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Dietary hypercholesterolemia in relation to cholesterol and fat absorption in cockerels. Lindsay, Owen Burnett 1963

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DIETARY HYPERCHOLESTEROLEMIA  IN RELATION  TO CHOLESTEROL AND FAT ABSORPTION I N COCKERELS  by  Owen B u r n e t t  Lindsay,  B.S.A.  A THESIS SUBMITTED IN P A R T I A L FULFILMENT OF THE REQUIREMENTS  FOR THE DEGREE OF  MASTER OF SCIENCE I N AGRICULTURE  in  t h e Department of  Poultry  We  accept  this  Science  t h e s i s as c o n f o r m i n g  standard required  from  candidates  to the  f o rthe  d e g r e e o f MASTER OF SCIENCE IN AGRICULTURE.  Members o f t h e D e p a r t m e n t o f  THE UNIVERSITY OF B R I T I S H  April,  1963  COLUMBIA  In presenting  t h i s t h e s i s i n p a r t i a l f u l f i l m e n t of  the requirements f o r an advanced degree at the U n i v e r s i t y of B r i t i s h Columbia, I agree that the L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and m i s s i o n f o r extensive purposes may  study.  I f u r t h e r agree that per-  copying of t h i s t h e s i s f o r s c h o l a r l y  be granted by the Head of my Department or  his representatives.  I t i s understood that copying or  by publi-  c a t i o n of t h i s t h e s i s f o r f i n a n c i a l gain s h a l l not be allowed without my w r i t t e n p e r m i s s i o n .  Department The U n i v e r s i t y of B r i t i s h Columbia,. Vancouver 8, Canada.  ABSTRACT  Differences levels  of adult  i n t h e mean p l a s m a c h o l e s t e r o l  White  Single  Comb L e g h o r n c o c k e r e l s  a c c e n t u a t e d when a d i e t c o n t a i n i n g oil  a n d 1% c h o l e s t e r o l was  levels  by  f e d f o r s i x days.  (p 0.05) w i t h t h e amount  the groups.  the  10% hydrogenated  rate  The d i f f e r e n c e s  negatively  of c h o l e s t e r o l  excreted  o b s e r v e d among g r o u p s i n  o f e l i m i n a t i o n o f t h e excess c h o l e s t e r o l from t h e  c i r c u l a t i o n were n o t f o u n d t o be s t a t i s t i c a l l y Variation  vegetable  The c h o l e s t e r o l  p r o m o t e d b y t h e d i e t were f o u n d t o be  correlated  were  i n cholesterol absorption  responsible,  i n large  may  significant.  therefore  be  measure f o r d i f f e r e n c e s i n t h e degree  of h y p e r c h o l e s t e r o l e m i a  i n d u c e d , by f e e d i n g  a diet high i n  cholesterol.  A significant  c o r r e l a t i o n (p 0.01) b e t w e e n t h e  amounts o f c h o l e s t e r o l a n d s a p o n i f i a b l e following the feeding suggests  that  t h e amount  major d e t e r m i n a n t  in  of cockerels  high-cholesterol  an i n c r e a s e  d i e t over  in lipid  410 d a y s o f f e e d i n g  excreted  high-cholesterol  o f d i e t a r y f a t a b s o r b e d may  o f t h e amount  Subjection fat,  of a high-fat,  lipids  of c h o l e s t e r o l  to the feeding  Cockerels  maintained throughout  be a  absorbed.  of a  a prolonged period  absorption.  diet  high-  resulted  which  the t e s t , a  survived lower  mean plasma c h o l e s t e r o l vessels of survivors evidence  l e v e l than n o n s u r v i v o r s .  compared to n o n s u r v i v o r s  of a t h e r o s c l e r o s i s .  The b l o o d  showed  little  ACKNOWLEDGEMENT  The his  author takes  gratitude  Department used  to Professor  of Poultry  i n this  study  encouragement. B.  this  Jacob  Science  Biely,  t o express  Head o f t h e  for providing  and f o r h i s constant  the f a c i l i t i e s  interest  and  I n a d d i t i o n t h e a u t h o r wants t o thank Mrs.  E . March o f t h e Department  supervision  opportunity  and a s s i s t a n c e  of Poultry  throughout  Science  for her  the course  of the  experiments.  Further Dr.  C. W.  Roberts  his  valuable  he w i s h e s t o e x p r e s s h i s a p p r e c i a t i o n t o o f t h e Department  criticisms.  of Poultry  Science f o r  TABLE OF  CONTENTS  Page  INTRODUCTION  1  LITERATURE REVIEW  . . . .  Characteristics  Which Render F a t s  The  Degree o f S a t u r a t i o n  The  Unsaponifiable  Essential  Fatty  Cholesteremic  . . . . . . . .  of  of Cholesterol  Absorption,  5  Concentration  Carbon Chain Length o f G l y c e r i d e Regulation  Levels  Excretion  4 4  Fat Content  Acid  .  4  6  Fatty  Acids  .  7  by t h e P r o c e s s e s  and S y n t h e s i s  . . . .  7  Cholesterol  Absorption  7  Cholesterol  Excretion  9  Cholesterol  Synthesis  10  Mechanisms R e s p o n s i b l e Cholesterol  f o r the Promotion o f  Absorption  by D i e t a r y  Fats  . . . .  EXPERIMENT I  13  Experimental Selection  13 of Experimental  Birds  Methods Analysis  12  . . . . . . . .  13 15  of Feces  16  Ether-Extractable Unsaponifiable  Lipids  Lipids  16 . . . . . 16  Page  Total  Cholesterol  Chromic  Oxide  Plasma C h o l e s t e r o l Results EXPERIMENT  16 17 .  and D i s c u s s i o n II . . . . . . '  17 18 23  Introduction  23  Experimental  23  Results  24  and D i s c u s s i o n  TABLES (EXPERIMENTS I a n d I I )  28  FIGURES (EXPERIMENTS I a n d I I )  38  SUMMARY AND CONCLUSIONS  47  BIBLIOGRAPHY  50  1  INTRODUCTION  The with  the  a s s o c i a t i o n of hypercholesterolemia  liberal  led  to  an  may  confer  advanced  consumption of  intensive search  from c h a r a c t e r i s t i c s the  constituent  these the as  acids  fat.  the  nature  of a c t i v i t y  type  and  cholesterolemic  potencies  these  Cholesterolemic  discovery  An advancement limiting  example of the  being,  that  by  solubility  in this theory  dietary fat.  a number o f  among c e r t a i n f a t s  cholesterolemic  effects  of  continue of a f a t  directed  measure o f c h o l e s t e r o l e m i c  direction,  appear  This  i s the theory  arguments, large  their  would potency.  is  the  which states that  factor i n cholesterol absorption  been d i s f a v o u r e d  of  characteristics  of c h a r a c t e r i s t i c s which  of e f f o r t s  of c h o l e s t e r o l i n the  derived  a t t r i b u t e d e x c l u s i v e l y to a s i n g l e  attention  a more r e l i a b l e  been  component  activity  Hence many w o r k e r s h a v e now  provide  be  factors  characteristic. to the  thus  may  unsaponifiable i n these  h o w e v e r , a l w a y s be  which  s a t u r a t i o n of  as w i d e l y  interest.  of f a t s  has  concentration  degree of  of the  fats  E v i d e n c e has  vary  of major  cannot,  the  type  a c i d s , the  Because f a t s  in their  t o be  s u c h as  fatty  and  activity.  this  man  more s a t u r a t e d  for characteristics  cholesterolemic  indicating that  the  in  the  the  solubility has  most  however recent  differences in  disproportionate to  the  2 slight  differences i n the solvent  a c t i o n of these  fats  on  cholesterol.  The  inconsistent cholesterolemic  f a t s when f e d t o v a r i o u s by  this  species  t o the d i e t a r y intake  recognized tion,  deserves  important  fat-induced  tibility  their  the species fats,  degree o f u t i l i z a -  i n studies  also,  concerned  with  of a difference i n  there  to  t o which adult  diet  has suggested  may be v a r i a t i o n i n t h e  undertaken t o determine  White L e g h o r n c o c k e r e l s response  high-cholesterol diet.  type  hypercholesterolemia,  l e a d i n g t o v a r i a t i o n i n Suscep-  s t u d y was t h e r e f o r e  hypercholesterolemic  high-fat,  the well  hypercholesterolemia.  