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1 This paper nt t be cited withut prir reference t the authr Internatinal Cuncil fr the Explratin f the Sea C.M. 1981/M:lS Anadrmus and Catadrmus Fish Cmmittee CHARACTERIZATION OF PROTEASES IN THE DIGESTIVE TRACT OF ATLANTIC SALMON (SALMO SALAR) IN COMPARISON WITH RAINBOW TROUT (~~~MO GAIRDNERI) By Krisna R. Trrissen Institute f Marine Research, Matre Aquaculture Statin, N-5198 Matredal, Nrway. Present by Dag MØller

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3 - l - ABSTRACT The prtelytic activities f the prteases extracted frm the digestive tracts f Atlantic salmn and rainbw trut shw similar characteristics in the ptimum temperatures. In Atlantic salmn, the ptimum temperature f the stmach extract is 37.5 C, f the pylric caeca and f the small intestinal extracts, 52.5 C. The extract frm the large intestine shws tw ptimum temperatures at 45 and 52.5 C. A similar pattern was bserved in the extracts frm the digestive tract f rainbw trut except fr the stmach extract, where the ptimum temperature is 35 C. The prtelytic activities f the intestine are very prnunced at the cnjunctin between the small and large intestines. The characteristics in electrphretic mbility f these prteases will als be studied. INTRODUCTION Salmnid fishes have high prtein requirement fr grwth. In species f salmn and traut, abut 40% f crude prtein level in diet is required fr ptimum grwth rate (Cwey,l979). The prtelytic activities in the digestive tract shuld be relatively high, and expected t influence n the grwth rate f these fishes. A lt f wrk n prteases in the digestive rgans have been dne in different fishes, such as rainbw trut (Kitamikad and Tachin,l960; Kawai and Ikeda,l973a; Rungruangsak and Utne,l98l), mackerel (Oshir,l971) and carp (Kawai and Ikeda,l972 and 1973b). The knwledge abut digestive enzymes in Atlantic salmn is hwever limitted.

4 - 2 - In a preliminary wrk (Rungruangsak and Utne,l981), the prtease activities in the digestive tract f rainbw traut was studied. In this experiment, the prteases f the different digestive tissues were characterized in mre detail t bserve the ptimum temperatures and electrphretic mbilities. The characteristics f the peptic-like and the tryptic-like enzymes were cmpared between tw salmnid fishes, Atlantic salmn and rainbw traut. ~ffiterials AND METHODS Tw grups f fishes, Atlantic salmen (Salm salar) and rainbw traut (Salm~ gairdneri), were used as experimental fishes in this study. Each grup was cultured separately in freshwater 3 in a rund fibreglass tank (1.7 m ) at Matre Aguaculture Statin. The fishes were fed a cmmercial dry pelleted feed (Tess laksef6r, T.Skretting Ltd.,Stavanger). The fishes ranged frm 62 t 192 g and 17 t 25 cm. The fish in each grup were sampled twice, 4 fish each time, fr duplicati and averaging the results. They were killed and the digestive tracts were remved at nce, emptied, defatted and cut int five sectins: stmach, pylric sectin and three equally-lng sectins f the intestines. During sampling the temperature was cntrlled at 4-5 C. The crrespnding sectins frm 4 fish in each grup were pled and the prtease f each sectin f the digestive tract was extracted and assayed fr prtelytic activity by the methd f Rungruangsak and Utne (1981). In rder t cmpare the ptimum temperatures f the prtease activities in the digestive tract between Atlantic salmn and rainbw traut, the assayed

5 - 3 - temperatures were varied frm 20 t 60 C. Anther experiment was carried ut in rder t see the difference in prtease activities f the intestinal sectins in mre detail. The intestines frm 2 fish in each grup were cut and divided int 4 sectins: the first half and the secnd half f small intestines, and the first half and the secnd half f large intestines. The prtease f each sectin was extracted and the enzyme activities were als determined by varying the temperature frm 20 t 60 C. RESULTS By varying the temperature, the ptimum temperatures fr the determinatin f prtease activities f the stmach extract and pylric caeca extract f Atlantic salmn are 37.5 and 52.5 C, respectively (Fig,l). The first sectin f the intestine has very lw prtease activity while the secnd and the third sectins have higher activities and shw tw ptimum temperatures at 45 and 52.5 C. As shwn in Fig.2, the similar pattern f the ptimum temperatures was bserved in the extracts frm the digestive tract f rainbw trut except fr the stmach extract, the ptimum temperature is 35 C. The prtease frm the stmach extract f bth Atlantic salmn and rainbw trut shwed peptic-like activity and has the ptimum ph abut 2. The tryptic-like enzyrnes in the pylric caeca and the intestinal extracts have a brad ptimum ph at abut 9. Since there was extremely lw prtelytic activity in the first sectin f the intestine (Fig.l and Fig.2), the first half

