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Liver

For other uses, see Liver (disambiguation). 1 Structure


The liver is a vital organ only found in vertebrates.[2][3]
The liver is a reddish-brown wedge-shaped organ with
four lobes of unequal size and shape. A human liver
normally weighs 1.441.66 kg (3.23.7 lb),[10] and has
a width of about 15 cm.[11] It is both the heaviest internal
organ and the largest gland in the human body. Located in
the right upper quadrant of the abdominal cavity, it rests
just below the diaphragm, to the right of the stomach and
overlies the gallbladder.[5]
The liver is connected to two large blood vessels: the
hepatic artery and the portal vein. The hepatic artery
carries oxygen-rich blood from the aorta, whereas the
portal vein carries blood rich in digested nutrients from
the entire gastrointestinal tract and also from the spleen
and pancreas.[9] These blood vessels subdivide into small
Labeled human liver capillaries known as liver sinusoids, which then lead to a
lobule.
Lobules are the functional units of the liver. Each lob-
ule is made up of millions of hepatic cells (hepatocytes)
In humans, it is located in the upper right quadrant of which are the basic metabolic cells. The lobules are held
the abdomen, below the diaphragm. The liver has a together by a ne dense irregular broelastic connective
wide range of functions, including detoxication of vari- tissue layer which extends into the structure of the liver,
ous metabolites, protein synthesis, and the production of by accompanying the vessels (veins and arteries), ducts
biochemicals necessary for digestion.[4] It also plays a role and nerves through the hepatic portal, as a brous cap-
in metabolism, regulation of glycogen storage, decompo- sule called Glissons capsule.[12] The whole surface of
sition of red blood cells and hormone production.[4] the liver is covered in a serous coat derived from peri-
toneum and this has an inner brous coat (Glissons cap-
The liver is a gland. It is an accessory digestive gland
sule) to which it is rmly adhered. The brous coat is of
and produces bile, an alkaline compound which aids in
areolar tissue and follows the vessels and ducts to support
digestion via the emulsication of lipids. The gallbladder,
them.[13]
a small pouch that sits just under the liver, stores bile
produced by the liver.[5] The livers highly specialized
tissue consisting of mostly hepatocytes regulates a wide 1.1 Gross anatomy
variety of high-volume biochemical reactions, includ-
ing the synthesis and breakdown of small and complex 1.1.1 Lobes
molecules, many of which are necessary for normal vital
functions.[6] Estimates regarding the organs total num- Gross anatomy traditionally divided the liver into two
ber of functions vary, but textbooks generally cite it being portions a right and a left lobe, as viewed from the front
around 500.[7] (diaphragmatic) surface; but the underside (the visceral
Terminology related to the liver often starts in hepat- from surface) shows it to be divided into four lobes and includes
-, the Greek word for liver.[8] the caudate and quadrate lobes.[14]
There is currently no way to compensate for the absence The falciform ligament, visible on the front of the liver,
of liver function in the long term, although liver dialy- divides the liver into a left and a much larger right lobe.
sis techniques can be used in the short term. Articial From the visceral surface, the two additional lobes are
livers are yet to be developed to promote long-term re- located between the right and left lobes, one in front of
placement in the absence of the liver. As of now,[9] liver the other. A line can be imagined running from the left
transplantation is the only option for complete liver fail- of the vena cava and all the way forward to divide the
ure. liver and gallbladder into two halves.[15] This line is called

1
2 1 STRUCTURE

ments have no known functional importance, though they


serve as surface landmarks.[19] The falciform ligament
functions to attach the liver to the posterior portion of
the anterior body wall.
The visceral surface or inferior surface, is uneven and
concave. It is covered in peritoneum apart from where it
attaches the gallbladder and the porta hepatis.[18]