This  fat  evidence  of various  to  extent  with  the f a c t o r of d i g e s t i b i l i t y  of s t r a i n s of poultry  that within  of t h i s  i n their  by a h i g h - f a t , h i g h - c h o l e s t e r o l  utilization  in  coupled  cholesterolemia.  susceptibility  the  c h o l e s t e r o l l e v e l s are  of fats  considerations  More r e c e n t  also,  that  v a r i a t i o n among f a t s  i t appears that  induced  are a l s o not accounted f o r  theory.  In view o f evidence related  effects of several  may  vary  to the feeding  of a  In addition, the r e l a t i o n  of v a r i a t i o n to the u t i l i z a t i o n  of the dietary  a n d c h o l e s t e r o l was t o be i n v e s t i g a t e d . It  animals,  was c o n s i d e r e d  cockerels  appropriate  t o use as  experimental  s e l e c t e d f r o m two s t r a i n s shown p r e v i o u s l y  3  to d i f f e r i n s u s c e p t i b i l i t y t o d i e t a r y h y p e r c h o l e s t e r o l e m i a . The e x p e r i m e n t a l d i e t was f o r m u l a t e d t o c o n t a i n 10% o f a h i g h l y s a t u r a t e d f a t ( c o n t a i n i n g f a t t y a c i d s whose c h a i n l e n g t h d i d not exceed 18 carbon atoms) and 1% c h o l e s t e r o l . The p r e s e n t study was a l s o concerned w i t h the effect  o f long-term f e e d i n g of the e x p e r i m e n t a l d i e t on the  c h o l e s t e r o l l e v e l of the b l o o d o f c o c k e r e l s and upon t h e development of a t h e r o s c l e r o s i s i n these  birds.  4  LITERATURE REVIEW  I  C h a r a c t e r i s t i c s which Render F a t s a)  The  Degree of  Kinsell were among t h e  Saturation  and  Michaels  first  to  s a t u r a t i o n are  regulating  blood  saturated the et  ( 1 9 5 5 ) and  not  the  feeding  f a t , produced higher  feeding  of  of  levels  of  Kinsell coconut  the  were i n v e r s e l y r e l a t e d .  Michaels  o i l ,a  highly  dietary fats  al.,  Kinsell  and  Ahrens  showed t h a t  Steinberg,  1958;  the  dietary fat  a l . , 1956; et  Ahrens  al.,  by et  1958  others).  The observed For  and  than  These f i n d i n g s have been c o n f i r m e d et  Avigan  i n man fat.  i o d i n e number o f t h e  numerous i n v e s t i g a t o r s ( B r o n t e - S t e w a r t 1957;  in  and  cholesterol levels  six different and  a l . (1955) varying  soybean o i l , a h i g h l y u n s a t u r a t e d  a l . (1955) u s i n g  cholesterol  fats  et  metabolically equivalent  cholesterol levels. that  Ahrens  demonstrate that  degrees of  (1955) r e p o r t e d  Choleresterolemic  cholesterolemic  i n man  instance,  are  when t h e  cholesterolemic (Biely  and  corn  i n other  were f e d a n o r m a l p o u l t r y  o i l t h e y were r e n d e r e d  protein level when t h e  March,  certain fats  always r e p r o d u c i b l e  when c h i c k e n s  supplemented w i t h lemic  not  e f f e c t s of  1959).  of the  diet  protein level The  reports  was  was  as species. diet  hypercholestero20%  and  r a i s e d to  hitherto  normo— 26%.  mentioned  5 describe  the e f f e c t s  of corn  o i l on man  as  hypocholestero-  lemic.  b ) The U n s a p o n i f i a b l e  Peterson mixed p l a n t 0.5%  Fat Content  ( 1 9 5 1 ) showed t h a t when c h o l e s t e r o l a n d  sterols  (at 1  a n d 1% o f t h e d i e t ,  hypercholesterolemia,  : 1 r a t i o ) were f e d t o c h i c k e n s t h e r e was  hepatic  a marked  lipidosis  and  Plant  sterols  have  experimentally  induced  hypercholesterolemia  al., and  1954; man  and S w e l l  1958,  protection against cholesterolosis.  a l s o b e e n shown t o  inhibit  i n rats  Farquhar  e t a l . , 1956;  Farquhar  r e t a i n s some o f i t s h y p o c h o l e s t e r o l e m i c  ing hydrogenation, et  of vegetable  Wood evidence vitamin  oils  t o show t h a t  saponifiable  The  Chaikoff,  depressant  (1960) have  presented  activity  between  of t h e un—  oils.  of experimentally sterols  followBeveridge  be some r e l a t i o n s h i p  of c e r t a i n marine  inhibition  1954;  may  corn  unsaponifiable f a t content.  and h y p o c h o l e s t e r o l e m i c  content  inhibition  activity  of the c h o l e s t e r o l  to their  there  c h o l e s t e r o l e m i a by p l a n t the  part  ( 1 9 6 0 ) a n d Wood and B i e l y  A content  that  have l e d J o n e s e t a l . (1956) and  a l . (1957) t o a t t r i b u t e  action  and  and o t h e r s ) .  These f i n d i n g s added t o t h e o b s e r v a t i o n oil  (Swell et  e t a l . , 1956), r a b b i t s ( P o l l a c k , 1958),  ( P o l l a c k , 1956;  Sokolow,  as  induced  hyper-  h a s b e e n shown t o be due t o  of c h o l e s t e r o l absorption  (Hernandez and  a n d H e r n a n d e z e t a l . , 1953) b u t t h e p r e c i s e  6 manner i n w h i c h t h i s i s a c c o m p l i s h e d r e m a i n s One  explanation  offered  with cholesterol  for  D u n c a n and  1956;  the  Best,  necessity  for  cholesterol.  the  1958  acceptor  for  are  or  that  itbinds  on  for  Essential  determinant  of  the  and  acids  10  20%  of  the  levels.  on  the  cast  doubts  on  1957),  intestine  and making  1953).  of  of  percentage  to  as  a  rats  a  the  fatty  role  fatty of  related acids  saturated  S i n c l a i r (1957) i n  ex-  hypercholestrolemic acids.  Ahrens et  such a r e g u l a t o r y  this factor  essential  r e c o g n i z e d the  investigations  mucosal  demonstrated that  K i n s e l l and  in essential  The  the  d i e t were i n v e r s e l y  the  effect  low  ex-  cholesterol for  Upon f e e d i n g  f a t they  percentage  diet.  and  Other  within  the  mucosa  a l . , 1958  Green,  (Pollack,  also  diets  and  lumen o f  periments w i t h r a t s of  either  the  not  Concentration.  a r a c h i d o n i c ) and  i n the  et  into  compete w i t h  a l . (1957) i m p l i c a t e d  or  of  esterification is  Swell  (Clover  1951;  postulated  absorption  been d i s c r e d i t e d .  i n the  acid  This  cholesterol  1955;  absorption  Fatty  of  lipoproteins  membrane  to  that  compete  (Peterson,  a l . , 1954).  sterols  l e v e l s reached  product  (Linoleic fatty  (Pihl,  cholesterol  containing  cholesterol to  passage  cholesterol  Hegsted et  diet  et  sterols  esterase.  evidence  plant  the  its cell  unavailable  c)  the  that  s i t e s on  cell  it  Swell  t h i s e x p l a n a t i o n has  planations  plant  esterification prior  i n t e s t i n a l wall  b),  that  cholesterol  In view of  a prerequisite of  was  obscure.  of  a l . (1959) however, these  acids.  These  7 w o r k e r s h a v e shown t h a t (e.g. at  f a t s low i n e s s e n t i a l f a t t y  menhaden o i l ) may d e p r e s s  l e a s t t h e same e x t e n t  acid  content  d)  (e.g.  as f a t s w i t h a h i g h  evidence  t o show t h a t  in  Fatty  produced  the carbon chain  length  When t h e p r e d o m i n a n t  fatty  c a r b o n atoms i n l e n g t h  of the blood.  of their acids  constituent  effect.  Absorption,  a)  Cholesterol  Excretion  with rabbits"and  further observed  fatty  acids.  Levels and  their  When t h e y  were  the hypercholesrespectively.  by t h e P r o c e s s  Synthesis  Absorption  Dubach and H i l l  poration  and i n t e r m e d i a t e  of Cholesterol  of  greatly  o f t h e g r o u p were 12 t o 14  8 t o 10 a n d 16 t o 18 c a r b o n atoms i n l e n g t h  Regulation  slight  (1962).  