lill 1 II I IlItl \ w l | i | | 5% III THE USE OF RABEOACTEVE ERGGSTEROL 5N THE STUDY OF LTPSD ABSORPTEON IN THE ALBENO RAT Thesis for the Degree of M. S. MICHIGAN STATE COLLEGE Robert Dougias MacKenzie 1954 MICHIGAN TTTTTTTT nnnnnnnnnnnnn THE US F “11-11;: 115101111”. ’IU} 153.3123. 113 TIL; {LUJY 0F LIEJID [11°12 1%‘IZJ1 1.; 11. 111.3311JI3 MT 31 R03: PT DJIJGLAS 13131:; 2113 A 1‘11 31.1313 Suhnit'bod to the School of Cr: 1111111."; Statics of I-iichiggan Sham College of 11721le taro and Ap :flia Li Scic .ce in partial fulfilhc.-t of the rcqui .. mts for the degree of 111133.111 ' SCLiI Dozm'tnent of (3101215. airy 1951+ n'f." -‘. “f . " - jII'Y '1" ~10 AU—J;J3¢:A. m — .;.LAI;Q The author is greatly ind eb ur a to Jr. Carl A. Haga-pcrt 1‘01: sugr'osting the problem an for much 01 ico an; 132- w1m1"~*,-.t flxrouglh out the course of the 93-1 :‘imental work and for 111113.110.le msistance with the preparation of the mscrint. Tharflzs are due Mr. Leo chvnr, Forarmn of Caraztzfi-cex's, Vi 1.1251111 Assay Lab. for 4.1.111ch n.11b1e help in caring, ' for the rats and 31*Muwlox1s d Irma the i..vc"t..{' tion. Tb an}: s are also duo to Dr. Edward U. err-um for his many 1111131111 nggostiona on use of radioactive material. f"!5".-a- ~l .- '1 H H H —.~..~..,‘..J .q k,’\.'. ‘ g.‘ T ‘-. 1'31 .‘rr \TIT.‘ .. --.J.-\p“.l._— -..\.,.,'.........v . Q . p . . . . . . . . O Y'-:""“ H.’|’I"RT' 1.‘ -".... QM.) . g o o o o n o o o o o 0 fi‘ . 0‘} Q". T '0 _ 1 3‘ j .m , . ,~ ‘ u. n J££3I301 133)-”. .2 ' ) .4 "‘ ' n? «-3... .4”: '3‘... .f,‘ r;_‘.‘.,.,.v._o (3"7 1‘; .~ T'n-vr‘. L).J‘_la‘ ' )..’_‘_r.a A L,..'. ‘_' ’ h ' L '\.'--'~. . . 4"... ‘”l"'~"-""""‘"' 'nA v , ~-v ‘P’-f¢ iJJ'Xi ..l._...’. ~ ¢.,..‘L .l t . ‘.‘ 'J— .2 . . . g . a g g g . I V V'p.. ‘,v1.)'1 qr-.- '- -~r J“ » - -v 0 o o O 0 o O O o o 0 Pt ‘ V " "fl ‘ . ' ‘ V " \dro“: IV " 1' " ’- “I ' "‘ ‘\“‘§‘fi . um I,.,.'\ .. r- .J ' '.‘. - ;' (Ix hr... ~J.'.‘f . .2. ' ‘ ”4‘4... 'J ,. \ <2ka xvu‘ "3. “H“.I. II ‘d-n— .‘ é....-.\4--t) r1 m1 .1..'3....-'.-O I g o o f" '1 $33.). .3 II . g “10‘?” l' A , ~‘ ' ~I‘-"‘-'-*‘—’-‘ Q o o o o o o a ..---I—»--7 ‘:4 ~7f u-J-..:....s. .... coo-o. v—‘-r~\l- ,‘r -1- f, ,l I» _ , v4.41» -,'\...-...-.L - J. ‘ . 344090 IEIITuOUUCI‘IOl-I IIJI'RODUCTIUH In order for 10 animal body to utilize foods that are ingwfiod, complex :m‘“ rials such as _v“:rot<:;;'-.no, carboi'mlru‘bos and possibly lipids mot be broken down to more soluble and diffuoible 081313011121218. The labia-r than pass timerstgh the intestinal. ragirzbrmm into the circflatory afghan. This ocmrs tbmmgh the epitholizzl calls of tho villi by two diffonrnt pa “Aways. 1) Host of the products of digestion enter the portal am; .123 by my of the capiuarioo, 11191100 to the intestinal. veins, the portal vein and finally reaching; the liver. 2) In the case of fats the hmfimtic firstom is an impor " it avfo'nuo of absorption and involves the lact:-al3 (lymph vowels), fonninrr tributaries to the thoracic Ctuct which ultimately ('mptics 111130 the blood otnrmn. t has gone tally been lold that. about. 60,3? of he ingratai lipids Inter the hmlphatic system and the rcmimlor, the porn: system. The mmct mothoai of transfer of lipid material from the intestine to this: lacteuls or crgapillmdeo has long: born a. subjcct of study and controvaroy. SrVoral theories of tho absorption of fat have been dovnloyal. A survey of tort: books and Kite-mature loam-a the impression that the 12.21;} 0221 ty of witors are of the ooxiavi ction that fats ore COK';pl-‘f“tle/” hydrolysozi’. by 1:233:15 a to free fat‘w acids and glycerol in the in nation bofor-a-- passing into the lactoala or canillarles. A summer number Etzold to the theory that, the fat may A b" absorbed with only slight l‘mlrolysis providcxi Lificiont -,rrols_3.fioation can be ochi .:v:d to films vat; 321.111 grafticlcs to trnv-zrrSr: the inmsfimfl. Micr. Tho px-zrpooo of thin :s‘ov‘vr horz;.-i‘or:: was to provide further ax; '.ori:u,z;t11 ovifionce that. 13.. gilt contaibuic to a batter mism- otmdinr' of tflifi tint-cixmzisz of fat absorption. L) . $.2- ‘-.-J 1113 all ‘ .5 Yv". .m,‘ -‘-§' .4 .‘ J;.Lu.-.Ul‘o.IU.'u‘r are ~ —- . ' ' .. ' ‘ - n T- :L ‘v .I‘ ' \p ‘v - ‘ 1‘ -‘ ' .I I) °‘ '1‘ .1] \ .‘f “" y .-< “ v, - Q. ~ A L44 ~ I I(!oo J'Q‘. J-.|' A-» -1- C'~.(/:‘( 1k) Lu: 4 ..I J 1“." (2L 1.8., n‘a\t‘ \'V~~‘J’.:r'\‘~ ‘1 w n"? 