1.1.3 Impressions

The upper surface of the liver showing two lobes

Impressions of the liver


Visceral surface showing four lobes and the impressions
There are several impressions on the surface of the liver
[16] which accommodate the various adjacent structures and
"Cantlies line".
organs. Underneath the right lobe and to the right of the
Other anatomical landmarks exist, such as the gallbladder fossa, are two impressions, one behind the
ligamentum venosum and the round ligament of other and separated by a ridge. The one in front is a shal-
the liver (ligamentum teres), which further divide the low colic impression, formed by the hepatic exure and
left side of the liver in two sections. An important the one behind is a deeper renal impression accommo-
anatomical landmark, the porta hepatis, also known as dating part of the right kidney and part of the suprarenal
the transverse ssure of the liver, divides this left portion gland.[20]
into four segments, which can be numbered starting at
The suprarenal impression is a small triangular de-
the caudate lobe as I in an anticlockwise manner. From
pressed area on the liver. It is located close to the right
this visceral view, seven segments can be seen, because
[17] of the fossa between the bare area and the caudate lobe
the eighth segment is only visible in the parietal view.
and immediately above the renal impression. The greater
part of the suprarenal impression is devoid of peritoneum
1.1.2 Surfaces and it lodges the right suprarenal gland.[21]
Medial to the renal impression is a third and slightly
On the diaphragmatic surface, apart from a small tri- marked impression, lying between it and the neck of the
angular bare area where it connects to the diaphragm, gall-bladder. This is caused by the descending portion of
the liver is covered by a thin double-layered membrane, the duodenum, and is known as the duodenal impres-
the peritoneum, that helps to reduce friction against other sion.[21]
organs.[18] This surface covers the convex shape of the The inferior surface of the left lobe of the liver presents
two lobes where it accommodates the shape of the di- behind and to the left the gastric impression.[21] This is
aphragm. The peritoneum folds back on itself to form moulded over the upper front surface of the stomach, and
the falciform ligament and the right and left triangular to the right of this is a rounded eminence, the tuber omen-
ligaments.[19] tale, which ts into the concavity of the lesser curvature
These peritoneal ligaments are not related to the anatomic of the stomach and lies in front of the anterior layer of
ligaments in joints, and the right and left triangular liga- the lesser omentum.
1.3 Functional anatomy 3

portal venule
bile ductule through which blood from the hepatic portal vein and hep-
Kuper cell
sinusoid atic artery enters via the portal triads, then drains to the
portal central vein.[22]
eld

central Histology, the study of microscopic anatomy, shows two


vein
major types of liver cell: parenchymal cells and non-
stellate cell parenchymal cells. 7085% of the liver volume is oc-
hepatic arteriole
space of Disse
cupied by parenchymal hepatocytes. Non-parenchymal
hepatocyte endothelial cell
cells constitute 40% of the total number of liver cells
Cells, ducts and blood vessels
but only 6.5% of its volume.[24] The liver sinusoids are
lined with two types of cell, sinusoidal endothelial cells,
and phagocytic Kuper cells.[25] Hepatic stellate cells are
non-parenchymal cells found in the perisinusoidal space,
between a sinusoid and a hepatocyte.[24] Additionally, in-
trahepatic lymphocytes are often present in the sinusoidal
lumen.[24]

1.3 Functional anatomy


The central area or hilum, known as the porta hepatis is
where the common bile duct, hepatic portal vein, and the
hepatic artery proper enter the liver. The duct, vein, and
artery divide into left and right branches, and the areas of
the liver supplied by these branches constitute the func-
tional left and right lobes. The functional lobes are sep-
arated by the imaginary plane, Cantlies line, joining the
gallbladder fossa to the inferior vena cava. The plane sep-
arates the liver into the true right and left lobes. The mid-
dle hepatic vein also demarcates the true right and left
lobes. The right lobe is further divided into an anterior
and posterior segment by the right hepatic vein. The left
lobe is divided into the medial and lateral segments by the
Microscopic anatomy of the liver left hepatic vein.

1.4 Couinaud classication system

Types of capillariessinusoid on right

1.2 Microscopic anatomy

Microscopically, each liver lobe is seen to be made up


of hepatic lobules. The lobules are roughly hexagonal,
and consist of plates of hepatocytes radiating from a cen-
tral vein.[22] The central vein joins to the hepatic vein to
carry blood out from the liver. A distinctive compo-
nent of a lobule is the portal triad, which can be found
running along each of the lobules corners. The portal
triad, misleadingly named, consists of ve structures: a
branch of the hepatic artery, a branch of the hepatic por-
tal vein, and a bile duct, as well as lymphatic vessels and
Shape of human liver in animation. Eight subsegments by Couin-
a branch of the vagus nerve.[23] Between the hepatocyte aud labelled.
plates are liver sinusoids, which are enlarged capillaries
4 2 DEVELOPMENT

unit VI is located more posteriorly


Right lobe Left lobe Left lobe Right lobe

Right (part of) liver Left (part of) liver Left (part of) liver Right (part of) liver

Right Right Left Left Left Left Right Right


posterior
section
anterior
section
medial
section
lateral
section
lateral
section
medial
section
anterior
section
posterior
section unit VII is located above unit VI

unit VIII sits above unit V in the superio-medial po-


sition

Umbilical ssure Umbilical ssure


(Falciform ligament) (Falciform ligament)
Cantlie's line Cantlie's line 2 Development
Anterior view Posterior view

