Studies on the site of fat absorption

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1 Gut, 1961, 2, 168 Studies on the site of ft bsorption 2 blnces fter resection of vrying mounts of the smll intestine in mn Fronm C. C. BOOTH, D. ALLDIS, AND A. E. READ the Deprtments of Medicine nd Chemicl Pthology, Postgrdute Medicl School oflondon SYNOPSIS This pper demonstrtes tht in mn, s in the rt, incresing mounts of ft rech more distl levels of the smll intestine s the dietry lod increses. Opinions hve vried s to the site of ft bsorption in the smll intestine of experimentl nimls (Bernrd, 1856; Frzer, 1943; Kremen, Linner, nd Nelson, 1954; Benson, Chndler, Vnsteenhuyse, nd Ggnon, 1956; Turner, 1958; Booth, Red, nd Jones, 1961). It is now cler tht the site where ft is normlly bsorbed depends on the dietry lod. When smll mount of ft is given to rt, for instnce, bsorption tkes plce lmost entirely in the. If lrger quntities re fed lrger mounts re bsorbed in the, but t the sme time n incresing proportion of the ingested ft escpes bsorption in the upper intestine nd psses on into the ileum where bsorption then lso occurs (Booth et l., 1961). The site of bsorption of ft in mn is uncertin. Borgstrom, Dhlqvist, Lundh, nd Sjovll (1957), using n intestinl intubtion technique, hve climed tht ft is bsorbed in the proximl 1 cm. of the. This finding, however, conflicts with the clinicl dbservtion tht stetorrhoe my follow resection of the ileum (Cox, Meynell, Cooke, nd Gddie, 1958; Klser, Roth, Tumen, nd Johnson, 1959). It seemed possible tht in mn, s in experimentl nimls, the site of ft bsorption might depend on the mount of ft fed. In order to test this hypothesis, we hve studied ft blnces in group of ptients who hd undergone resection of vrying mounts of the smll intestine. The ptients were grouped ccording to whether the proximl or distl smll intestine hd been resected, since experimentl studies in nimls suggest tht jejunl resection is less likely to disturb intestinl function thn resection of the ileum (Jensenius, 1945; Kremen et l., 1954). blnces were first crried out using diets 168 contining between 3 nd 75 g. dily. The results of these blnces indicted how much of the smll intestine is required to bsorb moderte mounts of ft normlly. Incresing ft diets were then given to four ptients, for if the site of ft bsorption is relted to the dietry ft, it might be expected tht incresing ft diets would cuse incresing degrees of stetorrhoe in ptients with intestinl resections. PATIENTS STUDIED blnces were crried out in seven ptients who hd undergone resection of vrying mounts of the distl smll intestine (Cses 1 to 7) nd two ptients who hd resections of the proximl intestine (Cses 8 nd 9). An pproximte estimte of the length of intestine resected in ech ptient ws obtined from the surgicl opertion specimens nd by subsequent brium followthrough exmintion. Clinicl detils, including the dt from which the extent of the resection in ech ptient ws ssessed re given in the Appendix, nd the lengths of intestine either resected or ssumed to be remining in the different ptients re given in Tbles I nd II. MATERIALS AND METHODS Ptients were studied in metbolic wrd. The diets contining from 3 to 1 g. of ft hve been described in detil by King nd Wootton (1956). Where dditionl ft ws given, it ws invribly dded in the form of butter ft. The mount of ech diet which ws rejected ws nlysed. ANALYSIS OF FAECAL FAT EXCRETION The methods were those described by King nd Wootton (1956). After preliminry equilibrtion period of three dys on ech diet, the stools were collected for two consecutive threedy periods into suitble metl continers. The specimens were first weighed nd homogenized nd smple of this Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.