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Log )1 .. «J O... n... ‘ J ~J-L L- 101 '-’ “" q \ 1" , “W‘ V k \ ”'Wu J _, _ . i 1 n '.“ . .‘ .1. ..'..'. . 4.! .u n-—y ran-fa. -‘ ~~n -’- —'~ -» -1 .n-w.‘ r ' n -" ".v "‘- “11“; s ‘5‘} ’ I \I . '.»- s..-IJ-1 VgJ. _’n ‘L’ k4.” U- .0 i)‘1 l “4“. ' 0" V‘l'“‘ "' U ‘ U J*-_"" I I_ 0 ‘1 ‘ . 1 .. ...-.‘ -'¢ ,. 31 ‘ - , _ - Fl . ' U QC 'J -i ‘J \L y J.I-.J_-v . T‘fl - '5 " b. r\ .J' '-,»- «o. -"'*.~'fl “- I : "/" r‘ 1".“'.‘¥~‘ ‘ " "‘ I. ["0" .- I ’\71 1:! (‘17:. ‘1. l . .. \~ .1 J”. L . 4...... ‘ _.Q 5.. .. ; JJ J.) v.1. L. Us! ‘3. J-.......~. m. Uh. 51...”. 0 \ I I O ' ‘ " .L’ 1 J - 1 - _ n , 1- .H .4-.. ‘ . ,., "H“ .H .. -«~"-'."2r‘- 11-, ‘ . .- - .«r -‘ , . ._ («o ‘ V | . . . . - . -: I ‘ ._, - Of 1 :3 'J U. .' SIC"... L5.) 5"! .-1 Karl .‘w ‘ .. . _ k u__ ~/-. v.-. \J- - ‘1 ....-- J. o I 0.5.. . . , {1.1.} k.» U; Q I a a 4. ‘\ I‘ ! 1 I - ‘ I ““ 1 \‘1 77 ‘fi"\f1 ir‘ ., ..‘,‘. ....: _ _ _ I . ._ ~ ‘ _ \ '_ - a. '~ 5- n p- " ‘f ‘ . . I . ‘ ‘ 5, ‘ ‘ U: 13:. 3.)O ~'.’ .:.r‘.' J x \r‘: U- l. l‘ T..- F- L . x. L. L... . x .d.‘ Lo. do» “ants-I- ‘ 5 '. ”I“ I.) ' ' *5 -.—'- dflo‘camtc to r2053 and collected the chylo, which he examined for Optical ‘0'! acti vi" 3'. 321 worm-mix} Qua: 'a‘im: 1:13 3 lootsx‘z 13 03118301? its strong 2 optical activity (lost noon 9: drolysia), low Inciting point, ability to :mflsfzi‘y and good utilization by the toot 5:111:22. ‘ . [he mzbmquont observation 0 ‘ t} a lack of Optical activity in the ether extract of ’19 cfzylc was iilt;f.1‘:i)r‘:tc'.3 as negating tho absozption of he unchag-od estvr. Vorzar (20) agreed in part xith Pflug. r, iz‘zoofzzr as the total hyirolysis of neutral fat by inta-stflml lipase was conch» .zcd. In own: .0 :rircxrzzvnt thv inability of soaps to remain Gable in acid solution, ho presented (avid 21cc to chm-J that bile azlts are 1:137:0th aids in tho. absorption of fat. {Io dcczmnstratod that both taurooholic and glycocholio acids 2:11:17: added to aqueous 11111310113 of palmitic, stcario or 012312 acids readily formed solutions; own at a pH of 6.25 and cubs quozz‘oly passed wrong: parcEm-w‘zt mxbranos. Iioyholonotric studios of solutions of bilo acids with fatty acids infzicctcd a zoo}. clam rintionship. ‘3 higher concentrafiion, cgregjai‘..c~s occurred. Lii‘fusion 63C_‘el*'171{;1'1t8 sizomxl that in very dilute solution, nool'ly all the fatty acid 1.53:: diffaoi‘olo whereas in concentrated solution, 011?}; a part diff-Jam. Fine emulsions of neutral fat did not :SLifi‘uo: at all, av 12 in “me proscznoo of glycocholic acid. Vorzar conclizlocl that only this action of glycocholic anti taro cholio acid on Line fatty acisi makes the absorption of fats 330335.th am ox 11:11:23 it wider 31111'3101031‘231 con- ditions . “’urflmr 12:: crinontotion 2:111 int-2.1121102... Lion of the :or’: of other 2.11172 3212.2) "at-ore 192:2 V 21""1 (:11) to 23 1.223170 that the act; 110:? bile salts was due to Ivar-om'oaiem (:1 tom describing: 2.21,- pro 2 rty by -... latch a substance can bring; on 52113012122152 sabotcmce into solution in 1.113023). 22in o:c;{2.o.11'aL.2.on of the rrlaLiono 21:23) 1."- 3 2.123.: 1:22.21 there is a 21.2.[21 fa?" acid concentration, the solution is of a polydiszwrso 1'.-3mm and tho 2'25'22rotro;11c effect. is br011;-2122 about by 221.2 form .:2Lion of a 12121;: or shell of oevorel bile salt molecules around a 3102.11) of fatty acid 1:101 .:culcs in such a way that 1:21»: water 3011222.”. e group 1113 o‘i-x 1.2-: d outwardly. Eli'Lh low conc»..=1:tr:2tion of fatty ac; 22L, 2.216 co: 1;): ex is rule up of 52.1131 :2 fatty 11052.22 1:12:21om11es 2211;170:2212} 02.2 by Gov-rel 2123.222 salt molecxflco. Scot-112.32: 1210.322 partici' can can pass through diffusion membranes, they can be regarded as gv'zzuino colocuhrly dispersr d :78 321213. 23:11:11 229.313.0113 of the amount of bile acids 110:2:3m‘17.‘or 1:" 1e aboorp 23.2011 of fatty acid haw-:2. 112.212. outed a';2px'o:2:2.r.atoly three 2:202 ocules of 11:21: acil to c och mlocule of fatty acid ( 2.212122 would 21‘; p2 2'o::2..'1a’Le 240 gr .113 of {51"00 2202.12: 9.ch for 72v 1112' 50 {TEL 18 of fat). It. is obviogz; that such 9.11 amount (GLISLEY‘LIXC the): 21.2.1 2 00:1.Lair :s a;~.;—:ro::2.2-;1::toly.."’ of the bile salt) is o .11. of ‘rohe (motion, in View of the: fact. that the 12mm]. socmLio: of bile in 112211 is 21.2 rozzimtoly only 530511. (2521' .y (22). Vernal“, however, erg-210121922 that there is obviomly mother 172301201" 0;; 1.122.. 