t h e f a t s were f o u n d t o e x e r t  e f f e c t was s l i g h t  fatty  Similar  groups o f f a t s v a r y i n g  most p r o n o u n c e d h y p e r c h o l e s t e r o l e m i c  II  only  h a v e b e e n made i n t h e d o g b y G r a n d e  i n v e s t i g a t o r f e d three  terolemic  Acids  i n man s a t u r a t e d  atoms i n l e n g t h ,  changes i n t h e c h o l e s t e r o l l e v e l  This  fatty  e t a l . (1959) and Hashim e t a l . (1960)  8 t o 10 c a r b o n  observations  essential  corn o i l ) .  Beveridge  acids  serum c h o l e s t e r o l l e v e l s t o  Carbon Chain Length o f G l y c e r i d e  have p r e s e n t e d  acids  (1946) and B o l l m a n  ( 1 9 5 1 ) who w o r k e d  r a t s r e s p e c t i v e l y i n d i c a t e d that  the incor-  o f f a t i n t o a n e s s e n t i a l l y f a t - f r e e d i e t p r o m o t e d no increases  i n the absorption  on t h e f a t - f r e e d i e t .  o f c h o l e s t e r o l beyond  Blomstrand and Ahrens  that  (1958)  8 investigated  the absorption  patient  c h y l u r i a who  her  with  urine.  of labelled cholesterol  shunted 40% of absorbed f a t i n t o  These workers  observed that  20% of the adminis-  tered  c h o l e s t e r o l was a b s o r b e d  given  s i n g l y or i n combination with f a t .  i n fifteen  led  c h o l e s t e r o l was f o u n d t o o c c u r  six  hours f o r t h e f a t .  absorption lation  constituent  thoracic  cholesterol  whether  The p e a k o f l a b e l -  a t n i n e h o u r s compared t o  i n man,  duct.  of t h e i r test-meal  esterified  hours  H e l l m a n e t a l . (1958) s t u d i e d t h e  of radiocholesterol  of the l e f t  in a  directly,  by cannu-  A l t h o u g h f a t was n o t a  they found that  i n t h e lymph i n c r e a s e d ,  the l e v e l of reaching a  peak i n s i x h o u r s .  Sano McCullagh the  (1924),  a n d Cook  (1939) have d e m o n s t r a t e d  presence  cholesterol  absorption.  cholesterol  absorption.  fats play This  s u p p o r t e d by e v i d e n c e  terol  absorption  1954;  Daskalakis,  and dogs)  consequently  an i m p o r t a n t  role i n  view i s i n contrast  t o that  mentioned i n v e s t i g a t o r s . that  dietary  i n the r a t (Albrink 1955, a n d B r o k e t t  fats  absorption  Iti s  enhance  e t a l . , 1955;  choles-  Morris,  e t a l . , 1934), and t h e  ( B i e l y a n d M a r c h , 1959; a n d Wood a n d B i e l y ,  The  that  i s obligatory to  These workers have  e x p r e s s e d by t h e p r e v i o u s l y  chicken  (in rats  of f a t i n the intestines  advanced t h e view that  well  ( 1 9 3 6 , 1 9 3 8 ) a n d Cook a n d  1960).  r e l a t i o n s h i p between f a t and c h o l e s t e r o l  becomes c l e a r e r when t h e l i p i d animals  (instead  content  feces  of experimental  terol  l e v e l ) i s u s e d as a measure o f c h o l e s t e r o l  of the  of the blood  choles-  absorption.  9 For  e x a m p l e , when G r a n d e  high-eholesterol excretion the  of  diet  steroid  he  Stanley  dogs a h i g h - f a t ,  observed that  an  substances  e x c r e t i o n of f a t .  o f Cheng and  (1962) f e d t o  increase  accompanied  These r e s u l t s ( 1 9 5 6 ) who,  are  using  an  i n accord  rats,  dietary fat intake  amount  of c h o l e s t e r o l s e c r e t e d ,  reabsorbed  indicate  that  absorption 1952;  Pihl,  to  free fatty a greater  1955;  extent  Swell,  1954;  and  to  cholesterol absorption  chicken  b)  (Stamler  Wilson  diets fat  et  Cholesterol  cholesterol  fatty  c e b u s monkey  of the  i n the  (Kim  and  Ivy,  et a l . , 1955).  The  rat  however,  ( L i n et a l . ,  a l . , 1 9 5 7 ) , and  S i p e r s t e i n (1958)  corn  the  oil.  administered  to three  groups of r a t s ,  oil,  lard  30%  They o b s e r v e d  administered  Hellman et  to  a c i d s have been r e p o r t e d  alpha-hydroxy, n e u t r a l s t e r o l s  25%  the  Excretion  30%  w h i c h were f e d c o r n  and  These  large  i n the  and  essentially  increases feces  sterols  prefed  in  no  the  of those  rats  a c c o u n t e d f o r 17  radioactivity.  a l . (1957) i n a s i m i l a r  experiment  14 injected diets  a  a l . , 1957).  and  respectively.  Swell  (Partman et  intravenously  containing  those  excreted.  seem  than glycerides  saturated  1955), the  with  acids potentiate cholesterol  long.chain inhibit  and  o f numerous e x p e r i m e n t s  in  demonstrated  r e l a t i o n s h i p between the  results  the  increase  direct  The  in  cholesterol-C  containing butter  intravenously or  corn  oil.  i n t o humans The  quantity  prefed of  to  10 labelled the  s t e r o l s i n the  serum c h o l e s t e r o l  feces  were f o u n d  concentrations  to  increase  when  decreased with corn o i l  feeding.  The not  observation  a f f e c t the  great  extent  w i t h the  excretion  foregoing  unsaturated  level  i n j e c t i o n of  When t h e  previously  described  bile  Bronte-Stewart  fed  et  following  oil.  cholesterol  excretion,  acids  accord  results  in rats  of  Haust  and  as  in  a  s i m i l a r to  those  Thus B y e r s  and  excretions  fed  and  decrease  i s used  of  cholesterol  coconut Beveridge  oil.  bile and  walnut  These  (1958)  re-  and  (1956).  Synthesis  Migicovsky  i n vivo  and  (1958) o b s e r v e d t h a t  in vitro  was  oleic  acid while  synthesis  acid.  i n vivo  was  synthesis  e n h a n c e d by  corn o i l , rapeseed o i l , erucic  lauric  in  any  Nicolaysen,1961).  containing  and  to  intraperi-  diets  oil  do  elimination  the  ( G r a n and  of b i l e  cholesterol  al.  Cholesterol  Wood and of  rats  the  shown i n t h e  have been o b t a i n e d .  agree w i t h those  c)  of  cholesterol  than i n r a t s  sults  i s also  (1958) have o b s e r v e d g r e a t e r  and  i s also  fats accelerate  exretion  cholesterol  acids  fats  circulating cholesterol  codliver  measure o f  Friedman  more s a t u r a t e d  Bauman, 1 9 5 7 ) ,  circulating cholesterol  toneal  the  findings.  plasma c h o l e s t e r o l  oil  of  ( C o l e m a n and  That of  that  acid  d e p r e s s e d by  feeding and coconut  11  On et  the  other  a l . (1958) have  rent  difference  between r a t s showed t h a t This  i s an  hand H i l l  a l . ( 1 9 5 8 ) and  demonstrated that  i n the  fed  et  rate  saturated  dietary  fats  essential  of  t h e r e was  synthesis  and  of  unsaturated  not  any  appa-  cholesterol fats.  enhance c h o l e s t e r o l  agreement w i t h the  Linazosoro  They  also  synthesis.  findings  of  Ahrens  (1957).  Murkherjee the  effects  oil  or  30%  greater  of  and  feeding  hydrogenated  increase  i n the  A l f i n - S l a t e r (1958) rats  diets  cottonseed rate  of  investigated  containing oil.  15%  cottonseed  They o b s e r v e d  synthesis  i n animals  a  fed  cottonseed o i l .  When c h o l e s t e r o l marked s u p r e s s i o n ( G o u l d and al.,  1957).  that  the  level  of  of  Taylor,  degree  of  dietary  There  Frantz  et  has  diet  supression  is directly  of  been  a l . , 1954;  also  rats  observed  and  Morris  advanced evidence related  to  to  et show  the  cholesterol.  i s thus  levels  quantity  f a t s are  cholesterol  synthesis  the  ( 1 9 5 1 ) has  cholesterol of  hepatic  1950;  Gould  i s added to  little  i n d u c e d by related  synthesis.  