2 RESULTS MODERATE DIETARY INTAKES The dietry intkes, fecl ft excretion, nd estimted ft bsorption from diets contining moderte mounts of ft (between 3 nd 75 g. dily) re shown in Tbles I (distl resections) nd IL (proximl resections). Distl resections The results of the ft blnces in the seven ptients who hd undergone resection of vrying mounts of the distl smll intestine (Cses 1 to 7) re given in Tble I nd illustrted in Fig. 1. Two ptients (Cses 1 nd 2) hd no significnt stetorrhoe (Fig. 1); these two ptients hd lost the ileocecl vlve together with only 6 or 8 ft. of the distl ileum. Studies on the site offt bsorption TABLE I FAT BALANCES DURING TWO CONSECUTIVE THREE-DAY PERIODS AFTER RESECTION OF DISTAL SMALL INTESTINE Cse No. 2 Amount of Intestine Resected 6 ft. of terminl ileum 8 ft. of terminl ileum 3 1 ft. of ileum 4 Ileum nd ll but proximl 6 ft. of 5 Ileum nd ll but proximl 6 ft. of 6 Ileum nd ll but proximl 4 ft. of 7 Ileum nd ll but proximl 8 in. of Dietry Fecl Absorbed * The ptients with lrger distl resections hd stetorrhoe nd its degree depended on the extent of their resection. One ptient (Cse 3), who hd undergone resection of pproximtely 1 ft. of the ileum hd mild stetorrhoe, excreting 9 5 nd 9.7 g. ft dily during successive blnce periods (Fig. 1). Two other ptients (Cses 4 nd 5), both of whom Cse no ii I & ~~. ~~~ W Dietry ft Fecl ft bsorbed 6 h 5s. 4 F 3b A- E 2h 1 Amount resected feet,feet 1 feet of terminl - -- ileum homogente ws then dried. This dried liquot ws treted with hydrochloric cid nd extrcted with ether, the extrcted ft being determined grvimetriclly. The men fecl ft excretion during ech of the three-dy periods ws then clculted. Using this technique, control subjects usully excrete less thn 6 g. of ft dily when tking diets contining between 5 nd 1 g. of ft. The difference between the dietry ft nd the ft excreted ws ssumed to be the mount bsorbed. While this clcultion does not tke into ccount the smll mount of ft in the feces tht my be of endogenous origin, it gives resonble pproximtion of ft bsorption. b~- All but proximl 6 feet 6 feet 4 feet 8 inches M,... *... ^... i FIG. 1. Dietry ft, fecl ft excretion, ndft bsorption during successive three-dy blnce periods in ptients who hd undergone resection of vrying mounts of the distl smll intestine (Cses 1 to 7). The interrupted line indictes the upper limit of norml fecl ft excretion. 169 Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.

3 17 hd more extensive distl resections, hd more mrked stetorrhoe; these ptients excreted between 11- nd 22. g. of ft dily. However, even though there might be severe stetorrhoe fter lrge distl resection, the proximl ws cpble of bsorbing considerble proportion of the dietry ft. Although only 4 ft. of the proximl remined in Cse 6, for instnce, more thn 4 g. of ft ws bsorbed by this ptient from dily intke of 62 g. (Cse 6, Fig. 1, Tble I). Furthermore, nother ptient (Cse 7), who hd only the duodenum nd proximl 8 in. of the remining, bsorbed s much s 15 g. of ft from dily intke of 3 g. (Fig. 1, Tble I). Proximl resections The results ofsimilr blnces in the two ptients who hd undergone resection of the proximl smll intestine re given in Tble II nd illustrted in Fig. 2. The first of these ptients (Cse 8) hd lost the proximl 8 ft. of the ; the dily fecl ft excretion ws 6.2 nd 6.8 g. during two successive blnce periods on diets contining 48-9 nd 46-7 g. of ft (Fig. 2, Tble II), indicting tht most of the dietry ft ws bsorbed (Tble II). The resection in the second ptient ws very much more extensive, only 15 in. of the terminl ileum remining. This ptient hd moderte stetorrhoe, excreting 12.1 nd 13.4 g. of ft dily, but he bsorbed more thn 35 g. of ft from the short segment of remining distl smll intestine. FIG. 2. Dietryft,feclft excretion, ndft bsorption during successive blnce periods in ptients