Organogenesis, the development of the organs takes place


Couinaud classication system from the third to the eighth week during embryogenesis.
The origins of the liver lie in both the ventral portion
The ssure for the round ligament of the liver (ligamen- of the foregut endoderm (endoderm being one of the 3
tum teres) also separates the medial and lateral segments. embryonic germ layers) and the constituents of the ad-
The medial segment is also called the quadrate lobe. In jacent septum transversum mesenchyme. In the human
the widely used Couinaud (or French) system, the func- embryo, the hepatic diverticulum is the tube of endo-
tional lobes are further divided into a total of eight sub- derm that extends out from the foregut into the surround-
segments based on a transverse plane through the bifurca- ing mesenchyme. The mesenchyme of septum transver-
tion of the main portal vein.[26] The caudate lobe is a sep- sum induces this endoderm to proliferate, to branch,
arate structure which receives blood ow from both the and to form the glandular epithelium of the liver. A
right- and left-sided vascular branches.[27][28] The Couin- portion of the hepatic diverticulum (that region clos-
aud classication of liver anatomy divides the liver into est to the digestive tube) continues to function as the
eight functionally independent segments. Each segment drainage duct of the liver, and a branch from this duct pro-
has its own vascular inow, outow and biliary drainage. duces the gallbladder.[31] Besides signals from the septum
In the centre of each segment there is a branch of the por- transversum mesenchyme, broblast growth factor from
tal vein, hepatic artery and bile duct. In the periphery of the developing heart also contributes to hepatic compe-
each segment there is vascular outow through the hepatic tence, along with retinoic acid emanating from the lateral
veins.[29] The division of the liver into independent units plate mesoderm. The hepatic endodermal cells undergo
means that segments can be resected without damaging a morphological transition from columnar to pseudos-
the remaining segments.[30] To preserve the viability of tratied resulting in thickening into the early liver bud.
the liver following surgery, resections follow the vessels Their expansion forms a population of the bipotential
dening the peripheries of each segment. This means that hepatoblasts.[32] Hepatic stellate cells are derived from
resection lines parallel the hepatic veins, leaving the por- mesenchyme.[33]
tal veins, bile ducts, and hepatic arteries intact.[26] After migration of hepatoblasts into the septum transver-
The classication system uses the vascular supply in the sum mesenchyme, the hepatic architecture begins to be
liver to separate the functional units (numbered I to VIII): established, with liver sinusoids and bile canaliculi ap-
pearing. The liver bud separates into the lobes. The left
umbilical vein becomes the ductus venosus and the right
Unit I is the caudate lobe and is situated posterior
vitelline vein becomes the portal vein. The expanding
l and it may receive its supply from both the right
liver bud is colonized by hematopoietic cells. The bipo-
and the left branches of portal vein. It contains one
tential hepatoblasts begin dierentiating into biliary ep-
or more hepatic veins which drain directly into the
ithelial cells and hepatocytes. The biliary epithelial cells
IVC.[26]
dierentiate from hepatoblasts around portal veins, rst
producing a monolayer, and then a bilayer of cuboidal
The remainder of the units (II to VIII) are numbered in a cells. In ductal plate, focal dilations emerge at points in
clockwise fashion:[29] the bilayer, become surrounded by portal mesenchyme,
and undergo tubulogenesis into intrahepatic bile ducts.
units II and III lie medial to the falciform ligament Hepatoblasts not adjacent to portal veins instead dieren-
with II superior to the portal venous supply and III tiate into hepatocytes and arrange into cords lined by sin-
inferior udoidal epithelial cells and bile canaliculi. Once hepato-
blasts are specied into hepatocytes and undergo further
unit IV lies lateral to the falciform ligament and is
expansion, they begin acquiring the functions of a ma-
subdivided into IVa (superior) and IVb (inferior)
ture hepatocyte, and eventually mature hepatocytes ap-
pear as highly polarized epithelial cells with abundant
Units V to VIII make up the right part of the liver:[29] glycogen accumulation. In the adult liver, hepatocytes
are not equivalent, with position along the portocentro-
unit V is the most medial and inferior venular axis within a liver lobule dictating expression of
3.2 Biliary ow 5

metabolic genes involved in drug metabolism, carbohy-


drate metabolism, ammonia detoxication, and bile pro-
duction and secretion. WNT/-catenin has now been
identied to be playing a key role in this phenomenon.[32]
At birth the liver comprises roughly 4% of body weight
and is at average 120 g. Over the course of further devel-
opment, it will increase to 1.41.6 kg but will only take
up 2.53.5% of body weight.[34]

2.1 Fetal blood supply


In the growing fetus, a major source of blood to the liver
Liver veins
is the umbilical vein which supplies nutrients to the grow-
ing fetus. The umbilical vein enters the abdomen at the
umbilicus, and passes upward along the free margin of livers oxygen demand is met by the hepatic portal vein,
the falciform ligament of the liver to the inferior surface and half is met by the hepatic arteries.[35]
of the liver. There it joins with the left branch of the por-
Blood ows through the liver sinusoids and empties into
tal vein. The ductus venosus carries blood from the left
the central vein of each lobule. The central veins coalesce
portal vein to the left hepatic vein and then to the inferior
into hepatic veins, which leave the liver and drain into the
vena cava, allowing placental blood to bypass the liver.
inferior vena cava.[23]
In the fetus, the liver does not perform the normal di-
gestive processes and ltration of the infant liver because
nutrients are received directly from the mother via the 3.2 Biliary ow
placenta. The fetal liver releases some blood stem cells
that migrate to the fetal thymus, creating the T-cells or
T-lymphocytes. After birth, the formation of blood stem
Liver
cells shifts to the red bone marrow.
After two to ve days, the umbilical vein and ductus
venosus are completely obliterated; the former becomes
the round ligament of liver and the latter becomes the
ligamentum venosum. In the disorders of cirrhosis and
portal hypertension, the umbilical vein can open up again. Perihilar
bile ducts