evidence feeding to  indicating different  changes i n the  that  types rate  of  and  12 III  Mechanisms  Responsible  Cholesterol  Swell that  Absorption  by D i e t a r y  o f exogenous  absorption  fats  i f adequate q u a n t i t i e s of b i l e  included  the  a c t i v a t i o n of c h o l e s t e r o l esterase a l . , 1953  i n the d i e t .  a, 1953  b).  quently  suggested  augment  the a b s o r p t i o n  flow  of b i l e  cholesterol reservations when h i g h  that  Swell  by b i l e  and F l i c k  salts  (1955) have  t h e manner i n w h i c h e x o g e n o u s  esterase.  Pihl  concerning  terol. the  potency  to salts  (Swell consefats  o f c h o l e s t e r o l i s by s t i m u l a t i n g t h e increase  the a c t i v i t y  ( 1 9 5 5 ) h a s however  the importance  of t h i s  of  expressed  his  mechanism  of f a t are f e d .  W i l k e n s e t a l . (1962) has a t t r i b u t e d t h e terolemic  shown  T h i s h a s b e e n shown t o be due t o  which i n turn w i l l  levels  (1957) have  i s n o t an a d j u n c t  are  et  Fats  e t a l . (1955) and Vahouny  the presence  cholesterol  f o r the Promotion of  of f a t s  They o b t a i n e d  to t h e i r  a high  cholesterol solubilities  serum c h o l e s t e r o l e f f e c t s  in  solvent  correlation  a c t i o n on  choles-  (+ 0.982) b e t w e e n  of the various man.  choles-  fats  and  their  13  EXPERIMENT I  Experimental  Selection  of Experimental  Twenty-four selected  from  third  generation  disorders  shown  (Biely  hypercholesterolemia  to d i f f e r  and March,  (Biely  strain  starting during  B) s u b s e q u e n t  the starting  and March,  1960).  T h e y were k e p t  i n battery  cages  and a t t h e end o f  g r o w i n g p e r i o d were t r a n s f e r r e d t o f l o o r  the  basal  I.  A a n d 40  were f e d s i m i l a r  and growing p e r i o d s  of Table  induced  The c o c k e r e l s  strain  the  diet  White  1958) and d i e t a r y  to hatching  and growing d i e t s .  Comb,  were  i n susceptibility to  f r o m w h i c h s e l e c t i o n s were made (35 f r o m from  cockerels  e a c h o f two s t r a i n s o f S i n g l e  Leghorns p r e v i o u s l y renal  adult  Birds  pens and f e d  Selection of the  experimental  b i r d s was c a r r i e d o u t a s f o l l o w s .  The fed  seventy-five  ad l i b i t u m , t h e b a s a l  cholesterol three  levels  days f e e d i n g  b i r d s o f s t r a i n s A a n d B were diet  of Table  o f t h e s e b i r d s were of this  diet.  These  I.  determined  ( a t s i x d a y s ) were made i n o r d e r  following  l e v e l s were  d e t e r m i n e d f o l l o w i n g s i x days o f f e e d i n g . tions  The p l a s m a  Repeat  to increase  again  determinathe r e -  liability  o f t h e c h o l e s t e r o l l e v e l s w h i c h w e r e t o be u s e d a s  the  f o r s e l e c t i n g and g r o u p i n g t h e e x p e r i m e n t a l  basis  birds.  14  From e a c h s t r a i n those w i t h t h e extreme for  levels  are given  of birds  from  those of b i r d s  range  but o n l y  highest  and l o w e s t  The six  I I and I I I .  s t r a i n A ranged from strain  B, f r o m  a f t e r the e l i m i n a t i o n  being valuable  lots  similar  cholesterol levels. i n t h i s study  birds  from  containing  generation The  grouping  are  as  126 — 194 mg %•  112 - 185 mg % .  Due  of birds with the T h e s e were  i s also  regarded included.  assigned  of birds with  to a shortage  the r e p l i c a t i o n of l o t s  of birds  The  h a v e b e e n made  and were t h e r e f o r e  a n e q u a l number  cockerels  Cholesterol  e a c h s t r a i n were t h e n  cholesterol levels.  levels  The c h o l e s t e r o l l e v e l s o f  i n Tables  from  (including  cholesterol  i n the l e v e l s o f both groups c o u l d  similar  as  and i n t e r m e d i a t e  t h e s t r a i n ) were s e l e c t e d .  these b i r d s  and  twenty-four birds  into  fairly  of t h i r d  was n o t p o s s i b l e .  shown i n T a b l e s  I I and I I I and  follows.  Lot  Mean  Cholesterol  Pretreatment Strain  1  133.7  2  149.0  3  156.5  4  165.8  5  175.3  6  187.0  Level(mg.%)  Strain  A  B  121.0 0  The cage u n i t s .  lots  156.5 158.2 162.5 169.8 181.3  were t h e n p l a c e d  Each u n i t  consisted  into  four  of three  individual  tiers,  each  15 capable  of housing  allocated into  t o one l o t , b i r d s  two c o m p l e t e  terol  four birds,  levels  w i t h one t i e r  s t r a i n were  housed  The mean p r e t r e a t m e n t (e.g. l o t s  n o t as s i m i l a r  choles-  A 1 and B 1)  a s was d e s i r a b l e .  None-  i t was c o n s i d e r e d a d v a n t a g e o u s t o compare t h e h y p e r — responses  were a s u n i f o r m  randomizing  the lots  the remaining  lots  were a d j a c e n t  t o each  was t h a t a  that  each  of corresponding lots  cholesterolemic that  from  cage u n i t s .  were i n a f e w c a s e s theless  such  lots  as p o s s i b l e . from  i n such  the s t r a i n s  of these  T h i s was a c c o m p l i s h e d  one s t r a i n a manner  other.  under c o n d i t i o n s  and then that  The r e s u l t  alternated  by  distributing  corresponding of this  lots  arrangement  i n f o u r cage u n i t s  placed i n  s i n g l e row.  Following  t h e assignment  the b a s a l d i e t  of Table  vegetable  1% c h o l e s t e r o l  oil,  T h e y were a l l o w e d f r e e  of lots  t h e b i r d s were f e d  I supplemented w i t h a n d 1% c h r o m i c  10%  hydrogenated  oxide  (a marker).  access t o f e e d and water.  Methods On t h e t h i r d and  and s i x t h  plasma c h o l e s t e r o l  levels  Rosenthal  e t a l . (1957).  bird  a twenty-four  over  d a y , f e c e s were  determined-by  withdrawn from  the diet  dried.  from  hour p e r i o d , p o o l e d f o r each  At t h e end o f t h e s i x t h  f e c e s were  t h e method o f  The f e c e s w e r e c o l l e c t e d  freeze-dried.  drawal  collected  and a f t e r  again c o l l e c t e d ,  Plasma c h o l e s t e r o l  l o t and  day c h o l e s t e r o l  f o u r and seven p o o l e d by l o t s  each  was  days o f w i t h and f r e e z e -  d e t e r m i n a t i o n s were a l s o  made  again  16 at  these  times.  Analysis  a)  of Feces  Ether-Extractable Extraction  of f e c a l  carried  out i n a s o x h l e t  extract  was f i l t e r e d  and  the ether  Lipids lipids  with  d i e t h y l ether  apparatus f o r s i x t e e n hours.  using  Whatman number 42 f i l t e r  e v a p o r a t e d on a 5 0 ° C w a t e r b a t h .  The  was d r i e d a t 8 5 ° C f o r one a n d a h a l f h o u r s a n d t h e n in  a dessicator  b)  Unsaponifiable  the " O f f i c i a l  the  following  The  Methods o f A n a l y s i s "  ether  recovered  described  gave a n e g a t i v e  s u l f a t e and t h e  i n a manner s i m i l a r t o t h a t  previously  lipids.  Cholesterol  freeze-dried  a mixture  thejunsaponifiable  sodium  f o r the ether-extractable  c) T o t a l  of alcohol test  dure o f R o s e n t h a l  samples  of feces  acetone  ( 1 : 1 ) until  were  extracted extracts  f o r cholesterol with the rapid  e t a l . (1957).  determined according procedure  cooled  o f t h e A.O.A.C. w i t h  solution containing  lipids  with  residue  modification.  was d r i e d w i t h a n h y d r o u s  The  paper  Lipids  lipids  described  The  p r i o r t o weighing.  T h e s e were d e t e r m i n e d b y t h e p r o c e d u r e in  was  Total  to the revised  proce-  c h o l e s t e r o l was  S p e r r y Webb  (1950)  i n which the p r e c i p i t a t e d d i g i t o n i d e i s reacted  with a sulfuric  a c i d - a c e t i c anhydride  (1 : 20) c o l o r  reagent.  