1.6-1.5 1.6-1.5 10.7-11.2 1.7-1.6 11.1-11.2 9.6-9.6 3.0-3.3 10.4-10.2 3.1-3.0 3.0-3.0 11.1-10.7 3.1-3.0 11.4-11.3 10.2-9.9 10.7-10.7 10.6-10.3 4.4-4.5 4.7-4.4 12.0-12.0 4.4-4.5 11.8-11.6 10.9-10.8 6.1-6.2 10.4-10.3 7.4-7.4 8.2-7.4 7.0-6.6 7.6-6,8 7.8-7.4 7.8-7.2 7.6-7.8 8.6-8.0 6.9-6.6 7.6-7.6 7.5-7.2 5.6-6.6 8.5-8.2 7.9-8.2 9.6-9.6 8.2-8.5 8.3-8.6 8.3-8.0 8.6-8.9 5.1-5.9 9.0-8.9 7.2-7.5 9.2-9.3 9.6-9.3 9.8-8.7 9.6-9.4 7.8-7.8 6.7-7.3 7.6-7.5 TABLE B (continued) Channel C 12 in. 3 in. 3 in.O. 12 in. Channel D 3 in. 10.5-10.8 6.u-6.0 9.0-8.9 6.1-6.1 12.0-11.9 6.1-6.2 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 10.5-10.5 6.0-6.0 8.8-8.5 6.1-6.1 12.0-11.8 6.1-6.3 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 8.2-8.0 7.8-7.4 7.1-7.3 8.4-8.4 8.8-9.1 9.1-8.9 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 6.1-6.6 10.2-9.8 6.0-5.6 10.9-11.1 6.2-6.2 11.6-11.6 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-i.7 9.7-9.8 5.9-6.6 10.2-9.7 6.0-5.6 10.9-11.0 6.0-6.0 11.4-11.6 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 10.2-9.8 9.8-9.7 9.7-9.8 8.5-8.6 9.0-8.8 9.0-8.8 9.8-9.8 8.6-9.0 9.0-9.0 9.8-10.2 10.2-10.2 1.7-1.9 9.3-9.4 1.6-1.7 9.5-9.2 S.2-i.4 7.1-7.1 6.1-6.8 7.6-6.9 6.3-7.1 8.9-7.4 9.8-9.4 1.6-1.7 9.3-8.9 1.5-1.5 2.9-3.1 9.6-9.2 1.7-1.6 9.1-9.0 1.5-1.b 6.2-3.2 8.9-8.1 6.1-6.5 7.8-7.3 ô.6-5.9 5.5-6.8 Experiment Number 55 56 57 58 59 60 61 62 63 64 65 66 67 68 3 in.0. 7.0-7.8 7.-7.5 '7.0-6.9 9.8-9.7 9.7-9.8 Silver Salmon 69 70 71 72 73 P7A I 75 76 77 78 79 80 81 1.6-1.7 9.3-9.5 1.7-1.6 9.4-9.0 9.4-9.7 C) Z C) C) 'Z 1 . J a. oJ . t. 9.4-9.3 3.2-3.8 6.1-3.0 4.4-4.6 9.1-9.1 9.4-9.4 4.6-4.6 9.3-9.5 . .. 3.2-.8 3.0-2.9 4.4-4.6 9.5-9.0 9.4-9.4 4.7-4.6 Cl i. a.. ) ) ,i . 'i 8.4-8.2 7.2-8.4 6.0-6.7 7.2-7.8 8.8-8.0 7.7-7.0 7.6-7.3 C) A . ' Ti . i 2.8-2.7 9.7-10.3 8.3-9.2 9.1-9.1 4.5-4.5 4.4-4.6 9.3-9.0 C) C) ' . i C) 7 . ta 3.0-2.8 9.8-10.2 9.1-9.4 9.1-9.2 4.6-4.6 4.7-4.7 9.3-9.0 0 Z 0 u . ti 7.5-8.1 7.4-8.2 8.4-7.8 8.3-8.0 5.1-7.6 6.4-6.6 7.7-7.4 TABLE B (continued) 12 in. Channel C 3 in. 3 in.0. 12 in. Channel D 3 in. 3 in.O. 9.7-9.2 9.-9.b 8.0-8.4 5.5-5.3 5.8-5.6 6.1-8.4 Experiment Number 82 03 84 85 86 87 88 89 U C b.2-6.0 9.4-9,4 .0-b.9 7.0-6.8 .8-6.8 6.8-6.8 La remouth Bass 90 1.5-1.4 9.6-9.2 91 1.5-1.5 92 93 9.2-9.0 94 9.0-8.4 95 1.6-1.4 96 97 98 99 100 101 102 103 104 105 106 107 108 1 .J_. 