Common
3 Physiology bile duct
Bowel
Pancreas
The various functions of the liver are carried out by the (duodenum)
liver cells or hepatocytes. The liver is thought to be re-
sponsible for up to 500 separate functions, usually in com-
bination with other systems and organs. Currently, there Biliary tract
is no articial organ or device capable of reproducing all
the functions of the liver. Some functions can be carried The biliary tract is derived from the branches of the bile
out by liver dialysis, an experimental treatment for liver ducts. The biliary tract, also known as the biliary tree,
failure. is the path by which bile is secreted by the liver then
transported to the rst part of the small intestine, the
duodenum. The bile produced in the liver is collected in
3.1 Blood supply bile canaliculi, small grooves between the faces of adja-
cent hepatocytes. The canaliculi radiate to the edge of the
The liver receives a dual blood supply from the hepatic liver lobule, where they merge to form bile ducts. Within
portal vein and hepatic arteries. The hepatic portal vein the liver, these ducts are termed intrahepatic bile ducts,
delivers approximately 75% of the livers blood sup- and once they exit the liver they are considered extrahep-
ply, and carries venous blood drained from the spleen, atic. The intrahepatic ducts eventually drain into the right
gastrointestinal tract, and its associated organs. The hep- and left hepatic ducts, which exit the liver at the transverse
atic arteries supply arterial blood to the liver, accounting ssure, and merge to form the common hepatic duct. The
for the remaining quarter of its blood ow. Oxygen is cystic duct from the gallbladder joins with the common
provided from both sources; approximately half of the hepatic duct to form the common bile duct.[23]
6 4 CLINICAL SIGNIFICANCE

Bile either drains directly into the duodenum via the com- 3.4 Breakdown
mon bile duct, or is temporarily stored in the gallbladder
via the cystic duct. The common bile duct and the The liver is responsible for the breakdown of insulin and
pancreatic duct enter the second part of the duodenum other hormones. The liver breaks down bilirubin via
together at the hepatopancreatic ampulla, also known asglucuronidation, facilitating its excretion into bile. The
the ampulla of Vater. liver is responsible for the breakdown and excretion of
many waste products. It plays a key role in breaking
down or modifying toxic substances (e.g., methylation)
and most medicinal products in a process called drug
metabolism. This sometimes results in toxication, when
the metabolite is more toxic than its precursor. Prefer-
3.3 Synthesis ably, the toxins are conjugated to avail excretion in bile
or urine. The liver breaks down ammonia into urea as part
Further information: Proteins produced and secreted by of the urea cycle, and the urea is excreted in the urine.[22]
the liver

3.5 Other functions


The liver plays a major role in carbohydrate, protein,
amino acid, and lipid metabolism.
The liver stores a multitude of substances, including
The liver performs several roles in carbohydrate glucose (in the form of glycogen), vitamin A (1
metabolism: The liver synthesizes and stores ap- 2 years supply), vitamin D (14 months supply),
proximately 100g of glycogen via glycogenesis, the vitamin B12 (35 years supply),[38] vitamin K, iron,
formation of glycogen from glucose. When needed, and copper.
the liver releases glucose into the blood by per-
forming glycogenolysis, the breakdown of glycogen The liver is responsible for immunological eects
into glucose.[36] The liver is also responsible for the mononuclear phagocyte system of the liver con-
gluconeogenesis, which is the synthesis of glucose from tains many immunologically active cells, acting as a
certain amino acids, lactate or glycerol. Adipose and 'sieve' for antigens carried to it via the portal system.
liver cells produce glycerol by breakdown of fat, which
The liver produces albumin, the most abundant pro-
the liver uses for gluconeogenesis.[36]
tein in blood serum. It is essential in the mainte-
The liver is responsible for the mainstay of protein nance of oncotic pressure, and acts as a transport
metabolism, synthesis as well as degradation. It is also for fatty acids and steroid hormones.
responsible for a large part of amino acid synthesis. The
liver plays a role in the production of clotting factors as The liver synthesizes angiotensinogen, a hormone
well as red blood cell production. Some of the proteins that is responsible for raising the blood pressure
synthesized by the liver include coagulation factors I (b- when activated by renin, an enzyme that is released
rinogen), II (prothrombin), V, VII, VIII, IX, X, XI, XIII, when the kidney senses low blood pressure.
as well as protein C, protein S and antithrombin. In the
rst trimester fetus, the liver is the main site of red blood The liver produces the enzyme catalase in order to
cell production. By the 32nd week of gestation, the bone break down hydrogen peroxide, a very toxic sub-
marrow has almost completely taken over that task. The stance due to it being a powerful oxidising agent,
liver is a major site of production for thrombopoietin, into water and oxygen.
a glycoprotein hormone that regulates the production of
platelets by the bone marrow.[37]
3.6 Relation to medicine and pharmacol-
The liver plays several roles in lipid metabolism: it per- ogy
forms cholesterol synthesis, lipogenesis, the production
of triglycerides, and a bulk of the bodys lipoproteins are The oxidative capacity of the liver decreases with aging
synthesized in the liver. and therefore any medications that require oxidation (for
The liver plays a key role in digestion, as it produces and instance, benzodiazepines) are more likely to accumulate
excretes bile (a yellowish liquid) required for emulsifying to toxic levels. However, medications with shorter half-
fats and help the absorption of vitamin K from the diet. lives, such as lorazepam and oxazepam, are preferred in
Some of the bile drains directly into the duodenum, and most cases when benzodiazepines are required in regard
some is stored in the gallbladder. to geriatric medicine.
The liver also produces insulin-like growth factor 1 (IGF-
1), a polypeptide protein hormone that plays an important
role in childhood growth and continues to have anabolic 4 Clinical signicance
eects in adults.
4.2 Symptoms 7

