TECHNICAL REPORT NO.

1969

FISHERIES RESEARCH BOARD OF CANADA

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FISHERIES RESEARCH BOARO OF CA~AOA

TECHNICAL REPORT NO. 148

SURVIVAL ANO GROIITH OF ATLANTIC SALMON FRY

I~ RELATION TO SALINITY ANO OIET

BY

R. L. Saunders and E. B. Henderson

This is the twenty-seventh Technical Report from the

Fisheries Research Board of Canada,

Biol09ica1 Station, St. Andrews, N. B.

1969

SURVIVAL AND GROWTH OF ATLANTIC SALMON FRY

IN RELATION TO SALINITY AND DIET

By R. L. Saunders and E. B. HendersonFisheries Research Board of Canada

Biological Station, St. Andrews, N. B.

INTROOUCTION

Atlantic salmon (SaZmo BaZar) are spawned in streamswhere the young hatch and remain for two or more years beforedescending as smo1ts to the sea. The smo1t migration representsa major transition from river to sea life with its attendantchanges in salinity, living space, and food supply. Smo1ts areable to withstand the increasing salinities encountered in theestuary and in the sea (Jones, 1947; Koch et a1., 1959; Parry,1961). Salinity tolerance of parr is less than that of smo1tsand varies directly with size (Hoar, 1937; Parry, 1958;Koro1eva, 1960; and Saunders and Henderson, 1969). Even salmona1evins and fry can tolerate dilute salt water (Hoar, 1937;Parry, 1958, 1960, 1966; Koro1eva, 1960; and Sydor, 1966).Saunders (1966) reports spawning of adult salmon in a tidalestuary and cites unpublished data of the late Dr. A. H. Leimwho successfully incubated Atlantic salmon eggs, which had beenfertilized in fresh water, in salinities of 3, 6, 9, and 12~.

The present experiments were designed to establishthe salinity tolerance levels of salmon a1evins and to studythe effect of salinity on the 9rowth rate. The experimentsalso gave an opportunity ~o study the beneficial effects ofsalt water in controlling fungus infections and the harmfuleffects of a diet of herring which ~ontained the vitamin­destroyin9 enzyme, thiaminase (Nei1'nds, 1947).

MATERIALS AND METHODS

Studies of salinity tolerance were made on hatcheryreared salmon fry just fo110win9 completion of yolk-sacabsorption. Details of the experiments are 9iven in Sydor(1966) .

Growth studies were done in two instalments. fromMay 1964 to June 1965 and from May 1965 to June 1966. Thesecond instalment served as a replicate of certain parts ofthe first and provided opportunity to test preliminaryconclusions.

2.

FIRST EXPERIMENT, 1964-1965

Salmon a1evins were obtained in May 1964 fromSaint John Fish Culture Station and held in fiber91ass tankswith limited exchange of fresh water so that the fish wouldbe quiescent. They were held in fresh water until they hadcompleted absorption of their yolk sacs and be9un to eat.In June four groups of approximately 148 fry each were placedin separate tanks of dilute salt water; a fifth group washeld in fresh water. Salinities for the saltwater groups weregradually increased to 2, 4. 8, and 12~ over 4 days and weremaintained within ±1~ by mixing fresh and salt water. Watertemperature varied seasonally but at any time was the same forall groups.

Fry were fed from June to September 1964 a mixtureof equal parts beef liver and fresh Atlantic herring (ClupeaharenguB) ground to a paste and mixed with a small amount ofPurina trout chow. The fish were given as much food as theywould eat at each feeding. Feedings were three or four timesa day during the summer but were gradually reduced to once aday as water temperature fell. Herring was excluded from thediet after September 16 because it was thought to be responsiblefor sickness and death of some fish. Herring containssignificant amounts of thiaminase (Neilands, 1947).

The fry were first measured at the end of July 1964and seven times subsequently, the final measurement on June 8,1965. They were not fed on the day they were measured. Fishwere anesthetized in a solution of tertiary amyl alcohol ofsuch strength (5-10 ml/~) that they were immobilized quickly(2-3 min). Higher concentrations of anesthetic were requiredwhen the water was cold. Only about 6 fish were anesthetizedat a time. Each was measured to the nearest millimeter(fork 1en9th) and weighed to the nearest 0.1 gram. They werequickly returned to clean water where recovery to thebreathing stage was usually made within 5 minutes.

SECOND EXPERIMENT, 1965-1966

Dn May 14, 3,373 fry were placed in running freshwater in a cylindrical fiberglass tank, 3 ft in diameter, tostudy the mortality rate before and after feeding began.Feeding was started on June 15 with 9round beef liver. Dailyrecords of mortalities were made.

Dn July 5 the surviving fish (about 1,800) weredivided randomly into six groups of 30D. Two groups wereput in fresh water. two in ~ sea water. and two in 1~.From June 15 to August 24 all fish were fed beef liver.From August 24 to March 4 three groups comprising Lot II(fresh water, 6~, and 12~) were fed a mixture of 50% herringand 5D% beef liver by weight, and the other three groups(Lot I) were fed beef liver only.

3.

Owing to a higher mortality rate in the freshwatergroups, there were fewer fish left by August 17 (the firstmeasuring date) than in salt water. Accordingly, all lotswere reduced to 100 fish on this date. A random samplingtechnique was used to avoid bias in selecting fish to keep.

Data processing for both experiments was done bycomputer analysis using' the iBM 1130 ·system. The computerprogram was written to provide: (a) mean length, weight, andcondition factor with standard error of the mean; (b) comparisonof batches with standard error of the difference between meansand "t" values; and instantaneous and linear growth rates!for length and weight.