9.9-9.8 9.1-9.4 U U 7 v.2-9.2 2.9-3.0 9,2-9 0 3.0-3.1 9.1-9.1 2.7-3.0 9 o-8 9 2.9-3.1 2.8-3.0 . - U7 0 .) 0?. . U L I I G.i-6.l 3.4-9.4 6.0-3.0 7.0-6.8 6.8-6.8 6.8-6.8 3.1-8.2 7.8-7.4 7.6-7.6 7.0-6.8 6.8-6.8 6.8-6.8 9.2-9.3 5.9-6.4 9.4-9.1 7.0-6.8 6.8-6.8 6.8-6.8 9.4-9.1 6.0-6.4 9.4-9.4 7.0-6.8 6.8-6.8 6.8-6.8 8.7-8.2 6.3-1.8 8.0-7.8 7.0-b.8 6.8-b.8 6.8-6.8 1.5-1.4 9.4-9.3 1.5-1.5 5.7-6.1 8.4-8.0 5.0-6.0 7.8-6.3 7.8-6.7 6.4-6.6 5.4-7.3 7.9-7.2 8.1-7.6 b.0-6.2 9.0-9.0 1.5-1.6 8.8-8.8 1.5-1.5 1.6-1.4 8.7-8.6 8.0-8.0 1.b-1.5 3.0-3.0 8.7-8.8 9.1-9.2 1.6-1.5 8.8-9.1 1.5-1.6 1.8-1.5 8.7-8.7 7.8-8.0 1.5-1.5 3.0-6.1 5.4-7.4 6.0-5.? 6.5-7.5 5.1-5.3 4.9-5.7 8.0-7.2 6.6-7.4 7.8-7.4 4.6-5.? 8.2-8.2 i.l-9.1 8.9-8.8 1.4-1.4 1.6-1.4 10.1-9.8 8.9-8.0 2.9-3.0 U -U 7 4 f 2.8-3.0 9.1-8.9 6.0-6.1 9.1-9.0 2.8-2.8 9.5-8.8 3.0-6.0 2.8-2.9 L 1 - I z.7-5.4 8.0-7.2 7.3-7.2 7.1-7.0 7.5-7.9 6.5-7.2 5.8-6.5 7.8-7.1 7 '. 9.0-3.6 3.1-3.0 8.9-9.0 3.0-3.1 9.0-8.9 3.0-3.0 9.3-9.3 10.2-10.0 9.C)-9.1 tJ S J. ' J, ,. 8.8-8.7 8.3-7.1 6.l-.0 4.-5.S 8.9-8.9 3.0-6.1 8.9-9.0 3.0-6.0 9.6-9.4 10.0-10.0 6.4-7.8 5.1-6.4 8.6-7.9 7.5-6.7 6.4-7.3 9.2-8.4 TPJ3T4E B (continued) Experiment Number 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 12 in. 9.7-9.5 8.7-8.8 4.7-4.7 8.7-8.8 4.7-4.6 9.5-d.9 2.0-6.1 4.-4.6 9.0-9.1 10.0-10.0 4.7-4.5 10.2-10.2 4.3-4.6 9.6-10.2 9.3-9.4 6.0-6.2 10.0-10.0 6.2-6.0 9.8-9.6 6.1-6.1 9.3-9.6 5.0-6.0 r/930 Channel C 3 in. 9.5-9.4 8.7-8.8 4.7-4.6 8.8-8.8 4.7-4.6 9.5-.3 3.0-3.0 4.5-4.6 9.0-9.0 10.0-10.0 4.6-4.5 10.2-10.0 4.6-4.4 9.6-10.2 9.5-9.4 8.2-6.0 10.2-10.0 6.2-6.0 9.7-9.6 6.2-6.1 9.6-9.6 6.0-6.0 7.9-8.0 8.4-8.3 8.5-8.8 7.3-7.6 134 8.4-8.3 8.5-8.5 7.6-7.6 135 7r7_79 7779 136 7.5-7.5 7.5-7.5 132 133 3in.0. 12 in. 8.4-8.5 7.4-7.2 3.1-3.0 4.4-4.5 8.8-9.0 4.6-4.8 8.4-8.7 Channel D 3 in. b.8-6.0 9.8-10.0 5.9-6.0 3.3-3.0 4.6-4.7 9.1-9,1 4.7-4.6 6.6-8.7 .0-3.0 9.3-9.4 9.0-9.0 4.7-4.7 4.0-4.0 10.0-9.4 4.5-4.7 9.b-9.6 4.6-4.8 6.0-6.0 9.8-10.0 6.0-8.0 8.6-8.6 8.7-8.6 7.9-8.0 8.4-8.3 8.5-8.5 7.3-7.t 6.1-6.0 10.4-10.2 6.1-6.0 9.2-9.? 7.9-8.0 8.4-8.6 8.5-8.5 7.3-7.6 6.1-5.8 10.0-10.0 6.4-8.1 9.6-9.5 7.9-8.0 8.4-6. 