4.1 Disease build up in the liver (cholestasis) and over time damages
the liver tissue in combination with ongoing immune re-
Main article: Liver disease lated damage. This can lead to scarring (brosis) and
The liver is a vital organ and supports almost every cirrhosis.
Many diseases of the liver are accompanied by jaundice
caused by increased levels of bilirubin in the system. The
bilirubin results from the breakup of the hemoglobin of
dead red blood cells; normally, the liver removes bilirubin
from the blood and excretes it through bile.
There are also many pediatric liver diseases, includ-
ing biliary atresia, alpha-1 antitrypsin deciency, alagille
syndrome, progressive familial intrahepatic cholestasis,
Langerhans cell histiocytosis and hepatic hemangioma a
benign tumour the most common type of liver tumour,
thought to be congenital. Diseases that interfere with
liver function will lead to derangement of these processes.
However, the liver has a great capacity to regenerate and
has a large reserve capacity. In most cases, the liver only
Left lobe liver tumor produces symptoms after extensive damage.
Hepatomegaly refers to an enlarged liver and can be due
other organ in the body. Because of its strategic location to many causes. It can be palpated in a liver span mea-
and multidimensional functions, the liver is also prone to surement.
many diseases.[39] The bare area of the liver is a site that is
vulnerable to the passing of infection from the abdominal Liver diseases may be diagnosed by liver function tests
cavity to the thoracic cavity. blood tests that can identify various markers. For exam-
ple, acute-phase reactants are produced by the liver in re-
Hepatitis is a common condition of inammation of the sponse to injury or inammation.
liver. The most usual cause of this is viral, and the most
common of these infections are hepatitis A, B, C, D, and
E. Some of these infections are sexually transmitted. In- 4.2 Symptoms
ammation can also be caused by other viruses in the
Herpesviridae family such as the herpes simplex virus. In- The classic symptoms of liver damage include the follow-
fection with hepatitis B virus or hepatitis C virus is the ing:
main cause of liver cancer.[40]
Pale stools occur when stercobilin, a brown pig-
Hepatic encephalopathy is caused by an accumulation of
ment, is absent from the stool. Stercobilin is derived
toxins in the bloodstream that are normally removed by
from bilirubin metabolites produced in the liver.
the liver. This condition can result in coma and can prove
fatal. Dark urine occurs when bilirubin mixes with urine
Other disorders caused by excessive alcohol consumption Jaundice (yellow skin and/or whites of the eyes)
are grouped under alcoholic liver diseases and these in- This is where bilirubin deposits in skin, causing an
clude alcoholic hepatitis, fatty liver, and cirrhosis. Fac- intense itch. Itching is the most common complaint
tors contributing to the development of alcoholic liver by people who have liver failure. Often this itch can-
diseases are not only the quantity and frequency of alco- not be relieved by drugs.
hol consumption, but can also include gender, genetics,
and liver insult. Swelling of the abdomen, ankles and feet occurs be-
cause the liver fails to make albumin.
Liver damage can also be caused by drugs, particularly
paracetamol and drugs used to treat cancer. Excessive fatigue occurs from a generalized loss of
nutrients, minerals and vitamins.
BuddChiari syndrome is a condition caused by blockage
of the hepatic veins (including thrombosis) that drain the Bruising and easy bleeding are other features of
liver. It presents with the classical triad of abdominal liver disease. The liver makes substances which help
pain, ascites and liver enlargement.[41] prevent bleeding. When liver damage occurs, these
substances are no longer present and severe bleeding
Primary biliary cirrhosis is an autoimmune disease of the
can occur.[45]
liver.[42][43] It is marked by slow progressive destruction
of the small bile ducts of the liver, with the intralobular Pain in the upper right quadrant can result from
ducts (Canals of Hering) aected early in the disease.[44] the stretching of Glissons capsule in conditions of
When these ducts are damaged, bile and other toxins hepatitis and pre-eclampsia.
8 4 CLINICAL SIGNIFICANCE