RESULTS AND DISCUSSION

Salinity tolerance

Fry survived indefinitely following abrupt transferfrom fresh to brackish water up to 1~. Within a salinityrange of 0-12~ and at 10-12 C there were no differences inmortality rate among groups (fresh water, 2, 4, 8, and 12~)attributable to salinity. Fry transferred directly fromfresh water to 15~ were obviously weak after 1 day and weredead or dying after 2 days. Prior acclimation to brackishwater (7~) prolonged survival of fry in 15~ but did not leadto indefinite survival in 15~. Fry transferred from freshwater to 22 or 3~ died .within 24 hours. It is concluded thatthe lethal level for fry is between 12 and 15~. Accordingly,the growth experiments were conducted with salinities of 12~and lower.

j964~1965 Experiment. There were no obvious adverseeffects of dilute salt water on the fry. On the contrary,it was soon obvious that fewer fish were dying in the saltwatergroups than in the freshwater one. Apparently salt water ata salinity as low as ~ prevented or reduced growth of funguswhich often accumulated on uneaten food and on fish in freshwater.

where Y ;s mean

Mortality ratesinitial feeding, June and

lInstantaneous growth rate

and linear growth rate

were highest during the period ofJuly, when it was inevitable that

loge YT - loge YtT -

~T - t

length or weight at times T and t, T being later than t.

4.

some food was uneaten and accumulated on the tank bottoms.Daily records of fish deaths were not kept but when the fishwere measured for the first time, July 29, there were fewerfry remaining in fresh water than in any of the other groups(Table 1).

During August there were few mortalities and nodifference in mortalities among groups. In September themortality rate increased, there being more deaths in thesaltwater groups than in the freshwater group. It wasthrought that the herring diet was responsible. Accordingly,herring was excluded from the diet thereafter. There werefew deaths between mid-October and the end of the experiment.

Mean lengths and weights were similar among groupsat each of the eight measuring dates from July 1964 to August1965 (Table 1). Towards the end of the experiment the fishin 12~ salinity were slightly heavier though no longer and there­fore had a higher mean condition factor than the other groups(Tab1 e 1).

It was meaningless to compare the growth rates amonggroups because of the divergence in numbers of fish followingdifferential mortalities among groups. It was decided,therefore, while the present experiment was still in progress,that we should repeat parts of the experiment with carefulattention to keeping equal numbers of fish among groups.In addition to studying further the possible effect ofsalinity on growth rate, we designed the subsequent experimentto test the hypotheses that dilute salt water reduces mortalityrates among fry through its therapeutic effect in reducingfungus infection and that a herring diet results in sicknessand death in salmon.

1965-1966 Experiment. Of the 3,373 fry placed in freshwater on May 14, only 18 had died prior to commencement offeeding on June 15. Mortalities increased after feeding wasstarted, apparently because of fungus growth which wasenhanced by accumulation of uneaten food. By July 5, 45% ofthe fry had died.

The surviving fish were divided into six groups of300 and put in water of different salinities on July 5.Further observations were made to relate mortality rates tosalinity. Between July 5 and August 17 mortality rates inthe various salinities were as follows:

/

~ .

Fresh water6~ sal inity12~ salinity

Lot I-.l!L~6.9

27.025.1

Lot II-.1!.L

60.019.628.3

Mortality rates in the freshwater groups in each lot weresignificantly higher than those in the saltwater groups.

Ilcasurelllents of grollth I'/ere begun on August 17 and onAugust 24 Lot II groups started receiving the herring-liverdiet. TIIere were fe'" mortalities in any group between August 17and October 29. Ouring November there was an upsur~e in themortality rate among the herring-fed groups (Lot II). Thecomparison of mortality rates between the herring- and liver-fedlots between October 29 and March 4 is as follows:

Fresh water6~ salinity

12)\. sal i nity

Lot I (liverdi et) (%)

3o2

Lot II (herrin~­

liver diet) (%1

456373

The inescapable conclusion was that inclusion ofherring in the diet of fry for a period over 3 months resultedin heavy mortality. In view of the similar, low mortalityrates among salinity groups in Lot I, it appears that increasedsalinity aggravated the adverse effect of the herring diet inLot ".

Mean lengths and weights were essentially the sameamong groups when the fish were measured for the first time(Table 2, Fig. 1 and 2). There was no difference in instan­taneous and linear growth rates between 1iver- and herring-fedgroups between August 17 and October 26 (Fig. 1 and 2,Appendix 1). Between October 26 and November 24, growth ratesof the herring-fed groups fell well below those of thecomparable liver-fed groups. This was a period of heavymortality in the herring-fed groups. Further comparisons ofgrowth between Lots I and II would be meaningless because ofdifferential mortality rates. Accordingly, the herring-fedfish were not measured after November 24.

In the liver-fed groups (Lot I) the fish in 6)\.salinity were significantly larger (P < 0.01) than the fresh­water fish on each measuring date subsequent to the first one(Fig. 1, Appendix 2). Although the fish in 12)\. were alwayslarger than those in fresh water, differences were small andnot statistically significant. The fish in 6)\. were alwayslarger than the fish in 12)\. but not significantly so(P < 0.01) until the last measuring date.

Condition factors were generally highest in the 12~group and lowest in the freshwater group (Table 2). The

6.

fastest-growing fish (6~) had condition factors intermediatebetween those of the freshwater and 12~ groups during mostof the experiment.

