Comp. Biochem. PhysioL Vol. 83B, No. 1, pp , /86 $ LIPOLYSIS POST MORTEM IN NORTH ATLANTIC KRILL
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1 Comp. Bochem. PhysoL Vol. 83B, No. 1, pp , /86 $3.+. Prnted n Great Brtan 1986 Pergamon Press Ltd LIPOLYSIS POST MORTEM IN NORTH ATLANTIC KRILL OLAV SAETHER, 1 TROND E. ELLINGSEN 2 and VIGGO MOHR 1 IDepartment of Bochemstry, The Norwegan Insttute of Technology, The Unversty of Trondhem, N-734 Trondhem-Nth, Norway (Tel: (7)59471); and 2Botechnology Research Unt, Dvson of Appled Chemstry, SINTEF, N-734 Trondhem-Nth, Norway (Receved 25 March 1985) A~tract--1. Three common speces of North Atlantc krll, Meganyctphanes norvegca (M. Sars), Thysanoessa nerms (Kroyer) and T. rasch (M. Sars), have been stored at C post mortem, and the lpolytc actvty followed by measurng changes n the lpd composton durng storage. 2. Both phosphoglycerdes and tracylglycerols were subjected to extensve hydrolyss wth the formaton of free fatty acds n all krll speces examned, whereas wax esters, consttutng a consderable proporton of the lpds n the Thysanoessa speces, were not hydrolysed at all. 3. In M. norvegca the tracylglycerols and phosphoglycerdes were hydrolysed at smlar rates, whereas n T. nerms and T. rasch the phosphoglycerdes were hydrolysed most rapdly. 4. For all krll speces examned, the rate of producton of free fatty acds was nearly constant durng the ntal phase of storage, and subsequently declned on prolonged storage. 5. At the end of the storage perod of 16- days, the free fatty acds consttuted about 35% of the total lpd n M. norvegca, and about 5% n the Thysanoessa speces. 6. The rate of producton of free fatty acds was about the same n all the three speces of krll and seemed to be ndependent of the total lpd content. INTRODUCTION Recent studes on Antarctc krll have demonstrated that both phosphoglycerdes and tracylglycerols serve as depot lpds n Euphausa superba, whereas E. crystallorophas deposts wax esters, tracylglycerols and phosphoglycerdcs (Ellngsen and Mohr, 1981; Ellngsen, 1982). Furthermore, Ellngsen (1982) has shown that storage of samples of krll post mortem resulted n extensve degradaton of tracylglycerols and phosphoglycerdes wth the formaton of free fatty acds, thus suggestng that these speces of Antarctc krll contan sgnfcant levels of actve lpases and phospholpases. The work by Ellngsen (1982) formed the bass of a program for examnaton of the chemcal composton and post mortem changes n three common speces of North Atlantc krll: M. norvegca, T. nerms and T. rasch. The lpd composton of North Atlantc krll has been examned n detal by Sargent and Falk-Petersen (1981) and Saether et al. (1985). Saether et al. (1985) have provded clear evdence that tracylglycerols are the only depot lpds n M. norvegca, whereas phosphoglycerdes, wax esters and tracylglycerols serve as depot lpds n T. nerms and T, rasch. The post mortem changes n the lpd of these speces have not as yet been studed. The present communcaton provdes data relatng to the post mortem changes n the lpds of these speces of North Atlantc krll. MATERIALS AND METHODS The krll used n the present studes were caught n Balst]orden and Ullsfjorden n Northern Norway durng the perod 23 February-21 Aprl Detals of the catches are gven n Table I. The catchng and handlng of the krll have 51 been descrbed n detal by Saether et al. (1985). On board the vessel 2-5 ml samples of whole and homogenzed krll were dstrbuted nto 1 ml plastc bottles wth screw caps wthn 15-3mn of catchng, and stored as descrbed below. Krll were also frozen n CO2-ce wthn 5-15 mn of catchng and stored at -8 C for no longer than I year before beng used expermentally. A closer examnaton of the dfferent Thysanoessa speces was carred out n the laboratory. Blocks of krll were thawed and sorted accordng to speces at C wthn 4-6hr of thawng. Each specmen was subsequently refrozen on CO2-ce after sortng. Krll belongng to the same speces and haul were pooled and homogenzed n the frozen state, usng an Ultra-Turrax TP 18-1 (Janke & Kunkel KG). Samples of 2-5 g of the homogenates were dstrbuted nto 1 ml glass tubes wth screw caps. The samples of both fresh and frozen/thawed krll were stored n an ce-water bath, and ndvdual samples removed as a functon of tme, and stored at -8 C untl analysed. The entre content of each sample was homogenzed and used n the analyss. The lpds were extracted wth chloroform-methanol-water accordng to Hardy and Keay (1972). The man classes of lpd were separated by thn layer chromatography n accordance wth Ellngsen (1982), as descrbed by Saether et al. (1985). The dry weght of each sample was determned by dryng I-2 g of homogenate for hr at 15 C. RESULTS AND DISCUSSION The post mortem changes n the lpd consttuents of three common speces of North Atlantc krll, M. norvegca, T. nerms and T. rasch, were examned at C. The studes were based on fve catches of krll from fords n northern Norway over a perod of 2 months (February-Aprl). Based on measurements of length, a majorty of the krll examned seemed to be sexually mature (Saether et al., 1985). RIMFROST EXHIBIT 7 page 1
2 52 OLAV SAETHER et al. Table 1. Data on the krll catches Total lpd Haul No. Date Localty Krll speces content (% of dry weght) Svartnes n T. nerms 35.3 Balsfjorden T. rasch 27.6 unsorted Thysanoessa* Ulsl]orden M. norvegca Svartnes n T. nerms Balsorden T. rasch unsorted Thysanoessat Ulsl~orden M. norvegca Svartnes n T. nerms 14.9 Balst]orden T. rasch 11.4 unsorted Thysanoessat 13.9 *Mxture of 7% T. nerms and 3% T. rasch. tmxture of 8% T. nerms and 2% T. rasch. In M. norvegca both tracylglycerols and phosphoglycerdes were hydrolysed wth the formaton of free fatty acds post mortem, suggestng that both lpases and phospholpases were actve durng storage. Judgng from the data, the hydrolyss of tracylglycerols and phosphoglycerdes proceeded at smlar rates. The rate of release of fatty acds gradually decreased durng storage n both catches examned, and at the end of the storage perod, free fatty acds accounted for about 35% of the total lpd (Fg. 1). The rate of release of free fatty acds, on a dry weght bass, was also smlar n the two catches (Fg. 2). In the Thysanoessa speces the ncrease n the free fatty acd content post mortem was prmarly due to extensve hydrolyss of phosphoglycerdes, but the hydrolyss of tracylglycerols was also sgnfcant (Fg. 3). The rate of free fatty acd producton was about the same n krll havng wdely dfferent total lpd contents ( % dry wt), thus suggestng that the rate of lpd hydrolyss was ndependent of the total lpd content. (Table 1 and Fg. 2). At the end of the storage perod free fatty acds consttuted about 5% of the total lpd n the Thysanoessa speces, whch s sgnfcantly hgher than n M. norvegca. A com- c~ 8 d- 64 ~'~ HAUL 23 BO J -, HAUL z~... o-.1 ~.j 48 el) b-- 'I1 ~ 32 ~l.i-.-.-:- 16 I /-..A I yafj-, _.~,dk IB 3 Fg. 1. Content of man classes of lpd durng storage ofm. norvegca (haul 23 and 27) at O Cpost mortem. Wax esters: O, tracylglycerols: A, free fatty acds: A, cholesterol + dacylglycerols: IS], phosphoglycerdes: m. COI 15 THYSANOESSA SP, _. + ~ /~_m 1 M, NO,RVEGICA ~W >->- 9 F-r~ ~-~ 6 t.././~/" /e u-o 3 v o 25 'lo 25 Fg. 2. Content of free fatty acds n dfferent hauls of Thysanoessa speces (haul 21:, : [] and 28: A) and n M. norvegca (haul 23: and 27: []) stored at C post mortem. RIMFROST EXHIBIT 7 page 2
3 Post mortern lpolyss n krll B HAUL 21 4B, HAUL /~////_A O--,ll F1 F-- V rl~ O- f / ,1J 6 HAUL 28 /~' 48.