4.3 Diagnosis 4.4 Biopsy / scan

The diagnosis of liver disease is made by liver function Damage to the liver is sometimes determined with a
tests, groups of blood tests, that can readily show the biopsy, particularly when the cause of liver damage is
extent of liver damage. If infection is suspected, then unknown. In the 21st century they have been largely re-
other serological tests will be carried out. Sometimes, an placed by high-resolution radiographic scans. The latter
ultrasound or a CT scan is needed to produce an image do not require ultrasound guidance, lab involvement, mi-
of the liver. croscopic analysis, organ damage, pain, or patient seda-
Physical examination of the liver can only reveal its size tion; and the results are available immediately on a com-
and any tenderness, and some form of imaging will also puter screen.
be needed.[46] In a biopsy, a needle is inserted into the skin just be-
low the rib cage and a tissue sample obtained. The tis-
sue is sent to the laboratory, where it is analyzed under
a microscope. Sometimes, a radiologist may assist the
physician performing a liver biopsy by providing ultra-
sound guidance.[48]

4.5 Liver regeneration


Axial CT image showing
anomalous hepatic veins coursing on the subcapsu-
Main article: Liver regeneration
lar anterior surface of the liver.[47]

The liver is the only human internal organ capable of nat-


ural regeneration of lost tissue; as little as 25% of a liver
can regenerate into a whole liver.[49] This is, however,
not true regeneration but rather compensatory growth in
mammals.[50] The lobes that are removed do not regrow
and the growth of the liver is a restoration of function,
not original form. This contrasts with true regenera-
tion where both original function and form are restored.
Maximum intensity pro- In some other species, such as sh, the liver undergoes
jection (MIP) CT image as viewed anteriorly true regeneration by restoring both shape and size of the
showing the anomalous hepatic veins coursing on organ.[51] In the liver, large areas of the tissues are formed
the anterior surface of the liver but for the formation of new cells there must be su-
cient amount of material so the circulation of the blood
becomes more active.[52]
This is predominantly due to the hepatocytes re-entering
the cell cycle. That is, the hepatocytes go from the quies-
cent G0 phase to the G1 phase and undergo mitosis. This
process is activated by the p75 receptors.[53] There is also
some evidence of bipotential stem cells, called hepatic
oval cells or ovalocytes (not to be confused with oval red
blood cells of ovalocytosis), which are thought to reside
Lateral MIP view in the
in the canals of Hering. These cells can dierentiate into
same patient
either hepatocytes or cholangiocytes. Cholangiocytes are
the epithelial lining cells of the bile ducts.[54] They are
cuboidal epithelium in the small interlobular bile ducts,
but become columnar and mucus secreting in larger bile
ducts approaching the porta hepatis and the extrahepatic
ducts.
Scientic and medical works about liver regeneration of-
ten refer to the Greek Titan Prometheus who was chained
to a rock in the Caucasus where, each day, his liver was
A CT scan in which the liver devoured by an eagle, only to grow back each night. The
and portal vein are shown. myth suggests the ancient Greeks may have known about
the livers remarkable capacity for self-repair.[55]
9

4.6 Liver transplantation magnetic resonance imaging (MRI). MDCT is good in


vascular anatomy and volumetry. MRI is used for bil-
Main article: Liver transplantation iary tree anatomy. Donors with very unusual vascu-
lar anatomy, which makes them unsuitable for donation,
Human liver transplants were rst performed by Thomas could be screened out to avoid unnecessary operations.
Starzl in the United States and Roy Calne in Cambridge,
England in 1963 and 1967, respectively.