Our results were generally similar to those ofCanagaratnam (1959) who demonstrated that coho salmon fry(Oncorhynchus kisutch) grew faster in brackish than in freshwater. However, his results differ from ours in that cohofry survived in 1B~ and growth rate was directly related tosalinities up to l~ , the highest salinity used.

CONCLUSIONS

1. Young salmon, after absorption of the yolk sac. livedindefinitely in salinities up to 12~ but died in 15~.Acclimation at 7~ increased the survival time in l5~.

2. Salinities between 2 and 1~ were effective in reducingmortalities associated with fungus which in fresh watercaused heavy mortalities.

3. Inclusion of uncooked herring in the diet of salmon fryfor more than 3 months resulted in heavy mortality ­probably owing to the high content of thiaminase inherring.

4. Growth rates of salmon fry in dilute salt water werehigher than those of fry in fresh water. Growth ratesof fry in water of ~ salinity were significantly higherthan those of fry in fresh water and slightly higherthan those of fry in water of 1~ salinity.

ACKNOWLEDGMENTS

We thank Misses Mary Holmes and Judith Sydor andMr. John Byrne who helped in various ways in this study.

Drs. M. W. Smith and P. F. Elson and Mr. K. R. Allenoffered suggestions for conduct of the experiments and analysisof data. Fish were provided by the Saint John Fish CultureStation through the Resource Development Branch of the CanadaDepartment of Fisheries.

r-Ir. K. R. Allen, assisted by Mr. J. K. Lindsey,prepared the program for computer processing of the data.Coding and processing were done by Mr. G. Fawkes and Mrs. ThoraDempsey, under the direction of Mr. A. Sreedharan.

7.

REFERENCES

Canagaratnam, P. 1959. Growth of fishes in differentsalinities. J. Fish. Res. Bd. Canada 16(1): 121-130.

Hoar, II. S. 1937. 1·largaree salmon investigation. Vol. II,1937. The reactions of young salmon to salt water.Bio1. Bd. Canada, /-IS Rept. Bio1. Sta., No. 127. 11 p,

Jones, J. N. 1947. Salmon smolts and sea water. Survivalof smolts of SaZmo saZar at various rates of increasesof salinity. Salmon and Trout ~agazine, No. 119, p. 63­76.

Koch, H. J., J. C. Evans, and E. Bergstrom. 1959. Sodiumregulation in the blood of parr and smo1t stages of theAtlantic salmon. Nature 184(4682): 283.

Koroleva, N. V. 1960. Tolerance of young salmon (Sa~mo8a~ar) to various degrees of water salinity. Iiauch.­Tekh. Byu1. Gos. Nauch.-Iss1. Ozern. Rech. Rybn. Khoz12: 33-34.

Iieilands, J. B. 1947. Thiaminase in aquatic animals offlova Scotia. J. Fish. Res. Bd. Canada 7(2): 94-99.

Parry, G. 195B. Size and osmoregulation in sa1monid fishes.Nature 161(4617): 1218-1219.

Parry, G. 1960. The development of salinity tolerance inthe salmon. Salma BaZar (L.). and some related species.J. Exp. BioI. 37(2): 425-434.

Parry, G. 1961. Osmotic and ionic changes in blood andmuscle of migratin9 salmonids. J. Exp. Biol. 38(2):411 -427.

Parry, G. 1966. Osmotic adaptation in fishes. BioI. Rev.41: 392-444.

Saunders, J. W. 1966. Estuarine spawnin9 of Atlantic salmon.J. Fish. Res. Bd. Canada 23(11): 1803-1B04.

Saunders, R. L., and E. B. Henderson. 1969. Survival andgrowth of Atlantic salmon parr in relation to salinity.Fish. Res. Bd. Canada Tech. Rept. 110. 147.

Sydor, Judith. 1966. Effects of salinity on salmon fry.Summer, 1964. Fish. Res. Bd. Canada, MS Rept. (Bio1.),No. B99. 10 p.

18

16 u0

14 ..12 :J

0

10 ..0-

E8 ..

~

14 .6,.

,,13 ' . 4"/ .

Fresh Water .'~E 12 ," ./ . 33u 6 0/•• Salinity " ./ I" " III 12 %. ,''''' I 30

~ , " ,,'". I

'" 10 ---_.....;" ". 27 ..c:..,~9=-=--;.-;.- =:: f E

...J 09 24 '"c: ",;; Length / /0 8 21.. ,,~/., ~"./::IE .<:;

~ '/ '"7 / /~/18 ..

~6 // 15"/ c:5 . ,''/ 12 0

----==-:.:::;:::/ ..::IE

....~ -- . 9.~

Weight/ 6

.~ 3

0A S 0 N 0 J F M A M J

1965 1966

Fig. 1. Effect of salinity on mean length and weight ofsalmon fry at various measuring dates. Data for lfver­fed fish, Lot I (batches 1-3) in Table 2.

~ /#," .......~ //=<>~ 7

Fresh Water

o • 6 %.2 12°"'.

:'~

f~

,.~0

:/ ~';c0.:0

Fig. 2. Effect of salinity on mean length and weight ofsalmon fry at various measuring dates. Data forherring-liver fed fish, Lot II (batches 4-6) inTable 2.

Table 1. Mean length, weight, and condition factor of salmonfry reared at different salinities .

.,'"~E en c

" .. u °.,° '"-;:; ~~ .... '" '"., .... .... _0 ....