17  The  intensity  of the color  p r o d u c e d was d e t e r m i n e d  i n a model  B Beckman s p e c t r o p h o t o m e t e r a t 625 m i l l i m i c r o n s .  d) C h r o m i c The Samples  Oxide  p r o c e d u r e o f S c h u r c h e t a l . ( 1 9 5 0 ) was f o l l o w e d .  of feces  ( a n d f e e d ) were a s h e d The c o l o r  peroxide.  tracts  o f t h e p r o d u c t o b t a i n e d was d e t e r m i n e d i n a m o d e l B  Plasma  sixth,  a t 550 m i l l i m i c r o n s .  cholesterol  tenth  (four  cholesterol)  days  vein  determined  The  withdrawal  ( s e v e n days  after  of heparinized  was o b t a i n e d f r o m two m l . s a m p l e s  the wing  Zlatkis  after  into  heparinized  tubes.  of dietary  withdrawal of  of t h e experiment.  were made on 0.1 m l . s a m p l e s plasma  d e t e r m i n a t i o n s were made on t h e  days  c h o l e s t e r o l ) and t h i r t e e n t h dietary  water ex-  Cholesterol  Plasma third,  of d i s t i l l e d  with  sodium  Beckman s p e c t r o p h o t o m e t e r  intensity  and then f u s e d  Determinations plasma.  The  o f b l o o d drawn  Total  from  cholesterol  was  spectrophotometrically using the procedure of  e t a l . (1953) as m o d i f i e d by R o s e n t h a l e t a l . (1957).  c o l o r r e a g e n t u s e d was made up a s f o l l o w s .  Iron  Stock S o l u t i o n  -  2.25 grams F e C l ^ dissolved phosphoric  Color  Reagent  -  i n 100 m i s . 8 7 % acid.  8.0 m i s . i r o n diluted  6 RY,0  stock  solution  t o 100 m i s . w i t h  centrated  sulfuric  acid.  con-  18 The  reaction  3 mis. g l a c i a l color 560  acetic  and  individual  and 2 m i s . c o l o r  Table  increases  cockerels  The  spectrophotometrically  i n plasma c h o l e s t e r o l  r e s u l t i n g from  I supplemented w i t h  Tables  I I and I I I .  The d a t a  at  l e v e l s brought  dietary  cholesterol  feeding  are also  cholesterolemic  response  (strain  i s graphically  from  cockerels  hydro-  hypercholesteroboth of which  The a d d i t i o n a l  changes i n of the  s i x days o f c h o l e s t e r o l I I and I I I .  of the various  groups  levels) t o these  shown i n F i g u r e s  mean p l a s m a c h o l e s t e r o l of s t r a i n A at three  a pre-treatment  respectively from  a n d 10%  diet  The h y p e r -  (as i n d i c a t e d dietary  1 (strain A), 2  B ) , 3 and 4 ( s t r a i n s A and B ) .  The fed  show t h a t  i n Tables  t h e mean p l a s m a c h o l e s t e r o l  changes  the basal  about by w i t h d r a w a l  following  presented  levels of  and s i x days a r e p r e s e n t e d  i n a l l b u t two c o c k e r e l s ,  b e l o n g e d t o l o t 1 o f s t r a i n B. cholesterol  feeding  1% c h o l e s t e r o l  o i l f o r three  l e m i a was i n d u c e d  level  whereas t h a t  of cholesterol-  and s i x days  of cockerels levels  significant correlation  post-treatment  level  increased  o f 161 mg.% t o 299 a n d 486 mg.%  159 mg.% t o c o r r e s p o n d i n g  A and  reagent.  Discussion  genated vegetable  by  o f 0.1 m i s . p l a s m a ,  millimicrons.  The  in  acid  consisted  i n t e n s i t y was d e t e r m i n e d  Results  of  mixture  cholesterol  from s t r a i n B changed o f 278 a n d 399  mg.%.  (p 0.01) b e t w e e n p r e -  l e v e l s was d e m o n s t r a t e d f o r  19 i n d i v i d u a l birds but when s t r a i n d i s t i n c t i o n s were made these l e v e l s were s i g n i f i c a n t l y r e l a t e d only for s t r a i n B birds.  The regression  of post- on pretreatment l e v e l s i s  shown i n Figure 5. It was observed that birds previously thought to be s i m i l a r on the basis of t h e i r pretreatment l e v e l s were not always so when fed the c h o l e s t e r o l - r i c h d i e t .  The v a r i a t i o n  i n response of birds within a l o t appeared to be more marked i n birds from s t r a i n A compared to s t r a i n B.  This probably  accounted f o r the absence of a s i g n i f i c a n t c o r r e l a t i o n hetween pre- and post treatment l e v e l s of birds of s t r a i n A . The extractable  quantities of l i p i d s ( c h o l e s t e r o l , etherl i p i d s and saponifiable l i p i d s ) excreted by i n -  d i v i d u a l l o t s following s i x days of c h o l e s t e r o l feeding are given i n Table V.  These r e s u l t s when analysed (Table  VIII)  showed that the mean peak c h o l e s t e r o l l e v e l and cholesterol excreted were s i g n i f i c a n t l y c o r r e l a t e d (p 0.05).  A signifi-  cant c o r r e l a t i o n between the f e c a l c h o l e s t e r o l and the f e c a l saponifiable l i p i d s (p 0.01) and the f e c a l c h o l e s t e r o l and the percentage saponifiable l i p i d s i n the f e c a l ether extract (p 0.01) was also demonstrated.  The regression of the peak  cholesterol l e v e l on the f e c a l c h o l e s t e r o l , the f e c a l cholest e r o l on the f e c a l saponifiable l i p i d s and on the percent saponifiable l i p i d s i n the f e c a l ether extract are shown i n Figures 6, 7, and 8.  20 These r e s u l t s i n d i c a t e d relationship i n d u c e d and  between the the  a three-way  degree of  amount o f  dietary  inter-  hypercholesterolemia f a t and  cholesterol  absorbed.  The lipids  and  mean q u a n t i t i e s  ether  of  extractable  e a c h s t r a i n were as  lipids  e x c r e t e d by  birds  Cholesterol  v  10  Kilograms  Saponifiable Lipids  Ether Extractable Lipids  48.8  187.9  259.2  B  58.7  322.2  404.4  d a t a show t h a t birds  of  cholesterol  greater  strain  A.  the  March  a strain  bility  the  That may well  view p r e v i o u s l y  s t r a i n s to  the  have been a illustrated  treatment ease of that  would p r o b a b l y  difference  probably responsible of  the  only  f o r the  dietary  determinant by  birds  of  retained  explain  the  higher  difference  refractory  to  of  in lots  B.  and  absorpsuscepti-  hypercholesterolemia.  the  dietary  peak c h o l e s t e r o l  This  strain  in  The  Biely  in cholesterol  induced  the  present  levels.  belonging to birds  were  e x p r e s s e d by  degree of u t i l i z a t i o n  cholesterol  birds  lipids  were p r o m o t e d i n t h e s e b i r d s .  data substantiate  t i o n was  amounts o f  This  l e v e l s that  (1960) t h a t  of  Diet  A  The by  saponifiable  follows:  Grams L i p i d s p e r  Strain  cholesterol,  with  the  is particularly Tables  II  and  fat  level lowest  was pre-  true  in  III  show  cholesterol-feeding,  as  the  21  indicated on  by t h e s t a b i l i t y  t e s t were p r e s e n t  creted  f a r greater  of t h e i r cholesterol  i n l o t B 1.  The b i r d s  amounts o f c h o l e s t e r o l  e x p e c t e d on t h e b a s i s  levels  while  w i t h i n l o t T B l ex-  and f a t t h a n  was  o f amounts e x c r e t e d by t h e r e m a i n i n g  lots.  The absorbed fat  also of  of  i n d i v i d u a l v a r i a t i o n may  has been p r e v i o u s l y  changes  reported  be  for rats  i n t h e mean p l a s m a c h o l e s t e r o l  levels  l o t s which o c c u r r e d upon w i t h d r a w i n g t h e  cholesterol  f o r four  circulating cholesterol  a n d s e v e n d a y s a n d t h e amount  eliminated  at these, times a r e  I I , I I I and IV and summarized  A  328.6  E x c e s s (mg.