2.11:; in fat absorption. T219 4:212:- ory, sup-gmrtcd vigorous..y by ‘2‘2;-rzar (23), that; the ros;.':'2".1<1235.8 of 22.1"” f‘ 2. from“ 2:2 thy acids 52212.22: humid-.1131 mucosa timough the 1333,0132: limbs: .31’0d‘.30‘tion 01' 9310033310 ’pids, 00.3 first introduced by 85.11.:21111' (24). 3.0 :102- 332.3 “brat: d a (2.2.3 333.001; 0i arm in the couposition of the-.2 pho. 213310... 3.332123 fatty 00:13:; in the intestinal 2.3110000 during thi‘: abs rpt .1011 of f0 . This 0130:3230 3mg; 01332::zwmd 153:: cats mm fax} 0 230-3 rich in u:-'.00turat-:2d fettzr acids. An appreciable increase in the ioniine mmimr of the pi 30:32:23 3031 21‘s of the ink-atlmfl. rmcosa was £017.06 Artom and 00-31013: era (25) 111300th or fod swim .itor-xpimto ntairdng rwiioactiw 3'2i30033210rus. After extraction of the grime-“.1101 pids of differe..t 01:50:30, they i'om'zd a pr-zgzohdcrmzco of the radioactim pho.phorus in the intos 13110.12“ 231130300 01’ liver and kidney. They 03031010'1 th: :11“ r0203. '08 as 0 dimct participation of thr- phos- 233303431 ids in fat. reap-123220330 in the int: stine 02minx 02135:: fed a 0:..-10130 00113000010 in tho 13.10000, not mar-.- 33 a :mbatitution of fatty acid rari- i 00.7. 0 . '3'? «Fm-'3”: a‘w “*1 - v n; M:;0:‘:“ t 52-1-01; 1‘ tJC‘LIISX IV“: ‘LTa-‘l i‘Vau:b One of.’ 1331:: 002'1121‘ views which 0.33.3 1110'ch invatigation was that loucoztzg'tc-s nigh ; 2303.131}; SOI'Vi as in “mule-L0 00.171. rs of lisid ”.0130“: 3.1 from the epithelial. 0013.3 to the lacteals. Clark and C100}: (26) in microscopic 0002:1308 of the 1:233:13 hatic c-ndo $0113.13 in vivo, Show“. that upon injection 01‘ 33.3.1312"; material (olive oil, cream, mid/or egg ~ 1 - “.:. .. . m . .2- , 8‘ JO.-L) 5.23.12 32.- mm 01‘ 3mg lemme (033015,) the lmphaum send out 033102-2330 toward tho {31013010. Lanicsacyi 00 ~00pa>nded quickly to the injected substance by migrating toward it in large mafimrs and 001.ch 1:, on.) wiring 2252:: 1:13:11: :mtarial. 1"“-1' obsomtion imiicatad an emlsii'i 1.20:1 of the 12230... 21.4606. ligsid. 22:32:20.2: of the lipid from the lcrucoq'te to the “Mm 0463100 appoorr‘d to be 0000220022501 by a 05: zmgo to 0 27102220321270 absoroablo fem. 'L’Lstcl 05:500.: :tvdies by Leach (27) produced no 0012125. rm 0me finid-inca, hover-r, that 122200.300 are active in lipid 01330363222325.0212. One of the prim? 2210+ .hoda for stud; .2213 2:520 absorptian med-5312:1331 has born the addition of .222 03001211010 compoznxls, such as H: , a. 4» . 1: 4n .- .v . 4. . h*'1r0~nwrbo..0, 1.0 0..., 1.22.1 fee-0-22;; .0. 21.31.2220 to unfit animals. Bragiley and Gasser (28) fed an {‘11. 3:1 “5.0:! mix'cur of olive 011 021.2 petroleum oil to dogs. The “:11 230010 lymph whi 0112 they obtained by fia'+ul a 00 $012.20; both 0113 in av~r0~dsmu-- the 80:30 prosortion s as in {2310 =r 27222101022 fad. '22051 a 0.2222 20m 2202: mmwt‘d a 2.00%}..521001 absorp- tion of are: 1:0’00 of 011m: 011 and ‘vdrocarbon oil zirturoa. In 1942, {1:230 and Pisa (29) mtmducod a new tcol in the study of 2‘02: absorption, namely the 11014: fie: 1d 211'. cros 003:0. Examina— tion of thoracic 11mph and circulatory blood by dark field illumination mailer the hir .2 pore}: of the 221020000330 .1222 1001701 the pm. n00 of abaorbod fat :23 2153-11120 3=rm103.03, 032202012051 half to one micron in 102201.013 T120301 *1v:..:..-b..cor0 u mg; 0011 0 '20 tzrm "daylorzlcr :20" far the minute 5... "’ 010800 1-0 counting of the 05 1ylomicrons aftnr fat bastion has 01220". 530011 :2 v0.3.2m’2210 122.102; of absorption. 1101'? 21:23 wi :51 the blood of humans, 15220 authors 4521;022:222 0d 2252:; tthe cfzylowim‘ons 1:01‘0 inc 00d fatty 2:1 “hol'al. 12030313213122; 2:023 obs :rv0d: 1. EoLther ,rotozn nor carbofyura;o foods gavo rise to chylozdcrons in the blood . 