'" - a. -" a. c en oc c ° c c" ... oc.,~ EU E E ",c '" '" '" ",cu '"'" ,i;...a. "'0 "'" '" '" "'''' '" "'''' "'0'" "''''c .... Z~ z_ V>E zen V>E z u .... V>E

29/7/64 Fresh 16.2 106 4.3 .042 0.8 .026 .938 .00812 16.2 135 4.3 .039 0.8 .022 .950 .00574 16.2 134 4.3 .039 0.8 .024 .958 .00778 16.2 156 4.3 .035 0.8 .022 .970 ,0059

12 16.2 117 4.4 .038 0.9 .027 1.038 ,0079

31/8/64 Fresh 15.7 98 6.0 .072 2.4 .103 1,055 .00922 15.7 132 5.9 .058 2.3 .076 1.076 .00684 15.7 131 5.9 .063 2.4 .079 1.094 .00678 15.7 152 5.5 .057 2.1 .071 1. 1gO .0091

12 15.7 117 6.0 .063 2.7 .092 1.204 .0097

3/11/64 Fresh 10.4 86 8.1 .125 6.4 ,390 .126 .00842 10.4 82 8.0 .097 6.2 .261 1.150 .00784 10.4 75 8.1 .125 6.8 .372 1.175 .00928 10.4 68 8.1 .144 6.6 .453 1. 158 .0088

12 10.4 68 8.2 .153 7.4 .474 1. 237 .0112

5/1 /65 Fresh 5.4 65 8.5 .180 7.2 .627 .055 .00962 5.4 75 8.4 .121 6.6 ,334 1.073 .00824 5.4 69 8.6 .154 7.6 .495 1.095 .00 28 5.4 67 8.4 .157 7.0 .500 1.089 .0069

12 5.4 69 8.7 .181 7.9 .614 1.088 .0077

9/3/65 Fresh 5.0 64 8.7 .192 7.8 .730 1.038 .01012 5.0 75 8.6 .133 7.4 .410 1.094 .00924 5.0 69 8.8 .172 8.5 .586 1.134 .00718 5.0 67 8.6 .166 7.8 .591 1,128 .0079

12 5.0 68 8.9 .202 8.9 .754 1. 116 .0088

8/4/65 Fresh 5.2 63 8.9 .202 8.8 .824 1.099 .00992 5.2 75 8.9 .146 8.2 ,472 1.102 .00754 5.2 68 9.2 .188 9.5 .684 1.124 .00978 5.2 67 8.8 .176 8.4 .648 1. 125 .0080

12 5.2 68 9.1 .219 9.8 .867 1.137 .0086

6/5/65 Fresh 6.0 62 9.5 .219 10.5 .955 1.103 .00822 6.0 75 9.5 .169 10.2 .610 1. 115 .00884 6.0 67 9.7 .211 11.4 .816 1.123 .00748 6.0 67 9.3 .195 10.1 .794 1. 116 .0090

12 6.0 67 9.7 .241 11.8 1 .062 1.135 .0086

8/6/65 Fresh 10.6 62 11. 1 .244 17.5 1.295 1.175 .01302 10.6 75 11.0 .202 16.7 .913 1.172 .01194 10.6 66 11. 2 .248 17.8 1.153 1.177 .01328 10.6 63 11.0 .226 16.9 1.200 1. 17 5 .0083

12 10.6 66 11. 2 .286 18.5 1.471 1.198 .0126

Table 2. Mean length, weight, and condition factor of salmon fryreared at different salinities. Groups designated "A"were fed liver (Lot I); those designated "B" were fedherring and liver (Lot II). Batch numbers correspondwith those in Appendices 1 and 2.

.,o<E

-'=~'"?; u .. u 0

" 0 0 .... -;; .... .... :5 ~ ., ..~ ~ -'= .. - .... .... _0 ....

., >, c- u c- DC- ;g' 0; ~ O~ ~" ., O~

.,"'''' ., E EE '" '" "'~u '"",0-~"'- '"

.,"'"

., ., "'., ., "'., "0'" "'.,0-., '" .... Z~ "'- V>E "'''' v>E '" u .... V>E

17/B/65 Fresh A 1 17.9 101 5.2 .050 1.5 .049 1.018 .0070(0) 6 A 2 17. g 100 5.2 .043 1.5 .040 1.024 .0057

12 A 3 17. g 100 5.1 .049 1.5 .042 1.065 .004Fresh B 4 17.9 100 5.3 .045 1.6 .045 1.039 .0063

6 B 5 17.9 101 5.1 .043 1.4 .038 0.991 .006512 B 6 17.9 100 5.0 .046 1.5 .042 1. 123 .0061

28/9/65 Fresh A 1 15.0 98 6.8 .076 3.9 .165 1. 162 .0092(42) 6 A 2 15.0 100 7.1 .058 4.4 .115 1.200 .0080

12 A 3 15.0 98 7.0 .071 4.4 .152 1.227 .0076Fresh B 4 15.0 99 7.0 .073 4.1 .134 1. 139 .0084

6 B 5 15.0 97 7.3 .073 4.5 .151 1. 150 .006712 B 6 15.0 100 7.1 .078 4.4 .161 1. 17 2 .0056

26/1 0/65 Fresh A 1 11.8 98 8.0 .102 6.4 .301 1. 168 .0085(70) 6 A 2 11.8 100 8.4 .080 7.3 .228 1.189 .0068

12 A 3 11.8 98 8.3 .094 7.1 .274 1.196 .0070Fresh B 4 11.8 99 8.2 .093 6.7 .264 1. 138 .0064

6 B 5 11 .8 97 8.3 .097 7.1 .266 1.181 .006912 B 6 11.8 100 8.1 .094 6.7 .255 1.186 .0059

24/11/65 Fresh A 1 7.B 98 B.9 .127 8.4 .441 1.0BB .0061(99) 6 A 2 7.B 100 9.4 .104 9.6 .342 1.106 .0061

