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Lingual lipase exists in
rats and some other rodent species - not in humans
lipase from human breast milk (bile salt-stimulated lipase) is take up by
infants and activated in the small intestine
majority of lipid digestion starts in the stomach:
-contraactions and churning results in fat/lipid droplets
-gastric lipase
-chyme
contractions and churning results in fat/lipid droplets causes
physical reduction in size
gatric lipase
responsible for up to 25% of TAG hydrolysis, pH optimum ~4.0
chyme (composed of fat/lipid droplets) is allowed into the
small intesting via the pyloric sphincter
microorganisms synthesize lipases and
secrete them into the rumen
Extracellular lipases completely hydrolyze
TAGs and other dietary lipids
unsaturated fatty acids are extensively hydrogenated by other species of microorganisms into
stearic acid, cis-9, trans-11 CLA, trans-10, cis-12 CLA, and trans—vaccenic acid (18:1 trans-11)
only small amounts of TAGs and phospholipids enter the small intestine in ruminants, free fatty acids primarily
enter the small intestine
lipid digestion in the small intestine occurs
in the duodenum
challanges of lipid digestion in the small intestine are
lipids are hydrophobic, TAGs can not be absorbed due to their size
solutions to the challanges faced by lipid digestionin the small intestine:
bile salts and pancreatic enzymes (lipase) are mixed with TAGs (TAGs become emulsified and become mixed micelles)
secretion of cholecystokinin from the intestinal mucosal cells (stimulated by the presence of fat in SI) ccauses:
gall bladder contractions, secretion of pancreatic digestive enzymes (~1200mL/day)
gall bladder contractions
holds ~40 to 70 mL of bile salts and bile acids and releases ~700 mL/day (extensive recycling)
secretion of pancreatic digestive enzymes (~1200 mL/day)
pancreatic lipase (very small amounts in ruminants), colipase
colipase
activates lipase in the presence of bile salts
bile acid functions:
emulsify fats, activation of pancreatic lipase (at low concentration), formation of mixed micelles
bile salt emulsification
bile acids and glycine or taurine conjugates serve as detergents, cause the formation of TAG particles of 1 microm or less, other emulsifiers
emulsifiers involved in lipid digestion other than bile salt
phospholipids (esp. lysolecithin), 2-monoacylglycerols
hydrolysis of dietary fat by pancreatic lipase wors at lipid-water interface and is activated by
bile salts at low concentrations
hydrolysis of dietary fat by pancreatic lipase is inhibited by
bile salts at high concentration, hydrolyzes TAG to fatty acids and 2-monoacylglyerol (2-MAG)
pancreatic colipase is secreted by
the pancreas with lipase
pancreatic colipase binds to bile salt emulsion droplets and reduces
inhibitory action of bile salts on pancreatic lipase
absorption in the small intestin involves the formation of
mixed micelles
mixed micells are 4-6nm in diameter and are formed when
bile salts and fatty acids reach a critical micellar concentration spontaneously form
mixed micelles incorporate 2-MAGs, lysolecithin, free cholesterol, and long-chain fatty acids essentially
detergents in themselves
absorption in the the enterocytes (intestinal mucosal cells) by 2-MAG, lyso-phospholipids, fatty acis, and cholesterol dissociate from
micelles at the surface of mucosal cells and produce locally high concentrations of 2-MAG, lyso-phospholipids and fatty acids
the production from absorption into the enterocytes are then absorbed by the entrocytes of the
duodenum and proximal jejunum
synthesis of TAG and phospholipids in encytes produce
triglycerols
75% of TAGs are synthesized via the 2-MAG pathway located on the smooth endoplasmic reticulum, 25% of TAGs are synthesized via the
standard TAG biosynthetic pathway located on the rough endoplasmic reticulum
triaclyglycerols include
phopholipids (especially lecithin) and cholesterol esters
synthesis of TAG and phopholipids in enterocytes cause the formation of
chylomicrons
chylomicrons synthesis starts when
lipid droplets form within the ER and goligi apparatus, these droplets contain TAG, phospholipids, cholesterol, cholesterol ester and apolipoprotein (with carbohydrates) complexes
apolipoproteins and phospholipids are formed in the rER and then TAG are formed in the
sER and transferred to apolipoprotein B to form a core for the chylomicron
after accumulating TAG and cholesterol ester, golgi vesciles form and CHO moietes are
added to the apolipoproteins
golgi vescles fuse with the cell membrane and are extruded into the lacteals, chylomicrons are then transported
via the lymphatics
composition of chylomicrons
70-90% TAG, 4-8% phospholids, 3% cholesterol, 4% cholesterol ester, 2% protein (apolipoprotein B)
apolipoprotein
protein that binds lipid for transport through circulatory and lymphatic systems
ruminants do not synthesize true chylomicrons, too little TAG because
they consume little dietary fat, very low density lipoprotein (similar to chylomicrons)
dietary cholesteral absorption in humans
~0.4-0.5 g/day
dietary cholesteral digestion in herbivores
virtually no cholesterol ingested
biliary
20-30 g bile salts enter the SI daily, cholesterol and cholesterol ester enter the sI via bile
a minor contributor to total cholesterol intake is
intestinal mucosa
a minor contributor to total cholesterol intake
ruminal microflora
mechanism of cholesterol absorption in the small intestine starts when cholesterol esters are
hydrolyzed by pancreatic cholesterol esterases
free cholesterol is incorporated into mixed micelles (bile acid presence is needed); free cholesterol is absorbed into
the intestional mucosal cells and re-esterfied to form cholesterol esters
sources of cholesterol loss from GI tract
bile salts, poorly absorbed, plant sterols
0.8 g/day bile salts are lost in feces while the rest is
taken up in the ileum for enterohepatic circulation
cholesterol is porly absorbed: ~0.4 g/day lost in feces and
only 30-40% dietary and biliary cholesterol is absorbed
200-300 mg/day of plant sterols are ingested and
absorbed in trace amounts
in large amounts, plant sterols
inhibit cholesterol absorption
0.2-0.4 g/day of cholesterol is lost as
sloughed skin
cholesterol ester is
cholesterol with long chain fatty acids linked to the hydroxyl group
cholesterol ester is much less polar when
compared to normal cholesterol
cholesterol ester preferred form of transport is
in the blood