%) E l i m i n a t e d at 4 days amt % 227 72  B  241.0  202  Excess Cholest e r o l present at 6 days  d a t a show t h a t  culating  dietary  e t a l . , 1958).  shown i n T a b l e s  The  that  of  absorption.  Marked i n d i v i d u a l v a r i a t i o n i n t h e u t i l i z a t i o n  the various  Strain  (reviewed) i n -  f a t s enhance c h o l e s t e r o l  indicates  lipids  The  dietary  by t h e amount o f  m a r k e d v a r i a t i o n i n t h e amount  marked.  dietary  extent  of cholesterol  s u p p o r t e d by e v i d e n c e  dietary  utilized  (Squibb  of  i s well  that  The lipids  t h e amount  i s determined t o a great  absorbed  dicating  assumption that  birds  cholesterol  from  per unit  86  below.  E x c e s s (mg.%) E l i m i n a t e d at 7 days amt % 314 97 239  strain A eliminated time than b i r d s  from  100  more  cir-  Strain  B  22  but  i n r e l a t i o n t o t h e amount  eliminated  lesser  elimination (i.e.,  amounts.  of significance.  peak c h o l e s t e r o l the absolute  nated per unit  cholesterolemia  levels.  This  The d a t a o f T a b l e  by-intravenous  with that  means.  elimi-  t o t h e amount o f previously  f e d a high-cholesterol a return Since  i n the present  hypercholesterolemia  hypercholesterolemic dietary  IV suggest  hyper-  i n j e c t i o n of hyperlipemic  however r e p o r t e d  noted  d i d not  difference  e t a l . ( 1 9 4 8 ) who r e n d e r e d b i r d s  i n twelve t o twenty-four hours.  variance  therefore  f o rthe s t r a i n  r e l a t i o n s h i p has been  and a s s o c i a t e s  assumed t h a t  by  difference  responsible  plasma o b t a i n e d from b i r d s  at  This  at the  amount o f c i r c u l a t i n g c h o l e s t e r o l  o b s e r v e d by H o r l i c k  levels  i n rate of  was a t i t s maximum  t i m e was d i r e c t l y r e l a t e d  excess present.  Horlick  When t h e d i f f e r e n c e  o f excess c h o l e s t e r o l  a p p e a r t o be g r e a t l y  that  they  a t 4 d a y s ) i t was f o u n d t o be n o n s i g n i f i c a n t  5% l e v e l  in  o f excess present  diet. t o normal  t h i s time i s  study  i t has been  i n d u c e d by i n j e c t i o n o f  p l a s m a does n o t s i m u l a t e  that  produced  23  EXPERIMENT  II  Introduction  In post was  experiment  treatment  cholesterol  demonstrated.  make a p r e l i m i n a r y pretreatment terol-fed  fed of  A  I t was  cockerels  investigation  to  (the  I I was  the  of  1 and  levels  of  the  2%  birds  f a t and  considered the  diet  also  for  to  used  and  interest  relationship  be  not  between  previously from  (10% on  fat  the  The  effects and  plasma  to  choles-  selected  months.  observe the  cholesterol)  cockerels  hypercholesterolemia)  several  diet  of  Birds  were t o  to  and  s u s c e p t i b i l i t y of  atherosclerosis.  experimental  levels  dietary  and  more s u s c e p t i b l e  experiment of  level  between p r e -  cholesterol—fed  into  cholesterol-feeding  a cholesterol-rich  feeding  l e v e l s of therefore  cholesterol  subjected to strain  I a relationship  and  purpose  of. l o n g - t e r m  alternating cholesterol  upon t h e i r u t i l i z a t i o n  of  the  cholesterol.  Experimental  Six 175  -  195  dividual water. that level  cockerels  mg.%  were s e l e c t e d  cages w i t h f r e e The  experimental  used i n experiment of  with cholesterol  cholesterol.  from  strain A  access to diet  was  I except The  levels ranging  the  and  experimental  similar  of  in in-  diet  i n composition  for periodic  duration  placed  from  feeding  changes i n on  each  and to the  24 r e g i m e n was  as  follows:  Cholesterol  Content  Duration  1%  7  2%  13  "  ( "•  390  "  ( "  l°/o F e c e s were c o l l e c t e d f r o m previously  cholesterol  l e v e l s were a l s o  Results  and  feeding  l e v e l s of  T a b l e VI  An days.  in  these  and  of  21  d e t e r m i n e d by  20)  -  410)  stored  and  I.  Plasma  methods u s e d  increase  the  2%  in  during  mg.%.  It  crease  in thyroid  this latter  i s assumed t h a t  more p r e v a l e n t  on  i n the  (Cruikshank,  crease  in thyroid  o i l and  f o r 410  I  alter-  days are  shown  a general  to  depression  mean  cholesterol  period  1171  mg.%  brought  the  observed  i n T a b l e VI  this decline  activity  months  experiment  l e v e l s was  t h e r e was  indicated  assumption r e s t s  of  which  9.  in cholesterol  As  diet  cholesterol  Beyond t h i s p e r i o d levels.  basal  levels  hydrogenated vegetable  Figure  fell  is  10% 1 and  levels  This  8 -  i n experiment  changes i n plasma c h o l e s t e r o l  supplemented w i t h  88  indicated  1-7)  Discussion  accompanied the  in  (days  I.  The  nating  Feeding-  individual birds,  a n a l y s e d as  experiment  days  of  on  findings  domestic  fowl  1930;  Winchester,  activity  produces  a  from  resulted by  to  440  f r o m an  in-  winter  that  hyperthyroidism  during 1940)  conditions.  the  and  winter  that  an  in-  hypocholesterolemic  25 effect  (Byers et  A  a l . , 1958).  s i g n i f i c a n t c o r r e l a t i o n between p r e t r e a t m e n t  peak c h o l e s t e r o l but  the  l e v e l s at  that  (r  = +0.811, p 0 . 0 5 ) .  time  part  birds  cholesterol  fed  variability  of  the  i n the  at  birds  variability for  various  times.  significant correlation  terol  l e v e l s and  during  the  the  period  cholesterol Consequently  the  between c h o l e s t e r o l and as  saponifiable i n experiment  I.  of  170  the  feces  could a  i n question  also 357 ship and  i n existence and  363  at  between c h o l e s t e r o l saponifiable  lipids  no  regarding period.  (134  be  demonstrated  the  of —  ether  170  extract  days) i t i s  interrelationship the  i n d i c a t i o n of  feces.  cholesterol  not  amount  During  l e v e l s and i n the  feces  significant correlation  t h r e e way  t h i s time.  d a y s t h e r e was  Data  amounts o f  S i n c e t h e r e was  the  choles-  a three-way i n t e r r e l a t i o n s h i p  i n the  assume t h a t  by  revealed  i n the  days.  excreted  reasonable to  data  collected for this  l e v e l s and  period  excreted  cholesterol  extract  l e v e l s and  the  lipids  these  between p l a s m a c h o l e s t e r o l during  levels.  between plasma  were n o t  lipids  by  (p 0.05)  and  con-  accounted f o r  t h e i r plasma  ether  a  of  of  b e t w e e n 134  existence  that  Analysis  amount o f  excretion  of  levels  levels  cholesterol  shows q u a n t i t i e s  shown  significance  in cholesterol  t w e n t y d a y s was  i n r e l a t i o n to  be  f o r these  These r e s u l t s i n d i c a t e d  pretreatment  Table VII individual  not  ( r = +0.785) c l o s e l y a p p r o a c h e d  siderable  a  days c o u l d  correlation coefficient calculated  at  levels  twenty  and  period  was  between  an i n t e r r e l a t i o n -  amounts o f  cholesterol  26 The lipids  following table  shows t h a t  a b s o r b e d by i n d i v i d u a l c o c k e r e l s  high-cholesterol  Bird Number  diet  was  increased  t h e amount o f on t h e h i g h - f a t ,  a s f e e d i n g was p r o l o n g e d .  Grams E t h e r - E x t r a c t a b l e L i p i d s E x c r e t e d p e r 10 k g . D i e t 170 d a y s 357 d a y s  1  163.9  106.5  2  197.3  108.8  3  168.1  101.4  4  221.6  110.1  200.1 Mean  190.2  106.7  It  c a n be s e e n a l s o t h a t  to  a reduction  lipids  diet  a t 357 d a y s  explained  i t r a n g e d f r o m 101 —  d e c r e a s e i n amount  lipids  110 grams p e r 10  shown t h a t  absorb l i p i d s  cholesterol-rich  was  They  excreted  kilo-  can best  e t a l . (1961).  