12 A 3 7.8 98 9.2 .116 9.1 .399 1. 113 .0055Fresh B 4 7.8 94 8.9 .110 B.2 .351 1.090 .0061

6 B 5 7.8 95 9.0 .104 8.3 .315 1. 114 .005912 B 6 7.8 96 8.7 .113 7.9 .342 1.123 .0055

24/2/66 Fresh A 1 3.7 98 9.4 .153 9.1 .513 0.990 .0052(191 ) 6 A 2 3.7 100 9.9 .129 11.2 .452 1.077 .0059

12 A 3 3.7 98 9.6 .143 10.3 .486 1. 03 .0289

10/5/66 Fresh A 1 10.7 96 11.4 .189 17.1 .912 1.056 .0052(266) 6 A 2 10.7 99 12.4 .156 21.8 .771 1.102 .0060

12 A 3 10.7 98 11.8 .191 19.8 .912 .124 .0075

7/6/66 Fresh A 1 12.8 96 12.9 .217 25.7 1.215 1. 117 .0082(294) 6 A 2 12.8 99 14.2 .166 32.8 0.974 1. 127 .0092

12 A 3 12.8 97 13.2 .220 26.7 1. 181 1.100 .0083

Appendix 1. Instantaneous and linear growth ratesfor length and weight of salmon fry.Batches 1, 2, and 3 are for liver-fedfish in fresh water, 6 and l~respectively (Lot I); 4. 5. and 6 arefor herring-liver fed fish in freshwater, 6 and l~ respectively (Lot II).Lot II discontinued after day 99.

BATCH DAY DAY -VAL MEAN 1 MEAN 2 INST. RATE LIN. RATE MEAN 1 MEAN 2 INST. RATE UN. RATE1 o. 42. 42. 5.206 6.818 o.6419t.~ Q.3836E-QI 1.485 3.864 O.2276E-Ql Q.5664E-Ol

42. 10. 28.

10. 99. 29.

6.81~

8.018

8.016 0.5169E-02 0.4265E-01

8.928 0.3707E-02 003138E-01

3.664

6.381

6.381 0.1191£-01 0.8998£-01

8.363 0.9325E-02 a.6933E-Ol

1 99. 191. 92. 8.928 9.446 O.6134t.-03 0.5634E-02 8.363 9.089 O.9U49t.-U3 o. 1692E-02

1 191. 266. 15.

1 266. 294. 28.

9.446

11.438

11.438 0.2550E-02 0.2655E-Ol

12.923 0.4360E-02 0.5305E-Ol

9.089

170131

17.131 0.8450E-02 001072E CO

25.663 0.1443E-Ol 0.3046E 00

2 O. 42. 42. 5.190 7.091 0~1449E:~-0"2--~'or":Ol L4'10---- 4.395--~1 0.6'}65E-'H

42. 10. 28.

2 10. 99. 29.

1.091

8.397

8.397 0.6006E-02 0.4642E-Ol

9.385 0.3835E-02 0.3406E-01

4.395

7.216

7.276 0.1199£-01 0.1029E 00

9.555 0.9397E-02 0.7860E-Ol

2 99. 191. 92. 9.385 9.936 0.6200E-03 0.5989£-02 9.555 11.150 o.1677e:-02 0.1733E-01.

-S---;TfT- ---7~038 0.1587E-02 O-~4$73£"~r--------r.468 4.434 0.2631E':-01 0.7061E-01c-...

2 191. 266. 75.

2 266. 294. 28.

~2.

42. 10. 28.

10. 99. 29.

99. 191. 92.

3 191. 266. 15.

3 266. 294. 28.

o. 42. 42.

42. 10. 28.

10. 99. 29.

O. 42. 42.

42. 70. 28.

10. 99. 29.

O. 42. 42.

42. 70. 28.

10. 99. 29.

9.936

12.384

1.038

8.213

9.191

9.641

11.825

5.280

1.015

8.234

5.122

1.257

8.335

5.011

10111

8.139

12.384 0.2936E-02 0.3263E-01

14.174 0.4821E-02 0.6392E-01

8.273 0.5772E-02 0.4409E-01

9.191 0.3629E-02 0.3166£-01

9.641 0.5195£-03 0.4891E~02

11.825 0.2121£-02 0.2911E-01

13.185 0.3887E-02 0.4857E-01

7.915 0.6750E-02 0.4128E-01

8.234 0.5722E:--02 0~"'354E-Ol

8.929 0.2795E-02 0.2398E-01

7.257 0.8294E-02 0.5083E-01

8.335 0.4942E-02 0.384 7E-0 1

8.957 0.2485E-02 0.2141E-01

.11 7 0 .8352E~-O~--------O;-50T4E-0 1

8.139 0.4788::-02 0.3641E-01

8.748 O.2491E-02 0.2102£-01

11.150

21.784

4.434

1.014

9;TTi

10.339

19.828

1.514

4.063

6.653

1.311

4.547

7.103

.455

4.401

6.665

21.784 0.8929£-02 0.1417:. 00

32.806 0.1462E-Ol 0.3736:; 00

1.01to 0.1668E-Ol 0.9428::-01

9.122 O.d768E-02 0.7062E-Ol

10.339 0.1361£-02 0.1322':::-01

19.828 0.8681E-02 0.1265E 00

26.612 0.1059E-01 0.2444E 00

''-'-063- -O~5 7E-01 0.5926£-01

6.653 a.P61E-01 0.9250::-01

a.178 C.7115E-OZ 0.5258::-01

4.547 0.2854£.-01 0.7562E-01

7.103 0.1593E-01 0.9129E-01

8.341 0.5537£.-02 0.4267::-0:

