Author(s): Rebecca W. Van Dyke, M.D., 2012

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Liver Physiology

Rebecca W. Van Dyke, MD

Winter 2012 Learning Objectives

• At the end of this presentation students should be able to: • 1. Describe the basic organization of the cell plate and its functional • consequences: • a. Blood supply • b. Configuration of • c. Configuration of other liver cells • d. Concentration gradients in sinusoidal blood. • 2. Describe the basic physiological processes the liver utilizes to accomplish function: • a. transport • b. metabolism • c. biotransformation • d. synthesis • e. secretion • 3. Be able to give examples of the consequences of liver damage on above processes. • 4. Be able to give examples of possible consequences of liver disease/injury on liver barrier function and hepatic regeneration. Industry Relationship Disclosures Industry Supported Research and Outside Relationships

• None

Cystic artery sole supply to Anatomy of Liver Acinus bile duct

hepatic artery portal blood flow portal triad bile

sinusoids

Michigan Histology Collection central vein Dual Blood Supply of Liver

liver Hepatic artery: 20%

Portal vein: 80% spleen

Liver has dual blood Regents of the University of Michigan supply: 80% portal vein 20% hepatic artery

What are the Functional Consequences of….?

• Increased vena caval pressure/hepatic vein obstruction?

• Decreased hepatic artery blood flow? and/or Decreased portal vein blood flow?

• Effects on the bile duct? Budd-Chiari Syndrome: Obstruction of the Hepatic Vein(s)

Normal Budd-Chiari

Hemorrhage in Dilated upstream pericentral area; sinusoids; hepatic vein atrophic/ischemic obscured hepatocytes Decreased inflow: ischemic infarction Bile Duct

• Sole blood supply to the bile duct is through the hepatic artery via the cystic artery

• Reduced blood flow through the hepatic artery causes ischemic injury to the extrahepatic bile ducts

• Ischemia or damage to the bile duct often leads to injury, fibrosis, stricture Liver Cell Anatomy:

Consider functional consequences

Image showing relationship between sinusoid, sinusoid lining cells, and removed. Scanning Electron micrograph of Liver Cells in Liver Sinusoid American Gastroenterological Association Functional Consequences of Fenestrated Sinusoidal ? Space of Disse

Other serum proteins

Albumin

Tissue macrophage

Filtration device , endotoxin

Releases inflammatory mediators that influence hepatocytes positively or negatively

Looking Down a Sinusoid at a Kupffer Cell Poised to Grab Passing Bacteria Kupffer cells (red) Ito Cell/Fat-storing Cell

Rare cell, located in sinusoids under endothelium Stores lipophilic materials such as Vitamin A Stimulated by chronic inflammation/alcohol converts to myofibroblast produces collagen and extracellular matrix Responsible for much of the excess fibrotic material in cirrhosis Ito Cell (Fat-storing Cell) on a Sinusoid (S) Hepatic Stellate (Ito) Cell (arrows)

The Liver: The Body’s Refinery

Defects in Glucose Metabolism in Liver Disease

Acute liver failure: Hypoglycemia (rapid neuronal death)

Chronic liver disease: Insulin resistance and diabetes (unknown mechanism) UREA

Fatty acid metabolism within hepatocytes albumin

hepatocyte FABP

free fatty acid pool

excess esterification to glucose, to mitochondria TAG amino acids for energy

Apo cholin B100 e VLDL cholesterol esters

Regents of the University of Michigan Greater Role of the Liver in Production and Metabolism of Lipoproteins and Lipids

LDL HDL Chol. Cholesterol Gut Liver

Chylomycrons CM remnants TG TG VLDL-TG BILE Cholesterol Bile acids Phospholipids

Fatty Liver with Inflammation Liver as Protein Synthetic Machine

Vena cava systemic outflow Liver synthesizes and secretes: Lipoproteins Albumin Clotting factors Anti-proteases (α1-anti-trypsin) Fibrinogen Complement factors Ceruloplasmin Transferrin and other binding proteins

Hepatic artery carries arterial blood with blood proteins

Portal vein carries venous blood from intestine, spleen and pancreas Protein Secretion Defects in Liver Disease

Example Clinical Consequence

Albumin Decreased plasma oncotic pressure/ edema Decreased binding of hydrophobic compounds

Clotting factors : Decreased factors II, VII, IX and X Increased bleeding

Fibrinogen Decreased fibrin formation in clotting

Defects in Protein Synthesis/release also cause liver disease: Alpha1-Anti-trypsin Deficiency