who hd undergone resection ofthe proximl smll intestine (Cses 8 nd 9). The interrupted line indictes the upper limit of normlfeclft excretion. C. C. Booth, D. Alldis, nd A. E. Red TABLE II FAT BALANCES DURING TWO CONSECUTIVE THREE-DAY PERIODS AFTER RESECTION OF PROXIMAL SMALL INTESTINE Cse No. Amount of Intestine Resected Dietry Fecl Absorbed 8 8 ft. of proximl All nd ll butdistl lsin.of terminl ileum 'Blnces were crried out during two consecutive four-dy periods in this ptient. EFFECT OF INCREASING DIETARY FAT The results of the blnces when incresing ft diets were given re lso shown in Tbles I nd II. Distl resections Incresing ft diets (between 3 nd g. dily) were given to three of the ptients who hd undergone distl resections (Cses 2, 4, nd 5). The results of the two successive three-dy blnce periods on ech diet re given in Tble I. The men fecl ft excretion during the totl sixdy period of ech diet is illustrted in Fig. 3, together with the corresponding mount of ft bsorbed. One of these ptients (Cse 2) hd lost only the terminl 8 ft. of the ileum nd hd no stetorrhoe when tking diet contining 63 g. of ft dily. However, when the dietry ft ws incresed to or g. dily, the men dily fecl ft excretion rose to 13 nd 16 g. respectively (Fig. 3, Tble I). Although this ptient developed stetorrhoe when tking the higher ft diets, the ctul mount of ft bsorbed incresed s the dietry ft ws incresed (Fig. 3). The two other ptients (Cses 4 nd 5) hd pproximtely 6 ft. of the proximl remining nd both hd stetorrhoe when receiving diets contining 5 g. of ft (Fig. 1). One of these ptients (Cse 4, Fig. 3) ws given diets contining 3, 5, nd 1) g. of ft dily. As the dietry ft ws incresed, there ws progressive increse in fecl ft excretion but t the sme time the mount of ft bsorbed from the remnnt of proximl smll intestine lso incresed (Fig. 3, Tble I). Similrly in the other ptient (Cse 5, Fig. 3) there ws n increse inboth feclftexcretion nd ft bsorption when the dietry ft ws incresed from 5 to 1 g. dily (Fig. 3, Tble I). Proximl resections Incresing ft diets (from 5 to g. dily) were lso given to the ptient who hd undergone resection of 8 ft. of the proximl Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.

4 o lso so 3,,. 'I. 1 CL Co. L Pr 1 Cose 2: Resection of 8 feet of terminl ileum 7 A ~ ~ ~ ~ ~ ~ ~~osnw Studies on the site offt bsorption FAT ABSORBED 4: 6 feet of proximl remining Cost5L 6 feet of proximl remining _ Dietry ft - 9. per dy FIG. 3. Effect of incresing ft in diets on ft bsorption nd excretion in ptients with resections of the distl smll intestine (Cses 2, 4, nd 5). L.. 1. hh F i 2 ID 11h Cse 8: Resection of 8 feet of proximl 7 FAT ABSORBED FAT EXCRETED ' pif, O_ 5 1 Dietry ft - 9. per dy FIG. 4. Effect of incresing ft in diets on ft bsorption nd excretion. I1. The results of the blnces re given in Tble II nd illustrted in Fig. 4. This ptient excreted men of only 6-5 g. of ft dily during the six-dy blnce when he received diet contining just less thn 5 g. of ft (Fig. 1). When the dietry ft ws incresed, there ws slight increse in fecl ft excretion (Fig. 4) but the men mount of ft excreted dily when the diet contined s much s g. of ft ws only 8-2 g. As shown in Fig. 4, the mount of ft bsorbed by this ptient incresed progressively s the dietry ft ws incresed (Tble II). DISCUSSION 171 The results in the ptients subjected to distl resections (Cses 1 to 7, Fig. 1) show tht in mn the cn bsorb ft well. They lso indicte tht lthough the proximl ileum my be required for completing the bsorption of ft (Cse 3, Fig. 1), the distl 6 to 8 ft. do not pper to be necessry when moderte diets re given, even when the ileocecl vlve hs been resected (Cses 1 nd 2, Fig. 1). The effects of incresing the dietry ft in three of these ptients suggest tht the smll intestine of -A- Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.