6 - 4 - f the small intestine was mitted in the secnd experiment. It is. clearly shwn, especially in the intestine f Atlantic salmn, that the prtease activity in the secnd half f the small intestine has the ptimum temperature at 52.5 C while bth sectins f the large intestine shw tw ptimum temperatures at 45 and 52.5 C (Fig.3). DISCUSSION The prtelytic activities in the digestive tracts f Atlantic salmn and rainbw traut shw the ptimum ph at abut 2 fr the peptic-like activity frm the stmach extract and abut 9 fr the tryptic-like activities frm the pylric caeca and intestinal extracts. These ptimum phs are typical fr pepsins and trypsins, respectively. Frm Fig.l and Fig.2, the ptimum temperature f the pepticlike activity in Atlantic salmn (37.5 C) is smewhat higher than that in rainbw traut (35 C). The tryptic-like activities have the same pattern f the ptimum tempera tures, 52.5 C fr pylric caeca extract, 45 and 52.5 C fr the intestinal extracts. The cnditins were similar t thse bserved by Kitamikad and Tachin (1960), the ptimum temperatures f the prteases in rainbw traut were bserved t be 40 C fr the extract frm the stmach and 45 C fr the extracts frm pylric caeca and intestine. Oshir (1971), fund the purified prteinase frm the pylric caeca f mackerel t be mst active at 43 C. The ptimum temperat~re f the tryptic-like activity frm the pylric caeca extract was fund t be higher in this experiment and the enzyme activities in the extracts f intestine exhibit tw active

7 - 5 - temperatures in bth Atlantic salmn and rainbw trut. As shwn in Fig.3, the prtease activity f the secnd half f the small intestine has high ptimum temperature at 52.5 C as in the pylric caeca, nly the large intestine shws tw active temperatures at 45 and 52.5 C fr tryptic-like activities& It is clearly shwn in Atlantic salmn. The enzymes in the first half f large intestine have higher prtelytic activities than the ther parts f the intestine. It can be cncluded that there must be at least tw different tryptic-like enzymes in the large intestine. The levels f prtease activities f the intestine are very high in the extracts frm the secnd half f small intestine and the first half f large intestine, but very lw in the ther parts especially in the first sectin f intestine. The digestibility in the intestine f Atlantic salmn and rainbw trut must be fairly high at the cnjunctin between the small and large intestines. The characteristics in electrphretic mbility f these prteases is n study. REFERENCES Cwey,C.B.,l979. Prtein and amin acid requirements f finfish. In: J.E.Halver and K.Tiews (Editrs), Finfish Nutritin and Fishfeed Technlgy, Prceedings f a Wrld Sympsium, Hamburg June,l978. Vl.!, Berlin, pp Kawai,S. and Ikeda,S.,1972. Studies n digestive enzymes f fishes- II Effect f dietary change n the activities f digestive enzymes in carp intestine. Bull.Jpn.Sc.Sci.Fish.,38:

8 - 6 - Kawai,S. and Ikeda,S.,l973a. Studies n digestive enzymes f fishes III Develpment f the digestive enzymes f rainbw traut after hatching and the effect f dietary change n the activities f digestive enzymes in the juvenile stage. Bull.Jpn.Sc.Sci.Fish.,39: Kawai,S. and Ikeda,S.,1973b. Studies n digestive enzymes f fishes~ IV Develpment f the digestive enzymes f carp and black sea bream after hatching. Bull.Jpn.Sc.Sci.Fish.,39: Kitamikad,M. and Tachin,S.,l960a Studies n the digestive enzymes f rainbw traut- II Prteases Bull.Jpn.Sc.Sci.Fish., 26: Oshir,Z.,l971. s tudies n prteinase in the pylric caeca f fishes- II Same prperties f prteinase purified frm the pylric caeca f mackerel. Bull.Jpn.Sc.Sci.Fish.,37: Rungruangsak,K. and Utne,F.,1981. Effect f different acidified wet feeds n prtease activities in the digestive tract and n grwth rate f rainb\v trut (Salm gairdneri Richardsn). Aquaculture,22:67-79.

9 Figure 1: The prtease activities at varius temperatures f the different sectins f the digestive tract f Atlantic salmen. STOMACH z w l g: 2 ~ "... a: :I:... w z u; ~ 1- w...l :E :::). PYLORIC CAECA INTESTINE 8 - first sectin 0-0 secnd sectin third sectin c c-----c-----c-----!-----c--c TEMPERATURE ( C) 50 60

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11 Figure 2: The prtease activities at varius temperatures f the different sectins f the digestive tract f rainbw trut. 35 ~ ~- 3 STOMACH O~..._0"" ~ " 0/ ~ ~ --.. w...j ~ ~ 8 INTESTINE -c first sectin - secnd sectin A-6 third sectin TEMPERATURE ( C) 50 60

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13 Figure 3: The prtease activities at varius temperatures f the different intestinal sectins f rainbw trut and Atlantic salmn. RAINBOW TRO UT 4 D z jjj l a: c. (:;l :!... a: ::.t:... w z U5 a: > 1- I.U...J :?- ~ ATLANTIC SALMON /l l l // ~rt::l ~ ~/ /'c l ~ 0-0 secnd half small intestine - first half large intestine ~.."_."~ secnd half large intestine

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