the capacity  increased  diet.  o f the mucosal  increased  of l i p i d s  by t h e f i n d i n g o f Murthy  workers have  These  by c o n t i n u o u s f e e d i n g  showed t h a t  with respect  This  to the saturated  of a  the e s t e r i f y i n g  increase fatty  containing  i n activity  acids  acti-  considerably  f o r c h o l e s t e r o l when r a t s were f e d a d i e t weeks.  be  of the i n t e s t i n e s of r a t s  enzymes o f t h e i n t e s t i n e s was  1% c h o l e s t e r o l f o r f i v e greater  of  diet.  The  vity  a t 170 d a y s t h e amount  r a n g e d f r o m 164 - 221 grams p e r 10 k i l o g r a m s o f  grams o f  to  of the diet l e d  i n i n d i v i d u a l v a r i a t i o n i n t h e amount o f  absorbed.Whereas  excreted  extended f e e d i n g  as  was  compared  27 to the u n s a t u r a t e d  The  ones.  survival  shown i n T a b l e V I . revealed every  that  test  of i n d i v i d u a l  Postmortem  The  cockerels i s  examination  c o r o n a r y o c c l u s i o n was  instance.  t a i n e d heavy  time  the  coronary vessels  of nonsurvivors  cause  of death  of these b i r d s  d e p o s i t s o f s u b s t a n c e s w h i c h gave a  for cholesterol,  in  con-  positive  w i t h the method o f R o s e n t h a l  et a l .  (1957) p r e v i o u s l y d e s c r i b e d .  With stances  small.  lowest  The  These  cholesterol  which f a i l e d  to  p o r t e d by F i s h e r  level  survive.  blood vessels  i s also  of s u r v i v o r s  test.  throughout  maintained  throughout  the  These r e s u l t s  diet  aorta.  the  a lower  experiment  mean  than  those  agree w i t h those  f e d to Leghorn  pullets  refor  c o n t a i n i n g 10% animal f a t .  of substances  deposited i n the  i n agreement w i t h t h a t  g r e a t e s t amounts were p r e s e n t i n t h e  ment o f t h e  was  levels  e t a l . ( 1 9 6 0 ) who  distribution  sub-  associated  three years a cholesterol—free  The  of  s m a l l q u a n t i t i e s were  b i r d s which s u r v i v e d  plasma c h o l e s t e r o l  Fisher:  t h e amount  d e p o s i t e d i n the b l o o d v e s s e l s  comparatively w i t h the  e x c e p t i o n ( b i r d #4)  one  observed abdominal  by seg-  TABLE I COMPOSITION OF BASAL DIET Ingredients  Per 100 l b s d i e t lbs  -  Ground -wheat Oats Corn Soybean meal D i c a l c i u m phosphate D i s t i l l e r s dried solubles Ground l i m e s t o n e Iodized s a l t ^Feeding o i l V i t a m i n B^^ Vitamin B Manganese s u l f a t e  gms  60.00. 10.00 5.00 16.50 1.75 .2.00 3.90 0.35 0.50 0.54 0.10 5.68  2  100.00  6.32  * 3000 U . S . P . u n i t s v i t a m i n A p e r gram and 300 I n t e r n a t i o n a l c h i c k u n i t s v i t a m i n D3 p e r gram.  TABLE I I PLASMA CHOLESTEROL LEVELS (mg %) OF COCKERELS OF STRAIN A IN EXPERIMENT 1 Lot  1  Mean  -2  P r e t r e a t - Average P.T. ment l e v e l level (i)  (ii)  123 130 150  129 129 131 136  136  131  14-1  140 141 140 145 146 159 145 152  126 135 131  143 134 146  1 41 146 152  Days  Cholesterol fed  Days  Cholesterol •withdrawn  3  6  4  7  324 246 225 253  538  204 4 00 390  172 130  262  383  224 210 I84  135 132 150  —  -  484  140 139 ^  m  154  678 398  4 10 152  — 210  143  Mean  152  146  149  319  538  281.  177  3  158 159 158 158  152 153  295 344 300  350  304  294  .200 152 194 186  158  1 54 159  155 156 156 159  Mean  158  155  157  311  359  183  162  4  168 160 179 155  160 165 166 170  190 194 210  157 138 180 220  166 185 165 173  Mean  5  Mean  164  488 30.2  470  145 134 209  163 173 163  .297 282 205 494  165  166  320  496  254  174  183 170 174 174  448  171 187 300  800 578  44O 310  1.7.2  180 175 175 175  205 169 223 272  174  176  175  277  462 250 800  420  6 66 782  400  707  398  440  217 (Continued)  30  Lot  P r e t r e a t - " Average P.T. ment l e v e l level  Days  Cholesterol fed  Days  Cholesterol withdrawn  3  6  4  7  (i)  (ii)  190 182 198 181  183 182 189 189  187 182 194 185  310 370 331 210  466 398 592 374  292 316 280  199 164 170 205  188  186  187  305  458  278  185  Grand Mean 162  160  161  299  490  263  176  6  Mean  224  3 1 TABLE I I I PLASMA CHOLESTEROL LEVELS (mg %) OF COCKERELS OF STRAIN B IN EXPERIMENT 1 Lot  1  Pretreatment l e v e l  (1)  (ii)  102  122  110 114 129  125 131  133  Average P.T. level  112 118 123  131 121  Days  Cholesterol fed  Days  Cholesterol withdrawn  3 124  6  4 100 94 110  7  131 240  140 114 124 394  198  150  193  126  117  368 282  432  I64  143 142 143  105  125  101 108  134  114  128  160 150 158 169  14? 151 158 157  158 163  238  350 494 356  Mean  159  154  157  319  408  168  3  158  158 157 161 157  378 187  476 476  162  163 155  157 159 158 159  156  346 140  174 148  Mean  158  158  158  230  360  172  4  155 168  160 163 165 168  158 166 158 168  172 260 246  441  454 372 476 520  198 152 184 198  Mean  161  I64  163  280  456  183  164  5  156 162 174 160  172 174 177 182  I64 168 176 171  255  218 250  240  429  384 610 354 560  278  169 173 195 183  163  176  170  351  477  247  180  Mean  2  155  168  151  Mean  154  151  388  200  388 330  172 160 174  204  154  146 154  160 168  134 154 169  148 179 158  (Continued)  32  Lot  Pretreatment l e v e l  Average P.T. level  Days  Cholesterol fed  Days  Cholesterol •withdrawn  3  6  4  7  (i)  (ii)  180 172 170 1.80  185 185 186 189  183 179 178 185  198 300 446  400  320 350 820 520  190 186 388 390  200 190 185 227  Mean  176  186  181  336  503  289  201  Grand Mean  155  161  . 158  278  399  197  160  6  33 TABLE IV QUANTITIES OF EXCESS CIRCULATING CHOLESTEROL  (mg %)  ELIMINATED FOLLOWING CHOLESTEROL WITHDRAWAL FROM THE DIET OF COCKERELS IN EXPERIMENT 1 Strain A Lot  T o t a l Excess Present  Amount E l i m i n a t e d A t  4 days  7 days  1 2 3 4 5 6  249.3 389.0 202.0 329.7 531.2 ,270.5  199.0 257.0 175.5 242.0 309.0 179.5  (79.8) (66.1) (86.9) (73.4) (58.2) (66.4)  243.7 361.5 197.0 321.7 489.2 273.0  (97.8) (92.9) (97.5) (97.6) (92.1) (101.0)  Mean  328.6  227.0  (71.8)  314.3  (96.5)  Strain B 1 72.0 2 251.5 3 201.3 4 293.0 5 307.2 6 321.2  67.5 240.5 187.5 272.5 230.5 214.0  (93.8) (95.6) (93.1) (93.0) (75.0) (66.6)  76.0 262.5 205.5 292.0 297.0 302.0  (105.6) (104-4) (102.1) (99.7) (96.7 > ( 94.0)  202.1  (86.2)  239.2  (100.4)  241.0  Mean (  ) % o f excess  eliminated  34  TABLE V LIPIDS (gm/10 k g d i e t ) IN FECES OF BIRDS OF STRAINS A AND B, FED HIGH FAT, HIGH CHOLESTEROL DIET IN EXPERIMENT 1 Strain A Lot  Cholesterol  Ether extractable lipids  Saponifiable lipids  231.5 231.2 201.5 223.3 122.2 118.0  75.6 75.1 73.4 77.0 65.3 62.4  %  Saponifiable lipids i n ether e x t r a c t  1 2 3 4 5 6  49.6  47.3  306.4 307.8 274.5 290.2 187.3 189.2  Mean  48.8  259.2  187.9  71.5  779.1 337.7 274.5  86.6 75.0 73.4 72.9 71.7 80.5 77.3  51.9 52.1 49.5  42.6  Strain B 1 2 3 4 5 6 Mean  42.5 59.8  217.0 504.5  674.8 253.2 201.5 175.5 155.5 406.0  58.7  404.4  322.1  77.9 61.0 52.1 57.3  240.8  TABLE VI c<-\  PLASMA CHOLESTEROL LEVELS (mg %) OF COCKERELS OF STRAIN A, FED HIGH FATy HIGH CHOLESTEROL DIET IN EXPERIMENT Bird no.  1  Pretreatment cholesterol level 7* 189 175 180  2  3 Mean (S)  4 5 6  4-60  320 -  195  195  676 580 640  14**  ,20**  35*-  88*  10 76 734 556  1372  10 82  1045 936  806  1111 795 678  1117.6  864.O  861.3 690.0 518.0 366.6 500.0 503.3  410  2059 1329 2371  1898 1020 14-69  1750  1270 900 1300  704  Mean (NS)  194.6 632.0 11.56.6 1919.6 14.62.3  Mean (S+NS)  188.0 535.2  *  Days o f survival  Days d i e t f e d  181.3 390.0 788.6  194  2  134*  275*  345*  363*  370*  808  858 343 353  550 250 300  670  670  330 500  340 500  757 505  1616  1520  1414  1635  1515  20 7  490  1050  1100  384 70 250  207  490  1050  1100  235  972.6 1518.6 1163.2 1170.8 1020.0 44O.3 397.5 637.5 652.5  1% d i e t a r y c h o l e s t e r o l  ** 2% d i e t a r y c h o l e s t e r o l  410 410 410  S survivors  NS n o n s u r v i v o r s  322  TABLE V I I PLASMA CHOLESTEROL LEVELS (mg %) IN RELATION TO LIPIDS EXCRETED (gm/lO kg d i e t ) BY COCKERELS OF STRAIN A IN EXPERIMENT 2  Bird No.  1 2 3 4  5  Mean  Pretreatment cholesterol levels  Plasma c h o l e s t e r o l l e v e l at 134- days  189 175 180 195 195  808 757 505 1616 141-4-  186.8  1020.0  363 days 670 330 500 1050 —  637.5  Ether extractable l i p i d s at 170 days  357 days  163.9 197.3 168.1 221.6 .200.1  106.5 108.8 101.4. 