5 172 mn responds to n incresed dietry lod in the sme wy s tht of the rt (Booth et l., 1961), for the bsorbed more ft when the dietry ft ws incresed. Two of these ptients (Cses 4 nd 5, Fig. 3) hd only 6 ft. of the proximl smll intestine remining, nd there ws progressive rise in ft bsorption with increse in the dietry intke (Fig. 3, Tble I). However, the incresed ft bsorption in these ptients ws only chieved t the expense of n incresed fecl ft excretion (Cses 4 nd 5, Fig. 3). This suggests tht, s lso occurs in the rt, incresing mounts of ft my rech more distl regions of the smll intestine when the dietry ft is incresed. The results in the third ptient (Cse 2) re in keeping with this concept. This ptient hd lost only the terminl 8 ft. of the ileum nd her remining intestine ws cpble of bsorbing norml mounts of ft from moderte dietry intke (Fig. 1). However, when greter mounts of ft were given, the fecl ft excretion incresed nd stetorrhoe developed (Fig. 3), suggesting tht some ft my rech the distl ileum nd be bsorbed there when the dietry intke is lrge. Like the upper, the terminl ileum ws shown to bsorb ft well. Cse 9, for instnce, hd only the distl 15 in. of the ileum remining, yet he bsorbed more thn 35 g. of ft from this remnnt of the distl smll intestine (Fig. 2). Similrly, Cse 8, who hd lost the proximl 8 ft. of the, bsorbed ft lmost normlly when given moderte dietry intke (Fig. 2). In this ptient, incresing the dietry ft resulted in progressive increse in bsorption nd cused only slight increse in the fecl ft excretion (Fig. 4); this contrsts with the stetorrhoe produced by similr diets in the ptient with comprble distl resection (Cse 2, Fig. 3). Repeted studies hve demonstrted tht when norml persons re given diets contining between 3 nd 35 g. of ft dily, there is slight increse in the fecl ft excretion, which my rech s much s 1 g. dy on the highest dietry intkes (Wolleger, Comfort, nd Osterberg, 1947; Annegers Boutwell, nd Ivy, 1948), but in terms of the mount of ft bsorbed there is progressive increse in ft bsorption s the dietry intke is incresed. Msterton, Lewis, nd Widdowson (1957) found tht two polr explorers were ble to bsorb 263 nd 276 g. of ft respectively from diets contining 273 nd 285 g. of ft dily, diets which were designed to meet their high energy requirements. The results reported in this pper provide rtionl explntion for this pprently unlimited cpcity of the humn smll intestine to bsorb ft. When moderte dietry intke is given, it seems likely tht ft bsorption occurs predominntly in the C. C. Booth, D. Alldis, nd A. E. Red (Fig. 1). As the dietry lod is incresed, the mount of ft bsorbed by the increses (Fig. 3). At the sme time incresing mounts of ft escpe bsorption in the (Fig. 3) nd presumbly pss on into the ileum, where bsorption my then normlly occur, for, s shown by the results in Cse 8, the distl intestine is lso cpble of bsorbing incresing mounts of ft when presented with n incresed lod nd very little ft psses through it unbsorbed (Fig. 4). The ileum therefore ppers to become progressively more importnt s site of ft bsorption when the dietry ft is incresed. Since both rid ileum re cpble of bsorbing ft well, nd since both respond to n incresed dietry lod with n incresed bsorption (Figs. 3 nd 4), it is not surprising tht in mn the mount of ft which cn be bsorbed is lmost unlimited. bsorption must therefore be controlled by the ppetite of the individul nd not by ny intrmucosl mechnism such s tht which regultes the bsorption of vitmin B12 (Glss, Boyd, nd Stephnson, 1954; Swendseid, Gsster, nd Hlsted, 1954; Booth, 1958). In conclusion, it ppers tht in mn, s in the rt, the is the mjor re of ft bsorption. As fr s ft is concerned, the ileum seems to represent the min reserve of the smll intestine which, like other essentil orgns such s the liver nd kidney, is equipped to del with fr more thn bsic metbolic requirements. Although the ileum my be of reltively minor importnce when moderte mounts of ft re tken, the results given in this pper suggest tht it is likely to be of greter significnce to those who enjoy grgntun ppetites. For individuls such s polr explorers who must tke high ft diet to meet high cloric requirements (Msterton et l., 1957), functioning ileum is probbly essentil. SUMMARY blnces were crried out in group of ptients who hd undergone resection of vrying mounts of the smll intestine. When moderte ft diets were given, resection of 6 or 8 ft. of the ileum did not interfere with ft bsorption. Resection of lrger mounts of the distl intestine cused stetorrhoe but considerble proportion of the dietry ft ws bsorbed when only 4 to 6 ft. of the proximl intestine remined nd some ws bsorbed even when ll but the duodenum nd proximl 8 in. of the hd been resected. The results of giving incresing mounts of ft in the diet to ptients with resection of the distl smll intestine indicte tht the responds Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.