MDCT image. Arterial


anatomy contraindicated for liver donation

After resection of left lobe liver tumor MDCT image. Portal ve-
nous anatomy contraindicated for liver donation
Liver transplantation is the only option for those with
irreversible liver failure. Most transplants are done for
chronic liver diseases leading to cirrhosis, such as chronic
hepatitis C, alcoholism, autoimmune hepatitis, and many
others. Less commonly, liver transplantation is done for
fulminant hepatic failure, in which liver failure occurs
over days to weeks. MDCT image. 3D image
Liver allografts for transplant usually come from donors created by MDCT can clearly visualize the liver,
who have died from fatal brain injury. Living donor liver measure the liver volume, and plan the dissection
transplantation is a technique in which a portion of a liv- plane to facilitate the liver transplantation proce-
ing persons liver is removed and used to replace the entire dure.
liver of the recipient. This was rst performed in 1989
for pediatric liver transplantation. Only 20 percent of an
adults liver (Couinaud segments 2 and 3) is needed to
serve as a liver allograft for an infant or small child.
More recently, adult-to-adult liver transplantation has
been done using the donors right hepatic lobe, which
amounts to 60 percent of the liver. Due to the ability of Phase contrast CT image.
the liver to regenerate, both the donor and recipient end Contrast is perfusing the right liver but not the left
up with normal liver function if all goes well. This proce- due to a left portal vein thrombus.
dure is more controversial, as it entails performing a much
larger operation on the donor, and indeed there have been
at least two donor deaths out of the rst several hundred
cases. A recent publication has addressed the problem of 5 Society and culture
donor mortality, and at least 14 cases have been found.[56]
The risk of postoperative complications (and death) is far In Greek mythology, Prometheus was punished by the
greater in right-sided operations than that in left-sided op- gods for revealing re to humans, by being chained to
erations. a rock where a vulture (or an eagle) would peck out his
With the recent advances of noninvasive imaging, living liver, which would regenerate overnight. (The liver is the
liver donors usually have to undergo imaging examina- only human internal organ that actually can regenerate it-
tions for liver anatomy to decide if the anatomy is feasi- self to a signicant extent.) Many ancient peoples of the
ble for donation. The evaluation is usually performed by Near East and Mediterranean areas practiced a type of
multidetector row computed tomography (MDCT) and divination called haruspicy, where they tried to obtain in-
10 6 OTHER ANIMALS

formation by examining the livers of sheep and other an- tional South African delicacy, namely skilpadjies, is made
imals. of minced lambs liver wrapped in netvet (caul fat), and
In Plato, and in later physiology, the liver was thought to grilled over an open re.
be the seat of the darkest emotions (specically wrath, Animal livers are rich in iron and vitamin A, and cod liver
jealousy and greed) which drive men to action.[57] The oil is commonly used as a dietary supplement. Tradition-
Talmud (tractate Berakhot 61b) refers to the liver as the ally, some sh livers were valued as food, especially the
seat of anger, with the gallbladder counteracting this. stingray liver. It was used to prepare delicacies, such as
The Persian, Urdu, and Hindi languages ( or poached skate liver on toast in England, as well as the
or jigar) refer to the liver in gurative speech to indicate beignets[60]
de foie de raie and foie de raie en croute in French
courage and strong feelings, or their best"; e.g., This cuisine.
Mecca has thrown to you the pieces of its liver!".[58] The
term jan e jigar, literally the strength (power) of my
liver, is a term of endearment in Urdu. In Persian slang,
jigar is used as an adjective for any object which is desir-
able, especially women. In the Zulu language, the word
for liver (isibindi) is the same as the word for courage.
The legend of Liver-Eating Johnson says that he would 6 Other animals
cut out and eat the liver of each man killed after dinner.
In the motion picture The Message, Hind bint Utbah is
implied or portrayed eating the liver of Hamza ibn Abd
al-Muttalib during the Battle of Uhud. Although there are
narrations that suggest that Hind did taste, rather than
eat, the liver of Hamza, the authenticity of these narra-
tions has to be questioned.
On November 26, 1987, the city of Ferrol, Spain, inau-
gurated what is believed to be the only monument to the
liver in the world. The then-mayor, Jaime Quintanilla,
also happened to be a doctor, and thought appropriate
to promote the monument. At an approximate cost of
$3.200, the monument stands in the village of Baln.
A plaque reads (In Galician language, free translation):
The Liver [is the] basis of Life, and below Through
History, Mankind tried to cure all illness. By helping it
on this duty, you are doing a great job. We are grateful
for it.[59]
Sheeps liver

5.1 Food
The liver is found in all vertebrates, and is typically the
Main article: Liver (food) largest visceral (internal) organ. Its form varies consid-
erably in dierent species, and is largely determined by
The liver of mammals, fowl, and sh are commonly eaten the shape and arrangement of the surrounding organs.
as food by humans. Domestic pig, ox, lamb, calf, chicken, Nonetheless, in most species it is divided into right and
and goose livers are widely available from butchers and left lobes; exceptions to this general rule include snakes,
supermarkets. where the shape of the body necessitates a simple cigar-
Liver can be baked, boiled, broiled, fried, stir-fried, like form. The internal structure of the liver is broadly
or eaten raw (asbeh nayeh or sawda naye in Lebanese similar in all vertebrates.[61]
cuisine, liver sashimi). In many preparations, pieces An organ sometimes referred to as a liver is found as-
of liver are combined with pieces of meat or kidneys, sociated with the digestive tract of the primitive chor-
like in the various forms of Middle Eastern mixed grill date Amphioxus. Although it performs many functions
(e.g. meurav Yerushalmi). Liver is often made into of a liver, it is not considered a true liver but a homolog
spreads. Well-known examples include liver pt, foie of the vertebrate liver.[62][63][64] The amphioxus hepatic
gras, chopped liver, and leverpastej. Liver sausages such caecum produces the liver-specic proteins vitellogenin,
as Braunschweiger and liverwurst are also a valued meal. antithrombin, plasminogen, alanine aminotransferase,
Liver sausages may also be used as spreads. A tradi- and insulin/Insulin-like growth factor (IGF)[65]
11

7 Additional images area

Liver
View of the various organs
and blood-vessels in proximity with liver.