8.5-6.o 7.3-7.6 7.5-7.5 7.6-7.5 6.09.1 8.2_r/.9 7.0-7.2 6.5-7.2 5.8-8.6 7.6-7.6 7.6-7.6 9.0-8.6 9.6-8.8 8.4-8.4 8.5-8.9 9.6-10.0 7.7-7.6 7.6-8.4 8.0-7.8 6.9-7.8 8.0-8.0 7.9-8.() 7779 .0-$.O 9.3-9.6 9.0-8.7 4.6-4.8 4.6-4.o 8.5-9. 4.5-4.6 10.0-10.0 4.-4.5 979 7779 77_79 7.5-7.5 3 in.0. 6.2-7.9 5.8-6.9 6.6-8.0 6.0-6.0 8.1-7.6 6.4-7.3 5.7-7.3 5.4-7.2 8.9-8.8 8.0-8.1 9.2-9,6 9.2-9.6 6_rI 6 7.8-8.2 6.7-7.2 8.8-8.1 6.7-7.5 8.0-8.6 8.5-7.5 8.7-8.6 7.9-8.0 3.4-6.3 ?.-7.6 r/7V/ 7.5-7.5 TABLE B (continued) Experiirimt Nuber 12 in. 137 '7.6-7.9 138 139 140 7.4-7.4 7.7-7.7 7.9-7.9 Bluegill Sunfish 141 1.5-1.5 142 8.8-9.0 143 1.5-1.5 144 8.7-8.9 9.2-9.2 145 1.5-1.4 146 9.0-8.9 147 148 1.5-1.4 2.9-3.0 149 3.0-3.1 150 9.1-9.0 151 Channel C 3 in. 7.6-7.9 7.4-7.4 r7777 7.9-7.9 1.5-1.5 1.0-9.0 1.5-1.5 9.0-8.9 9.2-9.0 1.5-1.5 8.8-.7 1.4-1.4 2.°-6.0 3.0-3.1 9.1-9.0 I ..J . £ t) . J 153 154 155 156 157 158 159 160 161 162 3.1-3.0 3.0-2.7 4.6-4.6 .3-9.5 10.0-9.8 9.6-10.2 10.0-10.2 4.4-4.6 6.0-5.8 9.4-9.1 t) I 4. 0 *1 3.1-3.0 2.9-2.8 4.6-4.6 9.4-9.5 10.0-9.8 9.6-10.2 9.7-9.7 4.4-4.6 6.0-6.0 9.4-9.5 Channel D 3 in.0. 7.6-7.9 74-'7.4 7.7-7.7 7.9-7.9 6.1-7.3 6.0-7.2 5.8-6.6 5.2-6.5 6.0-7.8 6.0-6.6 7.6-7.4 6.4-6.6 7.6-6.2 6.4-6.4 7.2-6.2 'i 1 I. U L) 5.5-6.2 6.7-7.5 6.5-7.1 8.4-7.4 8.6-8.4 9.6-10.0 8.6-8.4 8.2-8.6 7.4-7.4 6.2-7.8 12 in. 77 7,5...75 7.7-'7.7 8.0-3.0 8.8-9.9 1.7-1.6 9.2-9. 1.6-1.6 1.4-1.6 9.0-8.6 1.5-1.5 8.7-3.6 9.8-0.5 8.8-0.1 3.1-3.0 r L a I C) '. 9.1-8. 9.0-9.0 9.5-9.3 4.6-4.4 4.6-4.? 4.6-4.5 4.6-4.5 10.4-10.2 9.1-9.1 6.0-6.0 3 in. 3 in.0. 7/9 7..75 7.6-7.9 7.7-7.7 8.0-8.0 7.7-7.7 8.0-8.0 8.8-8.9 7.7-7.7 3.8-5.6 1.-1.6 9.1-3.9 1.8-1.6 1.6-1.6 8.8-3.9 1.6-1.5 8.7-8.7 9.5-9,5 9.0-9.2 3.0-2.9 C r J. I L S 5.d7.l 4.0-4.6 6.5-7.2 7.0-6.1 4.9-6.6 8.0-7.2 8.8-7,5 8.2-6.8 4.5-5.7 C 9.1-b. 8.8-8.9 9.4-9.3 4.6-4.6 4.6-4.6 4.6-4.5 4.7-4.6 10.4-10.3 9.0-9.0 6.1-5.9 I !.J 5 6.5-7.9 8.4-8.3 8.4-8.0 5.7-5.8 7.3-7.4 9.2-9.6 8.7-8.3 9.0-8.8 8.2-7.8 6.5-8.2 TABLE B (continued) Chnne1 0 Expe r .ment Numbcr 163 164 165 166 167 168 12 ifl. 1O.O-i.8 6.2-6.3 9.6-9.3 ------- O}anc 1 D 3 ira. 3 in.. 12 In. 10.2-9.8 8.7-8.0 6.06.1 9.6-.6 7.68.3 7.6-8.5 6.0-6.1 9.3-9.5 6.0-6.1 ------- ------ ------- 3 in. 3 In.0. 