Image of pathophysiology of alpha-1- anti-trypsin deficiency removed PAS Stain Showing Retained Globs of Mutant Alpha1 Anti-trypsin Protein in Hepatocyte ER Lipoprotein release: another liver synthetic function

VLDL: a combination of fat and protein

The unique position and blood supply of the liver also affect liver physiology Anatomy of Liver Acinus bile duct

hepatic artery portal vein blood flow portal triad bile

sinusoids

Michigan Histology Collection central vein

Consequences of Changes in Hepatic Blood Flow and/or Oxygen Delivery? Peri-central vein (hepatic vein) clotted off with ischemic damage to hepatocytes

Peri-central ischemia

HV clot

Peri-portal normal tissue Not all liver cells are alike.

Substances found in higher concentrations in the portal vein • Albumin • CPS • FABP • HMG CoA

Substances found in higher concentrations in the hepatic vein • P450s • ADH • C7αH • Cysteine • GR • Gluatamate •α 2µG • GS • GLUT-1 •α -KG Periportal Necrosis from Allyl Formate Toxicity

HV

PV Pericentral Necrosis from Carbon Tetrachloride Toxicity

Normal cells

PV

HV Necrosis

Bile Formation Functions of Bile

• Transports material to the intestine for excretion – Drugs, toxins, xenobiotics – Cholesterol – Bilirubin – Copper • Transports bile acids to the intestine to aid in fat absorption Bile Acids

• Organic acid synthesized in liver from cholesterol • Conjugated to amino acids • Secreted in bile - essential for fat digestion/absorption • Reabsorbed in distal ileum and returned to liver via portal vein Bile Acid Cholesterol

OH Cholesterol: Flat (planar) hydrophobic compound compound hydrophobic (planar) Flat Cholesterol: Lose the double bond bond double the Lose

OH Metamorphosis to a bile acid acid bile a to Metamorphosis Shorten the side chain chain side the Shorten

OH Metamorphosis to a bile acid acid bile a to Metamorphosis

plane of the rings rings the of plane

and bend this below the the below this bend and Add a carboxylic acid group group acid carboxylic a Add

OH

COOH COOH Metamorphosis to a bile acid acid bile a to Metamorphosis

that is bent down down bent is that

Add a hydroxyl group group hydroxyl a Add OH OH

OH

COOH COOH Metamorphosis to a bile acid acid bile a to Metamorphosis OH OH

OH

COOH COOH

hydroxyl group group hydroxyl

OH OH

Add another another Add Metamorphosis to a bile acid acid bile a to Metamorphosis OH OH

OH

COOH COOH

OH OH

you now have a tri-hydroxy bile acid: cholic acid acid cholic acid: bile tri-hydroxy a have now you Metamorphosis to a bile acid acid bile a to Metamorphosis OH OH

OH

NH NH COOH COOH CO

OH OH

Taurocholate (taurine conjugated cholic acid) acid) cholic conjugated (taurine Taurocholate

conjugation of an amino acid to the side chain yields yields chain side the to acid amino an of conjugation One more change - - change more One Conjugated tri-OH Bile Acid

Hydrophobic side

OH- OH- OH-

COO- Hydrophilic side Biliary Lipids Bile acids from intestine To intestine for fat digestion Enterohepatic Circulation of Bile Acids: recycling is efficient Bile acids cycle between the liver and the small Bile acid synthesis intestine.

Total bile acid pool is about 3 grams. Liver About 90% of bile acids are reabsorbed in the terminal ileum.

However about 5-10% of bile acids are lost daily into the colon. Effect?

Small bowel Liver synthesizes about Colon 5-10% of the total bile acid pool each day. Enterohepatic Circulation of Bile Acids: Ilial loss

Bile acid Resection of 40 cm of synthesis the terminal ileum will result in what problem?

Bile acid loss into the cecum Liver will increase. What will this cause?

Liver upregulates bile acid synthesis and bile acid pool remains normal. Fat absorption remains the same.

Colon 40 cm resection of terminal ileum Enterohepatic Circulation of Bile Acids: Loss of most of the ileum

Bile acid synthesis Resection of >100 cm of the terminal ileum will result in what problem? Liver Initially, bile acid loss into the colon will be massive. What will be the initial effect of this loss of bile acids into the colon?