2. Ingestion of fa uclways provoLod an obs crvablo increase in daylomi crons com: 3. ho chyle emulsion was dotcnziimkd to be fat, chausc: a) Particles roar to the *oj oft “( fluid b) They left a grease Spot on papnr 0) Comic acid, a s lcctive stain for fat, CCUSad a characteristic black stain. d) Fab solub 1m Eyes, such as Sudan IV, were charac Lori stically soluble. o) A qrpical ioLine absorption of fat was observad. f) Glyle mansion snowed the refractive index of fat. The observations of the abovo auti ors causcd than to oliovo that the partiC1fi .mttsr observofi in blood was a To csu7t of absor~l Lion of unhydrolyzod fat in globule form. A.fcu years later, Eollunbo (30), in studies of the G; go tion and absor"tion of fats in cats obsorvod that lipase is not found in fihe cat's int' “tin , thorcby nococsitaii r the absorption of unhyfiro— ZyZLd fat. Ho Lzrther moL mtuinLL than bile so]: a are necessary for the absorption of neutral fat. Eincrcl oil was found not to be absorbed. In 0:331'08‘ 4Lion to the 3a;t;r observation mention and Collinson (31), fOInd tha othc addition of mineral oil to the diet of cats produced an increase in the unscpon fiable fracm on of the liver. They mlggestod that $115 5.3 dun to th colubilizingj offs-ct of bile salts. 13 In 1938, Fraser (18) introduced a new theory of fat nbsomtion which supports the original work of kink, although in modified form. This concept, called the "Partition moor-f", is described as follows: "Fat is absorbed both as a fat id as fatty acid. Fat is absorb-ed by the lactocl—lmphatic route, short-drowning the liver and passes more or loss directl into the fat depot. Fatty acids, however, are absorbed by capillaries directly to the liver via the portal voin.’l He maintained mat if the unsaponii‘iable material incorporated (such as Vitamin A) is soluble in bile salts, it would pass to the liver. Triglycerides, insolublo in bile salts, would pass unchorgoi to the fat depots. During the past four years Frazer has published nmiorons srti 0188 in the fitsld of fat absorption. His original Partition E‘Exsory has been extonied by his 01m work an} the wo '11:: of otnors, not without (iisputc, however. In a con 3’); '"ative stmly, Eraser (32) fed one group of win 5 (albino rats) neutral fat; a second group was fed the some fat with a liberal addition of lipase. Characteristic :31le lactoals were observed in the first group of anims, microns the second group (with added lipase) was ”apical of fatty acid feeding. A second -..~:::perizi‘.<:nt was 1'>s;,~.ri‘omed, in whi ch 8. mixture of olive oil and 1 part of sodizm cowl sulfate to 2-00 parts solution (a lipase inhibitor) was fed. The rats, in which lipolysis was inhibited, absorbed as much fat as the control emimals. Samples of intestinal commute examined. to choc}: for inl'zibition, showed no increase in fatty acids after six hours. It was concluded that absorption occurred .ritho out hydro? 17.213, although sodium atyl cdfatc 130. surface active caspouxl and a good ozm'icii‘ying agent. Eraser mmgcsted that although lipolysie is not accesses? for triglycerd absorption, it affects the widow and fate of! absorbed fatty rats rim . In order mat mflmirolyscd fat be ab .30rb .d fro-:1 the 'ntaslanc, Frazer, Schu‘mon and ”towart (33) bczliev::d that it mist pass thro" 3h the intestinal wall it ..her in a s a c of molecular dispersion, as a dii‘ £23.: :21} 21c c0232]. ex, or in a state of fins;- emulsion. They fowzd :2. evidence in cup-2202.21; of either molecular dispersion or the fomation of a diffusiblc complex. ”he thin. stotz- , that of a fine 02.29.1131on was dc:2011r;t“at23 by 42.: oration of the civic of rats fed. olive oil. The 1‘ chylr- was 3302 d to consist i‘ .in 01:; disperse}, negativdu ch: rgcd globules of olive oil with an avrragr- diameter of 1332;332:912 half 8 micron. This spore 223...... 222299t the 35.2 e of both the ear ticles Seen in the intocidncd. cells and the hyl 0:.ficrons found in the bi cod at th. might of fat ab serption. In support of he necessity of ozziilsification, the absorption of paraffin oil 00:13:12.0: ?y b lieved mobsorbable be cause 01 its inability to form a (Eifi‘us. ...2lc co: 2.12-. : vi t2) bile cal to, was do:i3.2:2.9 crate; to as equal to that of olive oil if it was adrizzisterod in a l‘zijjhly (lief-2421‘scd i‘om. . TEo::1—-3...23:zuls ....ied paraffin oil was not absorbed because of the inability of 2.5320 into 8' incl tract .0 0'le 313' it. In vi tro c2::22c22i*1'22 .... I'-'.V{:3al'.3d that a bile solt/olcic/ amid/ Hemoglyceridc 3323123 m was ci‘i‘cctiv: in producing open ..oncous «AW-.11" .. .- . 1», 1 ”...-.7... £1.10 0.1.31; 11:1 313.. :(‘xJCLl :11111...-_..;-..y .2..:.-,..:.-.... ' .. -.p /. .:., fl [- 1:1ul;.,u;..::.;3. (:1 «.1 {3.0 u) 1).). a 7'. 3.1 7. poLnt: of the vi£C1rco for 5;;cfiyvu sysbsm, ncoawxaw'1ur cwontn;eous "mul”"' “qwoa, :a; fuuni in Lhc i1 vit"o snufy 03 fr 2;" “n1 Saxmons (3K) 3‘101 63:03.;r ted that aijr 3 "1v“ h: r Citospicn of 011v; oil ufbh fanczuatic 11rwsn, thc hydrolysi. U“0c1.ct. were fatty acid, {Eif'TyCariz‘zg can. 210110;""1311;331:219.