6.0-0.0 9.cs9.8 6.2-6.0 7.3-7.4 8.8-7.4 8.6-8.0 TABLE C NUMBJR OF ENTRIES INTO CHANNELS AiD NUMBER OF Fi&. COUNTED AT 60-SECOND INTEhYALS Exp. No. Date Entries First Marker Cant. ±xp. Cant. Exp. Chinook Salmon 1 326 6/22 2 291 6/22 3 285 6/25 4 360 6/2? 5 152 7/19 6 530 7/20 7 64 7/23 8 18 7/23 9 256 6/19 292 10 6/19 215 11 6/21 376 12 6/21 132 13 7/18 396 14 7/19 76 7/24 15 214 16 7/24 308 17 7/16 234 18 7/17 91 19 7/25 122 20 7/25 246 21 6/20 51 22 6/20 134 23 6/28 131 24 6/29 346 25 7/14 20 45 60 51 28 95 8 2 98 36 34 62 13 56 20 23 66 19 16 35 17 36 82 4 13 94 37 80 22 22 21 39 16 90 80 32 62 31 87 54 36 81 64 119 49 69 85 69 57 26 29 46 73 24 10 93 3 5 Entries Second Marker Cant. Exp. Cant. Exp. 30? 266 250 297 141 530 53 15 194 66 124 210 257 160 26 330 124 325 48 23 47 8 170 93 36 50 179 296 196 161 157 50 148 262 22? 55 79 174 34 103 107 296 8 20 18 6 0 2 2 0 67 11 21 20 2 3 2 1 33 50 11 12 67 35 66 55 38 13 Time Cont. Exp. Cont. Exp. 8 178 198 136 220 520 260 114 174 123 368 90 128 427 225 24 8 15 97 1 65 46 119 70 16 37 107 263 167 124 123 140 149 104 397 151 108 6 6 2 8 7 29 35 80 4 10 88 10 153 29 36 20 81 10 22 16 7 16 19 42 32 70 22 48 2 5 0 0 19 1 70 45 168 3 7 12 3 1 14 2 0 26 20 17 4 13 24 0 1 7 30 246 6 45 36 23 17 19 0 20 3 6 6 4 7 5 1 44 42 69 99 43 34 21 1 96 41 16 28 14 24 153 52 8 1 1 37 44 14 19 44 7 17 3 43 35 101 97 36 19 19 34 19 144 64 12 52 a 66 to TABLE C (continued) Exp. No. Date 26 27 7/16 7/26 9/29 9/29 10/1 10/2 11/11 11/11 11/16 11/16 10/3 10/4 10/5 10/6 11/5 11/7 11/7 11/8 10/6 10/9 10/9 10/11 10/24 11/2 11/3 11/3 10/12 10/15 10/16 28 29 30 31 32 33 34 35 36 3? 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 Entries First Marker nt 124 168 472 438 306 xp.Cont.}xo, 45 56 169 127 130 49 58 170 87 157 411 86 72 305 258 98 95 93 105 124 64 131 79 109 116 61 136 74 56 83 227 318 367 293 382 353 260 260 271 277 224 230 342 316 300 173 274 201 358 363 122 97 146 59 72 85 92 68 66 '79 68 119 96 8? 126 99 79 61 62 43 57 65 52 159 135 '78 69 47 56 134 104 82 183 '77 208 111 Entries aecox. Marker Cont. Exp. Cont. Exp. 108 114 167 43 34 7 39 31 109 61 8 4 126 39 26 32 35 39 67 78 35 58 91 41 117 72 33 113 14 34 37 33 17 201 351 346 210 348 252 215 183 248 311 255 313 291 201 204 234 208 187 157 234 282 241 144 235 160 234 268 173 265 303 56 31 94 101 399 87 47 185 170 93 207 65 110 149 191 198 169 164 158 