6 to n incresed dietry lod with n incresed bsorption, but incresing mounts of ft escpe bsorption nd re excreted in the feces when the dietry ft is incresed in such ptients. These observtions suggest tht in mn, s in the rt, incresing mounts of ft rech more distl levels in the smll intestine s the dietry lod increses. It seems likely tht the in mn is the mjor site of ft bsorption when moderte diets re given, but the ileum probbly becomes progressively more importnt with increse in the dietry ft Ḟt ws bsorbed well by ptient with resection of 8 ft. of the proximl even when diets of up to g. of ft were given. The terminl ileum cn bsorb ft, for ptient with only 15 in. of the terminl ileum remining bsorbed more thn 35 g. from diet contining 48 g. of ft. We wish to thnk Dr. F. Avery Jones for permission to crry out studies in Cse 8, nd Dr. Brbr Clyton for her kindness in llowing us to quote the results in Cse 9. We lso thnk the physicins nd surgeons of Hmmersmith Hospitl for their coopertion. APPENDIX Clinicl detils nd results of other intestinl function tests in mny of these ptients were given t the Interntionl Congress of Gstroenterology t Leyden, 196 (Booth nd Mollin, 1961). DISTAL RESECTIONS CASE 1 A 3-yer-old mn developed gngrene of the terminl ileum s result of volvulus. Approximtely 6 ft. of the terminl ileum ws resected (surgeon's estimte), the ileum being nstomosed to the cecum. A brium follow-through demonstrted norml pttern of the smll intestine but very rpid trnsit, brium reching the cecum within 15 minutes. CASE 2 A 36-yer-old womn hd intermittent dirrhoe, pssing two to three wtery motions dily. Eight yers before, lprotomy hd reveled Crohn's disese of the terminl ileum nd pproximtely 4 ft. of the terminl ileum nd the scending colon were removed. A recurrence two yers lter necessitted resection of further 3 to 4 ft. of terminl ileum, end-to-end ileo-trnsverse nstomosis being performed. A recent lprotomy for n obstruction due to n dhesion hd shown wellfunctioning ileo-trnsverse nstomosis nd no evidence of Crohn's disese. A brium follow-through reveled norml smll bowel pttern, brium pssing into the colon in two hours. CASE 3 A 4-yer-old mn hd 2-yer history of tuberculosis involving first the right nkle joint nd then the terminl ileum nd cecum. Approximtely 1 ft. of Studies on the site offt bsorption the distl intestine hd subsequently been removed (surgeon's estimte), the smll intestine being nstomosed to the trnsverse colon by side-to-side nstomosis. A brium follow-through reveled norml pttern of the residul smll intestine nd the brium pssed into the colon in two hours. There ppered to be smll blind end of the terminl ileum but there ws no stsis within this loop. CASE 4 A young womn ged 28 developed extensive gngrene of the distl smll intestine cused by volvulus. The ffected prt of the gut ws resected, the surgeon estimting tht pproximtely 6 ft. of the proximl intestine remined. End-to-side jejunocecl nstomosis ws performed. A subsequent brium follow-through reveled norml pttern of the remining smll intestine, nd brium reched the cecum within 2 minutes. CASE 5 A 3-yer-old womn hd first developed Crohn's disese of the terminl ileum ssocited with peri-rectl bscess 1 yers go. Resection of the distl smll intestine hd been crried out on four occsions. At her lst opertion, two yers before, blind loops of the distl intestine were resected nd the ws nstomosed end-to-end to the trnsverse colon. The surgeon estimted tht pproximtely 6 ft. of the remined. There ws no evidence of Crohn's disese t this time. A brium follow-through reveled norml pttern of the residul smll bowel but the brium hd pssed into the colon by three qurters of hour. CASE 6 An imbecile hospitl ptient ged 24 yers hd developed volvulus of the smll intestine cusing extensive gngrene two months before. The right side of the colon, together with ll but the proximl 3 to 4 ft. of the (surgeon's estimte) were resected, endto-end nstomosis being performed between the upper nd the trnsverse colon. A brium followthrough ws hmpered by some dely in gstric emptying, but the pttern of the remining smll intestine ws norml nd brium pssed into the colon within one nd hlf hours. CASE 7 An 8-yer-old widow hd developed thrombosis of the superior mesenteric rtery cusing extensive gngrene of the smll intestine. The ileum nd ll but the proximl 8 in. of the were resected, but jejunl remnnt being nstomosed end-to-end to the scending colon. A brium follow-through reveled slight dilttion of the residul smll intestine; some of the brium hd reched the colon within five minutes. (This cse hs been reported in detil by Hrrison nd Booth, 196.) PROXIMAL RESECTIONS 173 CASE 8 A 71-yer-old mn hd extensive jejunl diverticulosis requiring resection of 8 ft. of the proximl, the opertion specimen being mesured fter the opertion. End-to-end nstomosis ws performed. A brium follow-through two yers lter reveled slight dilttion of the bowel immeditely distl to the nsto- Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.