-;'=; 2:0 ”rm f“ ”c- 1‘01 CAL" bc; £311.11. firzuian 0L1 fir”i1 (2)! :3?" 3 sun tggb 72$;r 301 31 A incnrwmat1143ztlma cflrn oi] :xx:<:f;’:f“7'hbTy ab vrbcfi nxfi lurfl7y star 5 53 Lha liv1r 5%.: 3h y Lake intrcv Lucas injfiction; of In; inacbdv1""“13" ,ral 0:3 C.d.;:xhwm7 thw acténfiizréif‘vwinfls tissxcs, th¢y $112] the 11 r to be 3h fin'n xi}: of sooraf . ‘r3)s;;?ol 3 ans to E: r: 1v d by'uofi ‘f 1“: bil cu is b li:‘u° 93 Le COR" rt‘ to a bile sci ’. “u of cr:;:;.1,'~11"11’ :1 CI; :3..,;1".:' of this: 302:? rxiepzn are $03 jot ‘fiOtn at arc new Erin? luv 2?'*;u;0. \1.f~n) ~ u-o‘I .. .. IQqA‘ . *TT’. ...fi dub xv ..: ....L.“....TJ L’AL’../ .. Q; IE1?! iODOLOGY 1:10:13? stamens of lipid absorption ha.v:~::1 ~ 11:22 1d 1.111003011301110 compounds. menus:- 0f the :- 305:1: W. 0:12 0:: conv 211131100, rmlioacti ergosterol mas prn‘q‘xartfl am! used for a comparative 31:11-3; of the absorjét'ion of corn 011* and mi: 10:00.1 05.1. 3110 .ma..~»o.....i ..‘010 frac bion from tho liver and £103.13 £331.11}: than 2:10:13 rad for activity. SYIIL'. .31.. 3 CF MI) .OACIIVE H’fl‘LsE‘.T.I£QL :5 The 537115310010. rm 1:10;: ctivv ergostxwol was 0000:qu ished by 1 the momod of 110112.011 0.21:} 130.321 (3) using a strain of the yeast E‘accltarot'mcos 03r1::i.:10. The 000011110 was as follows: 1) Yeast was placed on a clezxroac agar slant, grown at. room t<'::::poratzr~.rc for 21, h 1:: 0:; stored in the refrigr era for until used (10:3: than three ucwks old). 2) T': 1:55 ant :0. used in grozrln: the yeast 0 13:3 in a writ nodima, doi‘icicnt in ~>011t0+'w...10 0.10 acid (.chicicnt rugdium I). All inmbations were for 2!. home at 30°C. 0110 100p m; was transfun‘ad to 50 7.11. of medium I and inmzbw 00 with 3:. “sing. 1‘1; volume was then at 0pcd um 00100 1:11., than to 600!“ 1:31., 1503 1:11., and fin ally to 10 \ 4 *E-iazola Oil liters, 03.1 with aeration. 15 1. . .. ...... ,. . 3) The 0011.. 110.0 2.21011 00110000 by c:31:tri;.1ca‘:ion, 1:017 210d .11 3.711310112101127 13022111111 II a... 30°C 1:22.21 aeration. 4) The 0011:: were 030.1211 007! 002.7012 by 00.113123215017102, 11-210th 2721200 tines, placed in 2.210 ace-”.7. 2.0 30°C 1:1 2:21 aura ion for 24. 21.: 2:00. 5) T210 0011:; were then collected washed three- 27103 1:: 2721 :10 272.1? 0:2 water. 6) ”P10 71 grid ..'2.2;:~.~.~ial 1.11.2: obtained from +110 03.113 120.1212: III and incubated at by centrifugati on and ‘I 0y r0flwdn1dth a ~~Lur of arsoluto e—mmolsethyl 0272103: (3:1) for .. 9 «u ur Q3 :- an a V 1 ? 6 hows. The 001m...t 110.: r.-..:ov.d :..... 27.10 02'... I” ‘6‘ 10233.01) 1213233531221 tnroe 2.3.3100. ’2’210 fractions were 0113123121011 and roduwd to 11 0:10.11 volm'le under Vacs-12m in a 002 atr100pl1er0. 7) Thr- real-2111.: 1:210 then 20110:..1‘10 ..b; 1'.-:.. ”1.1131553 221011.251 2; 5'53 0100210110 I‘ll. Distilled water 1.1110 012.206. 170 22011210 the 12012.10 and 172.0 00711131011 03351232002} 1.2127” other in a continuous liquid-liquid ema 2701' for 24 210.23 8) T20h1t210r 17.21:: r0:.:0v-:.d under V0022: :7. :3 pa :3 gr 6 .13 13: +6 and me r00 . 110 611 0:301v:d in 01)? 01:22.: ethanol. 9) The 01002101 solution 2. .3 then 0:22.300.- to an 0300210110 00111271011 of (”12.35.201.111 and 2.21:. C13:- .- 127012-120 was 0011002710 by centrifu- ga‘tion aft0r 12. :10 rs storm: in the rofrig raster. 10) The (31022011240 1.1113 dissolved in warm pyrimno, Nata tr“ 01.211011, t!1011<:‘*210r.1"220 other was rl'r'ovud 0.1122 the aqueous layer extracted ’ ' . + ' ' H v I y ' \ . o 7" ’7' h‘"‘} \" ' V' " ‘."‘." 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" ) .L LI J , ’SJ. . ‘_' 'J . . 1_ . .1 . ~-- - ...-......— f“ ... _. .. *2.) {Y'- \\...il JG" J . -. : 3 ,. . 1 -:1 2;:0 __ 13' ‘0 "’0 2. {:"1. mm / ' k W (" i. '. “I n ‘. r1- 0 1 -‘ ‘4. ‘ 1 .... ..P‘ih)‘ ' t ‘1 VJ— (NJ K—u -o.‘.§ . :) -.. -" .... A." .7 fi 1 :-':‘--1t :zr:-‘.:J-u .1'3l* {.‘d (1 IL...” .... A-s-. «C1 '3- r-rr .54‘._e:’--21: "117435130 ...”) .L. 1:133: 3.1. .45 . . I. I - u ’ ‘ -. ~r- 0.. .‘ -. \ - -nv-: ‘\l 122.34: -11., 4‘ 37:13, 1". co ....= ., (1.0.0., ”w". 1114--.. .0, “r Q . _? . 1 \ f I" 1 27.144 T)-CL‘14.." 3:42.12; :10 8.32.10. ) (3 cl 1 K 1.3.1.1.} 9.1.... ’1 ‘ 1 ... (a "' .7 2 X 1'0 .J—C g..c,.--\4. _ __ ::."_z-:;r:‘ l‘t;-'30r,rc:;€: 3 ._ o X A .1. r 'N . 