124 226 360 353 329 229 133 205 3 21 24 51 14 28 35 30 38 34 74 64 42 35 25 56 44 41 65 84 64 41 35 48 33 43 91 55 51 7 29 66 123 93 79 106 108 78 143 322 256 248 154 107 164 161 142 159 218 327 Time Cont. Exp. Corit. Exp. 36 45 149 173 188 126 218 190 143 128 151 195 134 148 123 46 21 138 100 158 135 17 95 91 82 116 156 94 165 147 1'73 164 46 148 63 9 50 48 39 42 4 63 6 22 25 34 29 4 21 26 29 35 68 12 13 36 36 35 26 55 12 21 18 27 13 12 12 19 20 41 18 20 15 22 30 8 59 50 40 45 35 37 22 34 29 33 37 21 16 13 35 30 34 50 39 20 28 36 68 25 35 82 45 69 70 127 90 79 92 41 102 109 127 93 129 204 C.' TABLE C (continued) Exp. No. Date 55 10/16 10/23 10/23 10/24 10/26 10/29 11/2 11/17 11/17 11/17 11/17 11/19 11/19 11/20 56 57 58 59 60 61 62 63 64 65 66 67 68 Entries First iarker Cent. X. Cont. Exp. 271 142 214 200 275 178 243 170 161 188 174 276 165 132 169 196 57 125 128 57 180 129 134 50 55 36 60 32 43 14 13 37 47 89 109 166 123 60 116 253 124 208 137 168 205 150 320 352 432 294 187 378 219 389 316 193 370 77 53 83 55 89 62 53 79 50 55 49 38 44 53 84 64 259 276 72 179 153 142 134 159 80 68 68 68 69 44 197 365 87 70 379 487 368 233 447 310 483 352 243 440 83 91 84 Entries Second 1arker Cont. Exp. Gont.Exp. 62 159 156 234 143 194 169 154 66 106 27 46 17 29 42 43 46 26 35 24 28 19 39 31 35 33 19 32 185 150 133 143 160 230 121 78 99 113 80 97 47 93 Time (iont.xp 129 123 25 26 24 28 91 111 154 62 132 125 101 80 90 126 113 52 40 32 19 15 13 59 21 8 8 33 4 15 7 63 22 47 22 16 44 32 27 29 49 Silver Salmon 69 70 71 72 73 74 '75 76 7? 78 79 80 7/7 7/9 '7/9 7/10 7/6 7/6 7/10 7/11 7/11 7/5 7/5 7/12 96 80 81 125 92 140 139 43 81 85 80 91 79 99 140 113 117 114 49 71 226 318 192 240 270 271 34 32 39 50 73 68 71 72 38 54 97 26 34 58 99 73 26 16 26 29 36 76 37 58 112 5 94 96 99 150 177 137 86 156 167 209 142 166 129 111 20? 215 Corit. Exp. 14 20 8 40 33 44 32 50 39 50 34 27 121 131 81 125 84 184 92 78 63 135 92 138 57 108 65 13 18 21 27 37 19 38 11 50 30 3? 47 13 10 168 143 29 44 38 38 25 76 74 13 13 18 50 103 38 6 TABLE C (continued) Exp. No, Date 81 82 83 84 85 86 87 88 89 7/12 7/2 7/3 7/5 7/13 7/13 10/17 90 8/3 'tri_F1rst_Marker Entries Second Marker 420 292 270 255 324 333 254 233 Cent. 1xp. Cont, Lxp. 266 256 266 10/17 160 10/17 Largemouth Bass 91 92 93 94 95 96 97 98 9 100 101 102 103 104 105 106 10? 