7 174 C. C. Booth, D. Alldis, nd A. E. Red mosis but no other bnormlity. Brium hd pssed into the colon within four hours. CASE 9 An 18-month-old child hd undergone three successive opertions on the smll intestine for ileo-ilel intussusception. Extensive resections hd been crried out, the stomch eventully being nstomosed to the terminl ileum. The mount of ileum remining ws estimted to be 15 in. (surgeon's estimte). A brium follow-through eight months lter reveled slight dilttion of the remining smll bowel; brium reched the colon in one nd three qurter hours. (This cse hs been reported in detil by Clyton nd Cotton, 1961.) REFERENCES Annegers, J. H., Boutwell, J. H., nd Ivy, A. C. (1948). The effect of dietry ft on fecl ft excretion nd subjective symptoms in mn. Gstroenterology, 1, Benson, J. A. Jr., Chndler, G. N., Vnsteenhuyse, F. E., nd Ggnon, J.. (1956). Studies concerning the site of ft bsorption in the smll intestine of the rt. Ibid, 3, Bernrd, C. (1856). Memoire sur le pncres et sur l role du suc pncretique dns les phenomenes digestifs, prticulierement dns l digestion des mtieres grsses neutres. Billiere, Pris. Booth, C. C. (1958). An Experimentl ndclinicl Study of the Intestinl Absorption of Vitmin B,2. M.D. Thesis, University of St. Andrews. nd Mollin, D. L. (1961). The effect of resection of the smll intestine in mn. Proc. Internt. Conf. Gstroenterol., Leyden 196. Red, A. E., nd Jones, E. (1961) Studies on the site of ft bsorption. 1. The sites of bsorption of incresing doses of "3'I-lbelled triolein in the rt. Gut, 2, 23. Borgstrom, B., Dhlqvist, A., Lundh, G., nd Sjovll, J. (1957). Studies of intestinl digestion nd bsorption in the humn. J. clin. Invest., 36, Clyton, B. E., nd Cotton, D. A. (1961). A study of mlbsorption following resection of the entire nd the proximl hlf of the ileum. Gut, 2, 18. Cox, E. V., Meynell, M. J., Cooke, W. T., nd Gddie, R. (1958). The folic cid excretion test in the stetorrhe syndrome. Gstroenterology, 35, Frzer, A. C. (1943). Differentition in the bsorption of olive oil nd oleic cid in the rt. J. Physiol. (Lond.), 12, Glss, G. B. J., Boyd, L. J., nd Stephnson, L. (1954). Intestinl bsorption of vitmin B,, in mn. Science, 12, Hrrison, R. J., nd Booth, C. C. (196). Mssive resection of the smll intestine fter occlusion of the superior mesenteric rtery. Gut, 1, Jensenius, H. (1945). Results of experimentl resections of the smll intestine on dogs. Nyt nordisk Forlg, Copenhgen. Klser, M. H., Roth, J. L. A., Tumen, H., nd Johnson, T. A. (1959). Reltion of Smll Bowel Resection to Nutrition in Mn. In Proceedings of the World Congress of Gstroenterology, Wshington Vol. I, pp Willims nd Wilkins Co., Bltimore. King, E. J., nd Wootton, I. D. P. (1956). Micro-nlysis in Medicl Biochemistry. 3rd ed. Churchill, London. Kremen, A. J., Linner, J. H., nd Nelson, C. H. (1954). An experimentl evlution of the nutritionl importnce of proximl nd distl smll intestine. Ann. Surg., 14, Msterton, J. P., Lewis, H. E., nd Widdowson, E. M. (1957). Food intkes, energy expenditures, nd fecl excretions of men on polr expedition. Brit. J. Nutr., 11, Swendseid, M. E., Gsster, M., nd Hlsted, J. A. (1954). Limits of bsorption of orlly dministered vitmin B12; effect of intrinsic fctor sources. Proc. Soc. exp. Biol. (N. Y.), 86, Turner, D. A. (1958). The bsorption, trnsport nd deposition of of ft. Amer. J. dig. Dis., n.s. 3, (Prt 1) , (Prt 2) Wolleger, E. E., Comfort, M. W., nd Osterberg, A. E. (1947). Totl solids, ft nd nitrogen in the feces: III. A study of norml persons tking test diet contining moderte mount of ft; comprison with results obtined with norml persons tking test diet contining lrge mount of ft. Gstroenterology, 9, Gut: first published s /gut on 1 June Downloded from on 28 June 218 by guest. Protected by copyright.