4!. q H _.I _"' ._]..Y I ' I) D'?! b; I.v-..:.l;c(1 :..-(11.25:. ~19) I ‘wl. V :3 4.)“ - ‘I'.~’ U0 J.‘- "‘ I v~ o “1.3."? . .' .. . . Ar r, “KT! . r :...Jp. C H‘ 1 ‘.;.. ...;1 I"), * £;::1_t {'2’}.3_.'.'.. 11: {).I..) i. 0. LL. ’ O.) u.’ ~ We, W“ q ' . r’fl ~—~ *1 -: ~ ". ,1 :.1" 1011 1 :33. 1 C13 .--. a. _- _ 4, . . . ,. -1 \ ‘ , . - ‘1 ..-._,.. '7 (‘_ 7‘1!“ - n‘ n. l " ’ 1 .. f" '1 7'(‘O . 7 .100. J. ;r.. I. “It,” (J1; 1. \4. J- .) )0), $‘* LID-3 :; {:‘3 . ‘1. I “’: O.“" ‘ .. 1")“; l} “J , LI 1 ’0’ (1 “- ‘Q'!-' )n . a . I- ' ‘ ~ ' ‘ "- " ."I v . . " WU 0.1 ..LI 3 1 .... a- 4 4-..»... 17 7 A a '9 ~83 v-u-q w‘ . ... wva ‘1 ‘ ‘... ‘1 23 (:1... ~ I 1‘.._{‘.'1+ “4“"...4‘4‘ ‘veiLglt-T‘: INT 63110030 5 :31 .....— 0 a -. TM P5“. :r-L " “4. h‘” 11 -.‘wfiué UOfirlur wow 3 A . ......__. ¢ Vt. Dis tilled wabor to l 111}? The accmtzz nodiwz. Indium I Sodimn acetates” 0,1 M Phosphate buffer awful. used with the sodium acetate) __ w_O.l Ii In the first attstmpt to use yeast cells for the biog/1° thesis of ra...oacmv on: or terol, it. was found that; old yeast 0 1.113 13:12} to build up cell wall carbaimiratr rather ‘han lipid material. ;..ccpt for the last stage, the cells had been 3014151 for 24 hours each t e the volume was stepped up. However, beams: of the 101.6; lag; 3.30:: 3’06: rem? icing from the use of the dufici 221: r Li‘zm the final 10 liter volume was incubatot fat. ‘w’hon the corn oil solution was place-ti in the feed about .835 was fowl in the liwr and about .05?) in the depot fat. Similar resulte were obtained with the rats killed a week after receiving the corn oil solution mixed with feed. (See Table II). The mineral oil fed rate Show d less total absorption but significant counts 5-. the d":>0t fat. {ice Table II). The remzlte of both ewerimmts are smmorized in Table III. TABEE i; I (each rat received 10,090 counts/min. f ergosterol) itbliuék. U;;I"0 QJb..:S RAT I30. WEEDII‘ION ILITIAL FIEAL 191413143 PLR 3.1-3833? if .:ICIIT 1‘."..‘133'1‘ "1115(5ij 1 liver corn oil by 255 295 .213 362 pipette 1.3 hrs. fat .030 0 2 liver corn oil by 250 240 .173 ..83 pigmtte [.8 hrs. fat .023 0 3 liver mineral oil by 269 303 .215 3’7 pipette 1.3 hrs. I. liver mineral oil by 231 254 .194 120 pipette [.13 hrs. 5 liver corn oil in food 224 260 .196 110 1 week fat .028 0 6 liver mineral oil in 256 315 .225 16/. food 413 hrs. fat .031. 30 7 liver mineral oil in 239 2‘35 .211 159 fat .031 34 8 liver corn oil in food 234 254 .196 72 43 hrs. fat .026 2 9 liver corn oil in food 250 275 .167 92 l Wok fat .025 6 10 liver corn oil in food 231 214. .130 141 [’8 hrs. fgt .016 Q (each rat reocivcd 10,030 counts min. of fifjHSfiCrOl) A . v1: Li; ..‘f ... 'I-b LU U. . -S RAT I} . ‘EDI'SIOH IICIL‘IAL FIEIIL 1713.73.35 PA: 2.7. ..IGCEJ.‘ 7.51131 21‘ 1",. .ICE If 3 131.1113 11 liver' corn oil by 272 201 .196 93 pipette 24 hrs. fat 0025 7 12 liver corn oil in feed 250 250 .194 58 21. hrs. fat .022 2 13 liver corn oil in food 271 262 .195 100 24 hrs. fat .03 8 14 liver mineral oil in 241 233 .192 92 feed 24 hrs. fat .020 20 15 liver mineral oil in 25 235 .191. 43 feed 24 hrs. fat . ” 13 16 liver corn.oil by 259 251 .193 317 pipette 21 hrs. fat .023 0 17 liver corn Oil in feed 245 2431. .203 73 1 wodk fat .024 3 13 liver min ral oil by 256 2A1 .193 Ill pipette 21. hrs. fat .020 17 19 liver corn oil in fuod 270 291 .214 85 l undk fat .031 6 23 livur i'filliiril oil by 24.0 21.3 .219 79 pipette 24 hrs. h 341.. 25 TABLE ” III (SKEW ' '1' is: or}. out {“1 .211 52) 0024331113253 TIM." (1:12:13 34:: 11‘": (_;1) AV§1QS Liver Gbrn oil by nipetto 24 398 317 357 Liver " " " “ 48 362 233 322 Fat " " “ " 21. 7 0 3.5 Fat " " " " 43 0 O 0 Liver Cbrn oil in feed 24 58 100 79 Liver ” “ " " 48 72 141 106 Fat “ " ” " 21. 2 8 5 Fat " " " " 1.3 2 O 1 Liver “ " " " 1 week 92 110 73 85 90 Fat " " " " l wool: 6 0 3 6 1. Liver Mineral on by 22-, 111 79 93 Pipette Liver " " " " [.3 37 120 '79 Fat " ” " ” 2/. 17 25 21 Fat " " " " 48 15 20 17 Liver Kineral oil in feed 24 96 48 72 Liver " " " " AB 164 159 161 Fat " " " " 21. 20 13 16 E151 u~v n n f“, 48‘- i.£§é:::::::éé&t_ ‘32 * T1120 be “’51) “ {st-Q +'~ a u -.L U}... and the killing: of the rat-s. C\ 43 fgam‘iing or a~’-"fif:iStti"'ri2’1f? of rrfiioactive (rgostoml DISCUSSION A " 1 Q (7 ‘,1-\"‘." --y-‘ l A p-\ ‘,- J-‘~r—.-" ‘v w-fi-‘z~ ». '5' 7 1 . I"‘. “[\-l -'.- .q‘ Y ' \HL-‘a‘.l'—'.-A' 24'.“qu ) .L I‘LLLLJ '9':ka '~.-.- 4‘ —"-..