4.407 o.2636!;-Ol O. 1029E-:l

6.665 O.1476E-Ol 0.3063E-01

1.913 0.5916£-02 0.4302E-01

Appendix 2. Comparison of mean length, weight, andcondition factor between batches atdifferent measuring dates. Batches 1,2, and 3 (Lot II are for liver-fed fishin fresh water, 6 and l~ respectively;4, 5, and 6 (Lot II) are. for herring-liverfed fish in fresh water, 6 and l2~ .Lot II discontinued after day 99.

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EXPT BATCH DEG. OIFFERENCF S. E. OF OIFF TDAY A 0 FREE

O. 1 2 LEN 199. 0015930[-01 O.62232E 01 0.2559WGT 199. 0015543E-Ol 0.63106E-Ol 0.2463CON 199. 0.56296E-02 0.90581E-02 0.6214

O. 1 3 LEcN 199, 0.88930E-Ol 0.66358E-Ol 103401WGT 199. 0.17143E-Ol 0.64254E-Ol 0.2668CON 199. 0.47303E-Ol 0.81216E-02 5.8243

O. 1 4 LEN 199. 0.74069E 01 0.63498E-Ol 101664WGT 199. 0.88956E-Ol 0.66085E-Ol 103460CON 199. 0.20575E-Ol 0.94456E-02 201782

O. 1 5 LEN 200. 0.84158E-Ol 0.61902E-Ol 103595'NGT 200. 0011425E 00 0.62114E-Ol 1.8394·CON 200. 0.27074E-Ol 0.95263E-02 2.8420

O. 1 6 LEN 199. 0019493 E 00 0.64072E-Ol 3.0423;IGT 199. 0.30143E-Ol 0.64197E-Ol 0.4695CON 199. 0.10490E 00 0.92720E-02 11.3135

O. 2 3 LEN 198. 0.73000E-Ol 0.65558E-Ol 101135\'GT 198. 0.16000E-02 0.57794E-Ol 0.0276CON 198. 0.41673E-Ol 0.70495E-02 5.9115

O. 2 4 LEN 198. 0.89999E-Ol 0.62632E-Ol 1.4369WGT 198. 0010450E 00 0.59842E-Ol 107462CON 198. 0014945 E-Ol 0.85550E-02 1.7469

O. 2 5 LEN 199. 0.68227E-Ol 0.61009[-01 101183\,GT 199. 0.98713E-Ol 0.55454E-Ol 1.7800CON 199. 0032704E-Ol 0.86547E-02 3.7787

O. 2 6 LEN 198. 0017900E 00 0.63219E-Ol 2.8313WGT 198. 0014599E-Ol 0.57730E-Ol 0.2529CON 198. 0.99271E-Ol 0.83610E-02 11.8730

O. 3 4 LEN 198. 0016300E 00 0.66773E-Ol 2.4410WGT 198. Q.I0610E 00 0.61064E-Ol 1.7375CON 198. 0.26728E-Ol 0.75457E-02 3.5421

O. 3 5 LEN 199. 0.47722E-02 0.65213E-Ol 0.0731WGT 199. 0.97113E-Ol 0.56757E-Ol 1.7110CON 199. 0.74377E-Ol 0.76690E-02 9.6983

O. 3 6 LEN 198. 0.10599E 00 0.67324E":01 1.5744WGT 198. 0012999E-Ol 0.58995E-Ol 0.2203CON 198. 0.57597E-Ol 0.73250E-02 7.8630

O. 4 5 LEN 199. 0015822E 00 0.62300E-Ol 2.5397WGT 199. O.20321E 00 0.58822E-Ol 3.4547CON 199. 0.47649E-Ol 0.90594E-02 5.2596

O. 4 6 LEN 198. 0.26899E 00 0.64478E-Ol 4,1719WGT 198. 0011909E 00 0.61003E-Ol 1.9523CON 198. 0.84325E-Ol 0.87834E-02 9.6004

O. 5 6 LEN 199. 0011077E 00 ().62885E-Ol 1.7615WGT 199. 0.84113E-Ol 0.56693E-Ol 1.4836CON 199. 0013197E 00 0.88783E-02 14.8648

42. 1 2 LEN 196. 0.27963E 00 0.95710E-Ol 2.9216

IWGT 196. 0.53090E 00 0.20074E 00 2.6446CON 196. 0.37725E-Ol 0012189E-Ol 3.0950

42. 1 3 LEN 194. 0.22040E 00 0010432E 00 201127\'GT 194. 0.56969E 00 0.22456E 00 2.5369CON 194. 0.64755E-Ol 0.11946E-Ol 5.4206

4? 1 4 LEN 195. 0019678E 00 0010603E 00 1.8558\'IGT 195. 0019904E 00 0.21247E 00 0.9367CON 195. 0.23664E-Ol 0012443E-Ol 1.9017

42. 1 5 LEN 193. 0.43936E 00 0010576E 00 401541

WGT 193. 0.682830 00 O.22398E 00 3.0 /.. 85CON 193. Oo12433E-Ol Oo11385E-Ol 1.0920