--I Z L2 e Fg. 3. Content of man classes of lpd durng storage of Thysanoessa speces (haul 21, and 28) at C post mortem. Wax esters: O, tracylglycerols: A, free fatty acds: A, cholesterol + dacylglycerols: IS], phosphoglycerdes: 1. parson of the rates of hydrolyss n North Atlantc and Antarctc krli reveals that the rates observed n M. norvegca, the Thysanoessa speces and E. crystallorophas are nearly the same, and hgher than that detected n E. superba (Fg. 4). The content of wax esters and cholesterol vared lttle durng storage of the North Atlantc krll speces post mortem, suggestng that there s no sgnfcant formaton of cholesteryl esters durng the frst week of storage (Fg. 3). The contents of wax esters and cholesterol were lkewse constant durng the frst week of storage of E. crystallorophas (Ellngsen, 1982). Esterfcaton of cholesterol has been observed durng storage of fsh post rnortem (Lovern et al., 1959)..)UJ >->- 6 u.w 4 ~u_ u-o 2 f TIME (DAYS) Fg. 4. Content of free fatty acds durng post mortem storage at C of M. norvegca (haul 27: A), Thysanoessa speces (haul 28: A) and Antarctc krll speces, examned by Ellngsen (1982) (E. crystallorophas: [S], E. superba: 1). 15 The studes of the post mortem changes n the Thysanoessa speces on board the research vessel were based on catches contanng 7-8% T. nerms and 2-3% T. rasch. A separaton of T. nerms and T. rasch requred mcroscopy of each ndvdual, whch was not possble on board the vessel, due to prorty gven to other experments. However, frozen krll from haul 28 were thawed n the laboratory, sorted accordng to speces, homogenzed and stored at C for a closer examnaton of lpolytc actvty n T. nerms and T. rasch. The results ndcate a smlar pattern of lpd degradaton n the two speces (Fg. 5). Furthermore, the rate of release of free fatty acds was also nearly the same n the two speces (Fg. 6). It s of nterest to consder the lpolytc actvty n relaton to the lpd class composton of the dfferent speces of krll. In M. norvegca tracylglycerols are the only depot lpds, whereas n T. nerms and T. rasch phosphoglycerdes, wax esters and tracylglycerols serve as depot lpds (Saether et al., 1985). E. crystallorophas also deposts phosphoglycerdes, wax esters and tracylglycerols, whlst E. superba uses only phosphoglycerdes and tracylglycerols as depot lpds (Ellngsen, 1982). A systematc comparson of the dfferent speces of krll reveal no correlaton between the relatve proporton of the man classes of lpd, and the rate at whch the lpds are hydrolysed. It appears, therefore, that the lpolytc actvty n each speces depends on other factors, such as the level of lpases and phospholpases n the krll, the extent of emulsfcaton of the substrates and the degree of contact between enzymes and substrates. Studes of dfferent speces of fsh have also faled to reveal any correlaton between RIMFROST EXHIBIT 7 page 3
4 54 OLAV SAETHER et al. T, INERMIS kº_ T, RASCH I I '-;~ 4 a. <~- 2 _I ~ ~'-~--~-... ZI~ lo I.... ~ v Fg. 5. Content of man classes of lpd durng storage of homogenzed, frozen and thawed T. nerms and T. rasch (haul 28) at C. Wax esters: O, tracylglycerols: A, free fatty acds: A, cholesterol + dacylglycerols: F-l, phosphoglycerdes: 1. the lpase- and phospholpase actvty and the content of tracylglycerols and phosphoglycerdes (Olley et al., 1962; Shono and Toyomzu, 1973; Nar et al., 1978). As ponted out above, phosphoglycerdes were subjected to more extensve hydrolyss than tracylglycerols n the Thysanoessa speces (Fgs 3 and 5). A smlar observaton was made n the study of the lpd degradaton n the Antarctc krll E. superba and E. crystallorophas (Ellngsen, 1982), whereas n M. 5 o.}-r,", co 4 ~ M >->- 3 I--,'-1 u.w 2 -r tall-- u.o v f,, 2 4 Fg. 6. Content of free fatty acds durng storage of homogenzed, frozen and thawed T. nerms (haul 28: A) and T. rasch (haul 28: FI) at C post mortem. B I norvegca the lpase actvty was of the same order as the phospholpase actvty (Fg. 1). It has been proposed that the more extensve degradaton of phosphoglycerdes than tracylglycerols n Antarctc krll speces may be due to the more hydrophlc character of the phosphoglycerdes, and hence a better enzyme to substrate contact (Ellngsen, 1982). The same may apply to North Atlantc Thysanoessa speces. Tracylglycerols are more hydrophobc, and may as a result, requre more effectve emulsfcaton for hydrolyss to take place. Wax esters are even less polar than tracylglycerols, and nadequate emulsfcaton and/or a low enzyme level may explan why the wax esters were not subjected to post mortem hydrolyss nether n the Thysanoessa speces nor n E. crystallorophas (present results and Ellngsen, 1982) The tracylglycerols n M. norvegca were subjected to much more extensve hydrolyss than those of the Thysanoessa speces, whch may be due to a hgher lpase content, and/or better emulsfcaton and, hence, a better enzyme-substrate contact. The lpds and lpolytc enzymes are presumably separated n the tssues of lvng krll. Thus, proteolytc actvty s probably requred to brng the lpds nto contact wth the lpolytc enzymes post rnortem, as suggested by Ellngsen (1982). An extensve proteolyss post mortem has been establshed BO M,NORVEG'ICA 6O THYSANOESSA SP, /.