Colon >100 cm resection of terminal ileum Enterohepatic Circulation of Bile Acids: Lost of Ileum Resection of > 100 cm of the terminal ileum will Bile acid synthesis result in what problem over time?

Liver upregulates bile Liver acid synthesis but cannot keep up with loss rate. Bile acid pool is reduced Fat is malabsorbed.

As the bile acid pool falls, loss into the colon is less per day and secretory diarrhea due to bile acids converts to steatorrhea Colon (+ secretory diarrhea > 100 cm resection from fatty acids). of terminal ileum Enterohepatic Circulation of Bile Acid: Cholestyramine

Bile acid Cholestyramine: synthesis bile acid binding resin that removes bile acids from the enterohepatic circulation Liver Liver upregulates bile acid synthesis (using up what compound in the process?)

If liver cannot keep up, what happens?

Small bowel Less free bile acid in Colon the colon causes what? Liver takes up and excretes many other organic compounds: bilirubin is the classic and historic example

Hepatic Bilirubin Transport

SER UDP-glucuronide RBC + breakdown Unconj BR in RES Conj BR Unconj BR Bile Unconj Canaliculus Bilirubin Conj BR MRP-2: Multispecific organic anion transporter Conj Conjugated bilirubin BR Glutathione S-conjugates other organic anions ATP

Blood Hepatocyte Jaundice of the Neonate

• Newborn infants have poorly developed bilirubin conjugation enzymes and jaundice is common.

• Premature infants are even more affected

• Unconjugated bilirubin in the brain causes permanent damage (kernicterus)

• How to prevent brain damage in neonates?

Regents of the University of Michigan Phototherapy for Unconjugated Hyperbilirubinemia of the Neonate Martybugs, Wikimedia Commons. Hepatic Bilirubin Transport and Mechanisms of Hyperbilirubinemia

Gilbert's syndrome (mild) Crigler-Najjar syndrome (severe)

SER Hemolysis

Unconj BR Bile Unconj Canaliculus Bilirubin Conj BR Multispecific organic anion transporter

Conj Conjugated bilirubin BR Glutathione S-conjugates other organic anions ATP

Blood Hepatocyte

Dubin-Johnson syndrome Rotor's syndrome ?estrogen/cyclosporin Consequences of Liver Disease

for Bilirubin Handling?

Bilirubin: Jaundice

The first liver disease test CDC Liver Biotransformation/Excretion of Endogenous/Exogenous Compounds

• Bilirubin conjugation is an example

• Many other organic compounds undergo two- step biotransformation – Example: cholesterol to bile acids

• After biotransformation, metabolites excreted – Larger, lipophilic molecules excreted in bile – Smaller (<400 Da) transported to blood and excreted by kidneys

Step-wise Synthesis of Bile Acids from Cholesterol Steps are analogous to Phase I and Phase II steps of drug/xenobiotic metabolism Cholesterol

P450- mediated hydroxylations

Conjugation OH group of side chain to glycine or amino acid taurine Liver and Gut Barrier Functions

Insoluble,Liver helps to remove/eliminate: nonabsorbable compounds Xenobiotics Xenobiotics: metabolism, excretion Liver RES/filter Drugs Metabolism Bacteria Drugs (acid, : physical metabolism, excretion Biliary excretion barrier, Bacteria gut immune : Kupfer cells system, liver RES)

Intestine mucosal barrier Pancreas Liver’s Magic Trick: Regeneration

Image of liver regeneration process removed Prometheus Bound

P.P.Reubens

An early case of of hepatic regeneration

P. P. Rubens Functional Consequences of Losing a Large Amount of Liver Due to Resection/Necrosis?

• Hypoglycemia • Poor blood clotting • Cholestasis and jaundice • Increased blood ammonia - affects cognitive function • Decreased drug disposition • Abnormal lipid metabolism Summary

• Liver exhibits a wide range of functions • Liver diseases may cause malfunction of one or more normal function • Functions regulated separately so any one liver disease can affect each to a different extent • Liver diseases cause: – Altered liver functions – Altered tests of liver injury Additional Source Information for more information see: http://open.umich.edu/wiki/AttributionPolicy

Slide 73, Image 1 (top): Martybugs, "Jaundice phototherapy," Wikimedia Commons, http://commons.wikimedia.org/wiki/File:Jaundice_phototherapy.jpg, CC; BY-SA 3.0, http://creativecommons.org/licenses/by-sa/3.0/.