42. 1 6 LEN 1%. 0.29963E 00 0010953E 00 2.7354,

\vGT 196 • 0.54320E 00 0.23106E 00 203508CON 196. Oo10092E-Ol 0.10742E-Ol 0.9395

42. 2 3 LEN 196. 0.59224E-Ol 0.91462E-Ol 0.6475\vGT 196. 0038789E-Ol 001 9022E 00 0.2039CON 196. 0.27029E-Ol 0.11058E-Ol 2.4441

42. 2 4 LEN 197. 0.S2848F-Ol 0.93493E-Ol 0.e861IlJGT 197. 0033186E 00 0017647E 00 1.8804CON 197. 0.61390E-Ol 0.11591E-Ol 5.2962

42. 2 5 LEN 195. 0015973E 00 0.92939E-Ol 1.7186WGT 195. Oo15192E 00 0018919E 00 0.8030

I

CON 195. 0.50159E-Ol 0010454E-Ol 4.797842. 2 6 LEN 198. 0.20000E-Ol 0.97487E-Ol 0.2051

WGT 198. 0012300E-Ol 001 9838E 00 0.0620CON 198. 0.27632E-Ol 0.97917E-02 2.8220

42. 3 4 LEN 195. 0.23624E-Ol 0010217E 00 0.2312WGT 195. 0037065E 00 0.20246E 00 1.8307CON 195. 0.88419E-Ol 0.11326E-Ol 7.8063

42. 3 5 LEN 193. 0.21895E 00 0010181E 00 201505WGT 193. 0011313E 00 0.21430E 00 0.5279CON 193. 0077188E-Ol 0.10126E-Ol 7.6228

42. 3 6 LEN 196. 0.79224E-Ol 0010584E 00 0.7485WGT 196. 0.26489E-Ol 0.22198E 00 001193CON 196. 0.54662E-Ol O.94/+00E-02 5.7905

42. 4 ,5 LEN 194. 0.24258E 00 0010361E 00 2.3411',;GT 194. 0.48379E 00 0.20159E 00 203997CON 194. 0011230E-Ol o.10733E-Ol 1.0463

42. 4 6 LEN 197. 0.10284E 00 0010747E 00 0.9569WGT 197. 0034416E 00 0.21001E 00 1.6387CON 197. 0033757E-Ol 0010075E-Ol 3.3504

42. 5 6 LEN 195. 0013973E 00 0010726E 00 1.3027WGT 195. 0013962E 00 0.22141E 00 0.6306CON 195. 0.22526E-Ol 0.87047E-02 2.5878

70. 1 2 LEN 196. 0037963E 00 0.12930E 00 2.9358WGT 196. 0.89476E 00 0.37662E 00 203757CON 196. 0.20512E-Ol 0.10765E-Ol 1.9054

70. 1 3 LEN 19'•• 0.25510E 00 0.13809E ,00 1.8473\;GT 194. 0.69295E 00 0.40713E 00 1.7020CON 194. 0.27888E-Ol 0010987E-Ol 2.5381

70. 1 4 LEN 195. 0.21597E 00 0013745E 00 1.5712WGT 195. 0.27240E 00 0.39982E 00 0.6813CON 195. 0.29769E-Ol 0.10598E-Ol 2.8088

70. 1 5 LEN 193. 0031668E 00 0014025E 00 2.2578WGT 193. 0072238E 00 0.40211E 00 1.7964CON 193. 0012487E-Ol 0010957E-Ol 101395

70. 1 6 LEN 198. 0012084E 00 0013832E 00 0.8736'iGT 198. 0.28415E 00 0039352E 00 0.7220CON 198. 0018108E-Ol 0010267E-Ol 1.7637

70. 2 3 LEN 196. 0012453E 00 0012326E 00 1.0102WGT 196. 0.20181E 00 0.35629E 00 0.5664CON 196. 0.73755E-02 0.96684E-02 0.7628

70. 2 4 LEN 19 0.1636n Ou u 012270E 00 103337WGT 197. 0.62236E 00 0.34861E 00 1.7852CON 197. 0.50282E-Ol 0.92464E-02 5.4379

70. 2 5 LEN 195. 0.62948E-Ol 0012548E 00 0.5016WGT 195. 0017238E 00 0.35014E 00 0.4923CON 195. 0.80251E-02 0.96221E-02 0.8340

70. 2 6 LEN 200. 0.25878E 00 0012407E 00 2.0857

- --- ------ I

WGT 200. 0.Gl0G1E O(j u o3430:>E 00 I07UOO(ON 200. (). ;:lIOl~2E-Ol 0.09137E-02 0.2697

70. 3 4 LEN 19,. Oo39125F-01 00l3172l 00 0.2970\v'GT 19'; • (J.,.. 2Q55[ 00 O.30053r 00 101051CON It?!J. CJ.576?7E-Ol 0.94702E-02 6.08B2

70. 3 5 LEN 193. 0.61502E-Ol 0013452E 00 0.4577WGT 193. 0.29427E-Ol o 038253E 00 0.0769CON 193. 0015400E-01 O.90472E-02 1.5639

70. 3 6 LEN 198. O.134?~)L 00 00l3200[ 00 1.0109WGT 198. 0.40880[ 00 U.37451E 00 1.0915CON 190. 0.97798E-02 0.91340E-02 1;.0706

70. 4 5 LEN 194. 00l0070E 00 O0l3391E 00 0.7520WGT 194. 0.44997E 00 0037495E 00 1~2000

CON 194. 0.42257[-01 0.94197E-02 4.485970. 4 G LEN 199. 0.95127[-01 0.13219E 00 0.7196

WGT 199. 00l1746E-01 0.36713E 00 0.0319CON 199. 0.47877[-01 0.87000E-02 5.5026

70. 5 6 LEN 197. O019583E 00 O.13493E 00 1.4512WGT 197. 0.43823E 00 o 036900E 00 101876CON 197. 0.56200 E-02 0.90820E-02 0.6188