- .L‘O .. ".:.-l .:. uk'. uf) Jul. AA, I L;(:O . ‘ ‘L .’ t. .5. ... 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' ‘ ‘V‘5' " “(5.. '1 f" 7' ’: 0‘ “r! '. 9 ‘ ....L: ..;-..}I‘ COLu- 0-3.11 01110; 1.....- u “Ls/J. -:.. Lluvs 4:19.!) ‘1 3 1‘.. .:.‘7 (3111131uy fa 5‘; revealed a "\ the method of artflnistration influences the absorption and deposition of ergosterol in certain tissues. mien rats were killed 21. hours after receiving the radioactive ergosterol by the some methods slightly more was found in depot fat but less in the liver. This would imicate toznporsnr storage of some of the ergosterol in depot fat ard probably in other ti mum anti sub- sequent trmlsfer to the liver. than the corn oil preparation was incorporated in the food and one wee-3: elapsed before the rats were killed, there was still con- siderable activity in the liver and slight activity in depot fat. This would iJrlicate that either ergosterol or some metabolite of ergosterol was still presemt in these tissues a week after the ingestion of the test done. In the case of the mineral oil there was less actiflty show in the liver except when the oil was mixed with the food. However, an appreciable activity was demonstrated in the depot fat. That some minral oil was absorbed and carried to both the liver and, fat depots is indicated by the higher content of the unsaponifiable matter. ’afl'zat is perhaps difficult to ozqzlain is that although the total absorption of :“ineral must be rolatiwly small, the activity found in the dr'pot fat was actually larger than in the case of the corn oil. This can only mean that on the basis of the amount of oil absorbed, Mneral oil appears to mm with it much more of the ergosmvrol. There is as yet; no basis for accomtting: for the $22.11 absorption of ergosterol when dissolved in corn oil . 3O {1' J. . .’ -L' . . . ‘ ' .. - ..- . n .L‘. _ ‘ z‘ 1‘ fl“ ‘ .1 '\ . 3‘. n, w.. erw "r . .‘ ‘ (a .1'\ ‘l‘ a f -; - Q". L. 3 LI. '3 O.)~~.?. JS 6 - —'—l. J. - vul- \A "D V J II J~JI \- ... u) .;. 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S cicnco 14, 579 (l‘Z-Wl) (1.8.-.- .. , $871101 J3, A auk'LL 22.21.,1. .13 2:10 (151) 011121 10:: of 1’.: :70 and D; i. - 4.1.1.1.»: T—h—T *wvcclwi: Liv q a b,- Q01:72 (1945) 00 J), (‘4. .‘ ‘1 1 '.. ..'.1 “fi in L.) ‘41er; . .L.. f‘: ...‘IJ ..ifll.‘ , .1 .1 4.2, f 1.2 .1 A3371" t 9:: in We 191.1110 v” Michigan State College East Lansing, Michigan ‘_1 - -‘ «___—— — ___—_— 'I'HE USE OF RADIOACTIVE ERGOS'CEROL IN THE STUDY OF LIPID ABSORPTION IN THE ALBINO RAT BY ROBERT DOUGLAS I~IACKEIJZIE AN ABSTRACT Submitted to the School of Graduate Studies of Michigan State College of Agriculture and Applied Science in partial fulfillment of the requirements for the degree of MASTER OF SCIE‘I CE Department of (hemistry Year 1954 APproved C} Q . ) f/‘k/V‘fl’ L54 V. ABSTRACT The mechanism for fat absorption has been the subject of extensive l) fats are completely study. Two theories have been proposed: hydrolyzed by lipolytic enzymes to fatty acids and glycerol before absorption, and 2) fats if highly enough dispersed may be absorbed without hydrolysis. Both theories are supported by experimental evidence. In 1938 Frazer (1) proposed a modified theory, according to which fatty acids are absorbed into the capillary system and then transported to the liver; whereas, the glyceryl esters are absorbed into the lymphatic system and eventually reach the fat depots. To shed more light on the subject it has been suggested that tracers be used so as to follow the path and conditions of absorption. The present study involved the use of radioactive ergosterol dissolved in corn oil and in mineral oil. t was found by Hanahan and Wakil (2) that ergosterol is absorbed when dissolved in corn oil. They showed that the major portion absorbed went to the liVer but small amounts were also found in other organs. The results of the present investigation indicate that when the corn oil containing radioactive ergosterol was dispersed in feed less absorption occurred than when given orally to rats deprived of food for 24 hours. In the case of mineral oil which is known to be slightly absorbed through the intestinal wall less ergosterol was absorbed. However, some was found in the depot fat; whereas, little or none was found in the depot fat when corn oil was used. The effect of the dispersion in the feed was the same as