~ - '-' 64 o-~ 48 "1..-I O')F- ~ & 4B, /// J 6 IB IB Fg. 7. Content of man classes of lpd durng storage of homogenzed M. norvegca (haul 23) and Thysanoessa speces (haul ) at C post mortem. Wax esters: O, tracylglycerols: A, free fatty acds: A, cholesterol + dacylglyeerols: F-l, phosphoglycerdes: 1. RIMFROST EXHIBIT 7 page 4 3
5 Post mortem lpolyss n krll 55 both n Antarctc krll (Ellngsen, 1982) and n North Atlantc krll (unpublshed results), whch may explan why lpolyss n all speces of krll examned so far proceeded wthout any dstnct, ntal lag-phase (Fgs 1 and 3; Ellngsen, 1982). Storage of homogenzed samples of M. norvegca revealed a three-fold ncrease n the ntal rate of hydrolyss of both phosphoglycerdes and tracylglycerols as compared to whole krll, whereas n the Thysanoessa speces, there were no sgnfcant dfferences n the rate of hydrolyss of whole- and homogenzed anmals (Fgs 1-3, 7). Separate studes have revealed that the bacteral actvty seems to be low durng the frst week of storage of M. norvegca and Thysanoessa speces at C (unpublshed results). Thus, there s reason to beleve that the post mortem changes observed durng the frst week of storage are prmarly due to autolytc processes. Low bacteral actvty durng the frst week of storage post mortem has also been observed n Antarctc krll (Ellngsen, 1982). In concluson, the present work and the studes of Ellngsen (1982) have provded evdence that the hydrolyss of both phosphoglycerdes and tracylglycerols s very rapd and extensve n both North Atlantc- and Antarctc krll, whereas the content of wax esters seem to be constant durng post mortem storage of wax ester rch speces. It s also concluded that the rate of producton of free fatty acds, on a dry weght bass, s about the same n M. norvegca, T. nerrns and T. rasch as n the Antarctc krll E. crystallorophas, examned by Ellngsen (1982). REFERENCES Ellngsen T. E. (1982) Bokjemske studer over antarktsk krll Dr. ng. thess, Department of Bochemstry, The Norwegan Insttute of Technology. The Unversty of Trondhem, N-734 Trondhem-Nth, Norway (n Norwegan wth summary n Englsh). EUngsen T. E. and Mohr V. (I 981) Lpds n Antarctc krll. Composton and post mortem changes. Proc. l lth. Scandnavan Symposum on Lpds. (Edted by Marcuse R.), pp SIK, Gothenborg, Sweden. Hardy R and Keay J. N. (1972) Seasonal varatons n the chemcal composton of Cornsh mackerel, Scomber scombrus (L), wth detaled reference to the lpds. J. Fd TechnoL 1, 5. Lovern J. A., Olley J. and Watson H. A. 959) Changes n the lpds of cod durng storage n ce. J. Sc. Fd Agrc. 1, 327. Nar P. G. V., Antony P. D., Gopakumar K. and Nar M. R. 978) Lpd breakdown n ol sardne (Sardnella longceps) durng frozen storage. Fsh. Technol. 15, Olley J., Pre R. and Watson H. (1962) Lpase and phospholpase actvty n fsh skeletal muscle and ts relatonshp to proten denaturaton. J. Sc. Fd Agrc. 13, Saether O., Mohr V. and Ellngsen T. E. 985) Lpds of North Atlantc krll. (To be publshed.) Sargent J. R. and Falk-Petersen S. (1981) Ecologcal nvestgatons on the zooplankton communty n Balsfjorden, northern Norway: lpds and fatty acds n Meganyctphanes norvegca, Thysanoessa rasch and T. nerms durng md-wnter. Mar. Bol. 62, Shono T. and Toyomzu M. (1973) Lpd alteraton n fsh muscle durng cold storage--i. Expresson of lpd hydrolyss and oxdaton n jack mackerel muscle based on decrease n C22:6 acd. Bull. Jpn. Soc. Sc. Fsh RIMFROST EXHIBIT 7 page 5
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