8/4 8/6 8/6 3/20 8/21 9/7 9/8 8/3 8/7 8/8 8/8 8/16 8/16 8/22 8/23 8/29 8/29 27 41 242 108 143 168 10 123 34 15'l 116 45 118 10 68 141 58 98 117 58 55 50 95 112 42 83 170 50 65 53 76 110 39 '79 64 40 15 17 3? 36 55 66 6i 17 20 47 30 43 74 62 64 43 13 66 43 45 62 61 56 31 19 64 35 48 255 99 222 245 270 220 187 204 299 19 22 48 67 94 33 13 88 37 72 77 127 44 71 74 162 68 144 13 109 53 137 64 50 42 108 Cont. Exp.Cont. Exp. 37 58 26 224 269 211 181 206 31 46 3? 30 50 56 19 2 2 123 20 96 58 10 13 86 20 3 90 7 89 14 52 58 16 73 89 2'? 118 37 82 3 18 10 5 38 12 23 20 10 28 17 14 Time Cont. xp. Corit. 1xp. 147 82 59 166 41 196 54 225 84 171 21 117 46 132 18 232 102 202 12" 17 127 124 91 127 6 81 60 9 101 67 107 89 117 76 5 8 20 7 72 42 9 18 21 15 6 9 27 24 49 23 24 8 51 17 19 19 10 35 49 20 40 36 4 26 29 22 1 92 11 97 '74 52 93 41 60 162 56 378 76 61 177 97 71 110 70 91 66 61 29 14 16 15 26 39 15 2 8 10? 35 29 111 50 31 25 27 44 65 178 37 18 15 14 33 41 48 40 18 87 40 72 74 41 35 127 194 76 97 134 103 70 52 22 105 123 86 50 16 42 44 81 130 95 83 145 13 6 27 14 23 19 7 49 58 62 130 39 103 75 79 10? 97 141 TABLE C (continued) Exp. 108 109 110 111 112 113 114 115 116 11? 118 119 120 121 122 123 124 125 126 12? 128 129 130 131 132 133 134 135 1ate 9/21 9/24 8/8 8/9 8/9 8/9 8/30 8/30 8/30 8/31 9/19 9/19 9/20 9/20 9/20 9/13 9/14 9/15 9/15 9/15 9/17 9/17 9/1? 8/15 8/15 8/29 8/31 8/31 Entries First Marker Cont. xp. Cont. Exp. 220 194 148 165 78 84 58 98 224 125 266 196 139 21? 37? 124 166 355 243 222 158 148 192 264 127 100 120 103 63 48 61 54 48 61 43 45 56 55 31 59 44 53 113 60 58 58 47 22 48 29 40 31 28 86 60 61 54 39 58 85 51 71 50 42 48 60 41 41 43 55 88 83 42 64 36 20 47 2 28 39 42 80 32 54 182 111 124 182 Cont. Tixp. Cont. 144 118 82 103 66 51 70 64 108 70 100 239 129 49 37 82 15? 69 271 169 163 108 260 163 160 220 254 236 23? 226 139 124 82 109 Time Entries Second Marker 145 126 77 133 307 96 104 182 161 144 124 72 133 111 79 74 92 77 44 36 24 26 22 23 4? 23 13 22 21 17 14 31 37 16 51 28 42 102 3? 40 47 36 19 46 24 31 4 9 47 40 43 25 29 22 31 42 35 36 35 34 78 56 24 56 26 17 33 18 19 11 21 41 21 34 xp. 92 76 54 104 26 13 41 60 48 82 133 68 157 123 155 76 165 125 104 138 141 128 154 130 70 85 57 77 Cnt. 112 101 14? 