when corn oil was used, less being absorbed. The results of this study are disappointing in that they do not apparently lend support to any of the current theories of fat absorption. It probably means that radioactive ergosterol is not a good compound to use for this purpose. Significant, however, is the observation that in the case of mineral oil appreciable amounts of radioactive ergosterol were found in depot fat. On the basis of the relative amounts of the oils absorbed, mineral oil showed a much greater capacity to transfer the ergosterol to the circulatory system. References: l) Frazer, A.C., Fat4§bsorption and Metabolism. Analyst ég, 308, (1938). 2) Hanahan, D.J. and S. Walzil, Stu___dy on _Arbsorption and IvIetaCbngicsm of Ergosterol-Ol4. Arch. Biochem. and BiOphys. 4,4, 150 (1953) . "WilliWills “w «’1 2—. -- we mmmocmazm -v.-r——~—.P __,,l ,- 'vJ‘l .-. ( ~~ -.K 'L v .- a _ .. ' 3‘ v‘ b -. .. ’ ' - - .‘ ’1'“ U'."‘ a “ ME“ I ‘ Iii-""- ;) '- - v‘-"’V " ' ‘“- 1' 'u “.-‘-“/‘l m- -J. nu. V». ..J- .... , _ .a- u' ... , .:.a'- .C . '0 g . 1' - \-.‘,.- 4;- w '..J .:.. '- 71". ‘ . ' H I ...-an 1.x. - a .. .-.-.-. ~.—-o_vor_-...m~ ' ‘r a - - - -.~ ."s ‘- ' . ." ‘T- , '7 " i"- A 1 . .. \I,-o--.-. . l.’ 1 s a,_, - 1? I - ‘ ‘ .' ave—- ~. - ...- “.--.“ .- .m , ‘ . "‘ ' . , "“ — V . ’ ‘ I ’ 2 w .__ v - ~ u - 1__ 3 {‘1 ' - . . . . I (. 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S 5100112101.: :1, R. ., 1.:11 63111111131. 10;-3 1:1 11:11:13.- .- science 14, 579 (1: 31) I”- 3'" 1110:, 30311101 5., A ”Stud: of Tat AM, :11 111' 0:11: the 57111371 whnL.’ '1';1(‘,~3_’Lg EILC (19;)1) -—..-V" - ».. k .P'» ”990 Michigan Stator Co ing, Michigan of Lane Ea THE USE OF RADIOACTIVE MOSTEROL IN THE STUDY OF LIPID ABSORPTION IN THE ALBINO RAT By ROBERT DOUGLAS I~IACKENZIE AN ABSTRACT Submitted to the School of Graduate Studies of Michigan State College of Agriculture and Applied Science in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Chemistry Year 1954 Approvecl C Q )fiff/J’ L51 II ABSTRACT The mechanism.for fat absorption has been the subject of extensive study. Two theories have been prOposed: l) fats are completely hydrolyzed by lipolytic enzymes to fatty acids and glycerol before absorption, and 2) fats if highly enough dispersed may be absorbed without hydrolysis. Both theories are supported by experimental evidence. In 1938 Frazer (l) prOposed a modified theory, according to which fatty acids are absorbed into the capillary system and then transported to the liver; whereas, the glyceryl esters are absorbed into the lymphatic system and eventually reach the fat depots. To shed more light on the subject it has been suggested that tracers be used so as to follow the path and conditions of absorption. The present study involved the use of radioactive ergosterol dissolved in corn oil and in mineral oil. It was found by Hanahan and Wakil (2) that ergosterol is absorbed when dissolved in corn oil. They showed that the major portion absorbed went to the liver but small amounts were also found in other organs. The results of the present investigation indicate that when the corn oil containing radioactive ergosterol was diapersed in feed less absorption occurred than when given orally to rats deprived of food for 24 hours. In the case of mineral oil which is known to be slightly absorbed through the intestinal wall less ergosterol was absorbed. However, some was found in the depot fat; whereas, little or none was found in the depot fat when corn oil was used. The effect of the dispersion in the feed was the same as when corn oil was used, less being absorbed. The results of this study are disappointing in that they do not apparently lend support to any of the current theories of fat absorption. It probably means that radioactive ergosterol is not a good c0111pound to use for this purpose. Significant, howeVer, is the observation that in the ease of mineral oil appreciable amounts of radioactive ergosterol were found in depot fat. On the basis of the relative amounts of the oils absorbed, mineral oil showed a much greater capacity to transfer the ergosterol to the circulatory system. References : l) Frazer, A. 0., Fat Absorption and Metabolism. Analyst £13, 308, (1938). 2) Hanohan, D.J. and S. Wakil, Study_on Absorption and I—Ietabolism of Frgosterol-Guf. Arch. Biochem. and Biophys. 54, 150 (1953). HICHIGQN STATE UNIV. LIBRQRIE