110.1  190.2  106.7  —  Fecal cholesterol a t 357 days  34.9 49.2 55.6 48.2  -  47.0  TABLE V I I I SUMMARY OF CALCULATIONS OF REGRESSION AND CORRELATION COEFFICIENTS Experiment Strain  Y  X  A + B  Pretreatment cholesterol level  n  f e c a l cholesterol f e c a l saponifiable lipids  ti  tt  Peak cholesterol level H  fecal cholesterol tt  % fecal saponifiable lipids  Experiment  *  1 X*  Y  b  r  df  p  44  0.01  -9.11  +0.473 -0.710  10  0.05  53.7  0.05  +0.856  10  0.01  74.4  53.7  1.19  +0.782  10  0.01  160.3  444* 5  4.30  53.7  444* 5  255.0  2  A  Pretreatment cholesterol level  Peak cholesterol l e v e l a t 20 days  188.0  1519.0  52.40  +0.785  4  0.05*  B  Ether extract a t 170 days  Plasma cholesterol a t 134- days  190.2  1020.0  16.29  +0.830  4  0.05  Approaching  significance  38 F i g . 1.  Mean plasma c h o l e s t e r o l l e v e l s o f l o t s o f c o c k e r e l s o f s t r a i n A, i n experiment 1.  39 Fig.  800-j  70CH  2.  Mean plasma cholesterol levels of lots of cockerels of s t r a i n B, i n experiment 1.  40 Fig. 3.  Mean plasma cholesterol levels of cockerels of strains A and B, i n experiment 1.  50O "i  A1-6  AOO-\  SooJ  zoo  150 A  I BAYS Of  a  i  r  3  4  CHOLESTEROL  5 FEEDING-  DAYS  OP  CHOLESTEROL  WITHDRAWAL  41 F i g . 4.  Degree of hypercholesterolemia induced i n lots of cockerels of strains A and B fed a high-fat, high-cholesterol diet i n experiment 1.  QOO-i  )*:»j\ CRlA$>fi XN  IH CHOLESjEWot LEVEL  Pf?eT«6ATME/VT  7oo-  50o-  -J Ul  > LU  440-  o ui H  (0 ui -J  O  u < —i  Q.  loo-  100-  A4  &4  n  Fig.  5.  Regression of peak cholesterol on pretreatment cholesterol l e v e l s of i n d i v i d u a l cockerels of s t r a i n s A and B i n experiment 1.  43  F i g . 6.  8oo  -i  Regression of mean peak cholesterol levels on the amount of cholesterol i n feces of cockerels fed a high-fat, high-cholesterol diet i n experiment 1.  Fig. 7.  Regression of cholesterol on the saponifiable l i p i d s i n the feces of cockerels fed a high-fat, high-cholesterol diet i n experiment 1.  GRAMS PER  SAPONIFIABLE IO  KILO&KAMS  LIPIDS OF  DIET  IN  FECES  45 Fig. 8.  6  0  Regression of the amount of cholesterol excreted on the percent of sponifiable l i p i d s i n the ether extract of feces of cockerels fed a high-fat, high-cholesterol diet i n experiment 1. Q  I  r  66  70  PERCENT  9APONIF/ABU  1—  70 LIPIDS  T  1  "i  SO  85  90  IN ETHER  EXTRACT  OF  FECES  &  F i g . 9.  Plasma cholesterol l e v e l s of cockerels of s t r a i n A, fed a high-fat, high-cholesterol d i e t i n experiment I I .  DAYS  ON  TEST  '47  SUMMARY AND  CONCLUSIONS  Two e x p e r i m e n t s were c o n d u c t e d w i t h a d u l t Leghorn cockerels levels  promoted  of u t i l i z a t i o n  o f t h e s e two d i e t a r y  of the diet  components. (1)  a n d (2) t h e e f f e c t  on t h e d e v e l o p m e n t  In  shown t o d i f f e r cholesterolemia. a wide range  were s e l e c t e d  small  contained birds  terol  f r o m two s t r a i n s  were s e l e c t e d  l o t s on t h e b a s i s with small  cockerels.  previously  induced hyper-  i n order t o ensure l e v e l s a n d were  of these l e v e l s .  differences  i n their  Each  choles-  levels.  Cholesterol  a n d h y d r o g e n a t e d v e g e t a b l e o i l were  fed  a t l e v e l s o f 1% a n d 10%  the  end o f t h i s t i m e marked d i f f e r e n c e s  cholesterol two  i n adult  feed-  (experiment I ) the  i n pretreatment c h o l e s t e r o l  into  by e x t e n d e d  of t h i s type of  i n susceptibility to dietary The b i r d s  In addition  t h e manner i n  of atherosclerosis  the s h o r t - t e r m experiment  experimental birds  assigned  cholesterol  of a hydrogenated  t h e a b o v e r e l a t i o n s h i p may be a f f e c t e d  feeding  lot  10%  containing  experiments attempted t o determine  which  ing  by a d i e t  plasma  o i l a n d 1% c h o l e s t e r o l were r e l a t e d t o t h e d e g r e e  vegetable  the  t o determine whether  White  groups  respectively  f o r s i x days. i n t h e mean  At  plasma  l e v e l s w e r e o b s e r v e d among l o t s a n d b e t w e e n t h e o f s t r a i n s A a n d B.  48  When c h o l e s t e r o l plasma The  cholesterol  was w i t h d r a w n f r o m t h e d i e t  returned to the pretreatment l e v e l s .  l e v e l s observed a f t e r four  indicated  that  the differences  and seven days i n the degree  h y p e r c h o l e s t e r o l e m i a were n o t p r i m a r i l y i n the rate  of elimination  withdrawal of  dietary  due t o  differences  of excess c h o l e s t e r o l  from the  circulation.  The lots  amount  following  of cholesterol  e x c r e t e d by t h e  various  s i x days  feeding  of the high-fat,  diet varied  greatly  a n d was  f o u n d t o be  signi-  ficantly  c o r r e l a t e d w i t h (1) the plasma  cholesterol  levels  (p 0.05)  a n d ( 2 ) t h e amount  cholesterol  (p 0 . 0 1 ) . terol  These  absorption  Differences  In  may  i n consequence  experiment  i n contrast  evident.  differences  diet  be a t l e a s t  I I cockerels  were f e d a  f o r an extended p e r i o d I w h i c h was  choles-  absorption. hyper-  partly  high-fat,  of time.  This  a short-term t e s t .  t h e r e l a t i o n s h i p between hyper-  and d i e t a r y  lipid  utilization  was n o t c l e a r l y  I n b o t h e x p e r i m e n t s however t h e degree  hypercholesterolemia  in  to dietary  absorption.  t o experiment  excreted  in fat  in fat  In the second experiment cholesterolemia  lipids  a r i s e from d i f f e r e n c e s  to differences  high-cholesterol is  may  suggest that  i n s u s c e p t i b i l i t y of cockerels  cholesterolemia attributable  results  of saponifiable  high-  and p r e t r e a t m e n t c h o l e s t e r o l  f o u n d t o be s i g n i f i c a n t l y c o r r e l a t e d .  of induced l e v e l s were  I t was 410  observed t h a t b i r d s which  survived  days on a h i g h - f a t , h i g h - c h o l e s t e r o l d i e t ,  maintained  throughout the t e s t a lower mean plasma c h o l e s t e r o l l e v e l than n o n s u r v i v o r s . to n o n s u r v i v o r s  The  b l o o d v e s s e l s of s u r v i v o r s  showed l i t t l e  compared  evidence of a t h e r o s c l e r o s i s .  50  BIBLIOGRAPHY  A h r e n s , E . 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