99. 1 2 LEN 196. 0.45742E 00 0016412E 00 207870WGT 196. 0011925E 01 0.55683E 00 201416CON 196. 0018634E-01 0.86301E-02 201591

99. 1 3 LEN 194. 0.26326E 00 0017l96E 00 1.5309WGT 194. 0.75938E 00 0.59466E 00 1.2770CON 194. 0.25446E-Ol 0.82331E-02 3.0907

99. 1 4 LEN 190. 0012158E-02 0016873E 00 0.0072WGT 190. 0018443E 00 0.56673E 00 003254CON 190. 0.25514E-02 0.86486E-02 0.2950

99. 1 5 LEN 191. 0.29323E-Ol 0016517E 00 0.1775WGT 191. 0.21893E-Ol O.54499E 00 0.0401CON 191. 0.26272E-Ol 0.85543E-02 3.0712

-99. 1 6 LEN 192. 0017961E 00 o 017043E 00 1.0538WGT 192. 0.45003E 00 0.55948E 00 0.8043CON 192. 0.35488E-Ol 0.82400E-02 4.3067

99. 2 3 LEN 196. 0019416E 00 0015533E 00 1.2499WGT 196. 0.43314E 00 0.52481E 00 0.8253CON 196. 0.68127E-02 0.81903E-02 0.8318

99. 2 4 LEN 192. 0.4562lE 00 0015114E .00 3.0184WGT 192. 0013769E 01 0.49051E 00 2.8072CON 192. 0016082E-01 0.86012E-02 1.8697

99. 2 5 LEN 193. 0.428 OE 00 Ool4741E 00 2.9040WGT 193. 0.12144E 01 0.46658E 00 2.6027CON 193. 0.76380E-02 0.85079E-02 0.8977

99. 2 6 LEN 194. 0.63704E 00 o ol5334E 00 4.1542WGT 194. Ool6425E 01 0.48389E 00 303944CON 194. 0016854E-01 0.81979E-02 2.0558

99. 3 4 LEN 190. 0.26204E 00 Ool5963E 00 1.6415WGT 190. 0.94381E 00 0.53322E 00 1.7700CON 190. 0.22895E-Ol 0.81807E-02 2.7986

I99. 3 5 LEN 191. 0.23394E 00 0.15596E 00 1.4999

WGT 191. 0078128E 00 0.51044E 00 1.5305CON 191. 0.8253lE-03 0.80861E-02 0.1020

99. 3 6 LEN 192. 0.44287E 00 0016162E 00 2.7401WGT 192. 0012094E 01 0.52624E 00 2.2982CON 192. 0010041E-01 0.77582E-02 1.2942

99. 4 5 LEN 187. 0.28107E-Ol 0015141E 00 0.1856WGT 187. Ool6253E 00 0.47l82E 00 0.3444CON 187. 0.23720E-Ol 0.85104E-02 2.7872

99. 4 6 LEN 188. 0018082E 00 Ool5756E 00 101476

WGT 188. 0.26560E 00 0.49012E 00 0.5419CON 188. 0032936E-01 O.81832E-02 4.0248

99. 5 6 LEN 189. 0.20893E 00 0.15380E 00 1.3584WGT 189. 0.42614E 00 0.46527E 00 0.9201CON 189. 0.92159E-02 0.80875E-02 101395

191. 1 2 LEN 196. 0.49006E 00 0.19970E 00 2.4539WGT 196. 0.2061OE 01 0.68336E 00 3.0166

,

CON 196. O. 87190E-0 1 0.78591E-02 11.0942191. 1 3 LEN 194. 0.19489E 00 0.20986E 00 0.9286

WGT 194. Ool2503E 01 0.70688E 00 1.7687CON 194. 0.11325E 00 O.29339E-01 3.8601

191. 2 3 LEN 196. 0.29516E 00 Ool9252E 00 1.5331WGT 196. 0.81120E 00 0.66341E 00 1.2227CON 196. 0.26066E-01 0.29188E-01 0.8930

266, 1 2 LEN 193. 0.94630E 00 O.24494E 00 3.8633WGT 193. 0.46529E 01 Ool1911E 01 3.9062CON 193. 0.45451E-01 0.79384E-02 5.7255

266. 1 3 LEN 192. 0.38696E 00 O.26918E 00 1.4375WGT 192. 0.26962E 01 0012894E 01 2.0910CON 192. 0.67782E-01 0.91301E-02 7.4240

266. 2 3 LEN 195. 0.55933E 00 0.24682E 00 2.2661WGT 195. 0.19566E 01 0.1l927E 01 1.6404CON 195. 0.22331E-01 0.95704E-02 2.3333

294. 1 2 LEN 193. 0012507E 01 0.27237E 00 4.5921WGT 193. 0.71431E 01 0015525E 01 4.6007CON 193. 0.10482E-01 0012367E-01 0.8475

294, 1 3 LEN 191. 0.26160E 00 0.30867E 00 0.8475WGT 191. 0010097E 01 o016946E 01 0.5958CON 191. 0.16705E-01 o011732E-01 1.4238

294. 2 3 LEN 194. 0.98918E 00 0.27474E 00 3.6003WGT 194. 0.61333E 01 0.15281E 01 4.0135CON 194. 0.27187E-01 0012425E-01 201880

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