125 78 100 66 61 143 202 108 60 59 91 143 131 114 99 101 126 80 55 84 208 79 98 140 80 xp. Corit. ixp. 70 33 100 36 63 77 65 178 120 137 40 63 35 63 50 81 70 33 52 22 82 14 43 12 24 133 145 165 46 109 223 116 161 96 123 84 83 59 137 103 56 97 93 115 48 141 43 95 72 68 24 46 83 161 86 51 121 56 64 87 20 4? 29 22 40 105 70 65 113 98 9? 91 132 87 105 12 133 105 66 64 163 94 91 8? TABLE C (continued) Exp. Date No. 136 137 138 139 140 9/12 9/12 9/13 9/13 9/13 Entries First Marker Cont.}xp. Gont. 1xp. 113 51 33 210 61 80 92 133 140 151 82 76 52 56 79 Bluegill Sunfish 147 141 8/10 142 153 8/11 70 143 8/11 144 150 8/11 219 145 8/17 74 146 8/17 283 147 8/20 148 168 8/21 31 149 8/2 77 150 8/13 96 151 8/13 72 152 8/13 '74 153 8/16 154 lüS 8/23 64 155 8/2 64 156 8/2 196 157 9/19 377 158 9/20 238 159 9/26 157 160 9/27 137 '75 161 7/26 56 65 69 44 64 69 95 76 72 38 70 17 17 50 50 61 58 60 52 45 74 113 79 40 114 114 100 125 117 Entries Second Marker Cont. Exp. (oat. Exp. 102 34 24 83 43 73 165 82 99 124 127 101 75 127 31 99 78 29 86 62 56 59 81 31 125 102 38 70 38 54 68 59 87 '77 84 83 25 142 63 65 228 128 44 51 49 67 260 163 288 341 83 88 42 63 '78 56 86 -16 34 21 64 67 42 69 118 59 3 87 9 20 0 30 23 44 50 19 12 29 3? 8 12 10 27 2 3 -2 3 12 19 13 26 16 36 13 10 22 193 307 204 143 64 6 2 63 102 55 2'? 31 -- 36 0 8 29 56 9 65 76 52 61 93 67 85 52 47 58 43 13 47 22 90 2 5 54 18 34 42 23 187 114 92 114 128 23 -- 9 6 7 78 Time Cont. ±xp. Oont. Exp. 107 102 130 136 89 48 136 103 127 132 123 97 77 136 134 104 147 135 110 79 9 29 16 225 155 231 283 113 143 85 49 31 84 10 69 26 7 115 145 13 40 15 77 73 58 22 17 2 4 27 23 34 67 30 33 43 35 16 58 50 27 58 67 65 54 46 50 45 64 18 45 17 93 57 51 15 20 27 10 20 28 26 19 6 105 26 25 81 52 33 44 120 70 189 150 72 TAI3L Exp. 162 163 164 165 166 167 168 Date 8/i 9/18 9/18 9/iS 8/15 8/15 8/is Entries First Marker Cant. ixp. Coot. xp. 82 300 274 2176 224 73 116 60 47 73 89 100 54 77 72 35 60 71 92 82 40 114 269 303 165 143 120 47 C (continued) Entris Second Marker Cort. ixp. Cont. rxp. 24 291 238 216 83 23 23 17 45 12 33 57 44 63 51 4 5 67 264 254 97 28 16 1 2 11 7 26 Time ont. kTxp. Cent. Lxii. 85 80 102 110 111 95 114 100 177 66 40 37 52 43 54 103 58 31 32 174 51 53 77 66 78 81 59 46