8 See also
Articial liver

Liver lifted to show gall Enterohepatic circulation


bladder and stomach in situ.
Hepatectomy

Liver shot (martial arts strike)

Polycystic liver disease

Portacaval anastomosis
Cross section showing the
liver as the large brown mass in the left of the Portal hypertension
images, right of the individual.

9 References
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[55] An argument for the ancient Greeks knowing about liver


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(1994). The myth of Prometheus and the liver. Jour-
nal of the Royal Society of Medicine. 87 (12): 754755. Liver at the Open Directory Project.
PMC 1294986 . PMID 7853302. Counterarguments
are provided by Tiniakos, D. G.; Kandilis, A.; Geller, VIRTUAL Liver online learning resource
S. A. (2010). Tityus: A forgotten myth of liver re- Liver enzymes
generation. Journal of Hepatology. 53 (2): 357361.
14 11 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

11 Text and image sources, contributors, and licenses


11.1 Text
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0, Berek, Taw, Alex.tan, BenBaker, Camembert, Icarus~enwiki, Sfdan, Someone else, Stevertigo, JohnOwens, Liftarn, Ixfd64, Dori,
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Diberri, Ancheta Wis, Giftlite, Christopher Parham, Marnanel, Nlevitt, Tom harrison, Peruvianllama, Everyking, Moyogo, Alison, Jfd-
wol, Jrdioko, Kandar, OldakQuill, Chowbok, Utcursch, Knutux, DCrazy, Antandrus, Rdsmith4, CnorthMSU, Ilgiz, Balcer, Borameer,
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cospinster, Ardonik, Bishonen, Tim Peterson, Paul August, MarkS, Edgarde, Bender235, Rubicon, Kbh3rd, Jnestorius, Tv@nlntv.com,
Violetriga, Aecis, NickGorton~enwiki, Art LaPella, RoyBoy, Jpgordon, Causa sui, Bobo192, Smalljim, Clawson, Olve Utne, Clarkbhm,
Arcadian, Jag123, Nk, Larryv, PiccoloNamek, Obradovic Goran, MPerel, Merope, Espoo, Alansohn, Gary, Keenan Pepper, Tabor, Wouter-
stomp, Riana, Kurt Shaped Box, SlimVirgin, YDZ, Gaurav1146, Pion, VladimirKorablin, Ombudsman, MoraSique, Wtmitchell, Velella,
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decency, User A1, DerHexer, JaGa, Lenticel, Rickterp, Scottalter, Hedwig in Washington, Flowanda, Ksero, MartinBot, Ectoplasmi-
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carrot, NewEnglandYankee, Russellfsm, Bigdumbdinosaur, Shshshsh, KylieTastic, Don Cdfd!!!, Jamesontai, Jto410, Pdcook, Permafrost,
TheNewPhobia, Ichormosquito, Squids and Chips, VIOLENTRULER, Smrolando, Idioma-bot, GylesM, Frammo, Wikieditor06, Roy-
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LeaveSleaves, Slimshaidy, KC Panchal, DeathbyChiasmus, Ilyushka88, Unoluigi, Mwilso24, Liveranalyzer, Lerdthenerd, Andy Dingley,
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soccer, Keilana, MilFlyboy, Andymac345, Flyer22 Reborn, Oda Mari, Chris112358, Cablehorn, Momo san, Oxymoron83, KPH2293,
Steven Crossin, Tombomp, Kahnusc81, Crisis, Nskillen, OKBot, LearnAnatomy, Hackmefoo, Anchor Link Bot, Superbeecat, Bditwa,
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Jncraton, Kobeybryant, Doctormanhattan, CanadianLinuxUser, Diptanshu Das, EconoPhysicist, Suseno, D.c.camero, Glane23, Twanjik,
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11.2 Images 15

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Tommy2010, Your Lord and Master, Wikipelli, Dcirovic, K6ka, Chloeebaybee, AvicBot, HiW-Bot, ZroBot, Wiensgov, Anir1uph,
Fomenka, RAWRMM, Ebrambot, SporkBot, Wayne Slam, Insane.ShaneRP, OnePt618, Lji1942, Brandmeister, Monostitch, Donner60, I
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11.2 Images
File:2104_Three_Major_Capillary_Types.jpg Source: https://upload.wikimedia.org/wikipedia/commons/8/8d/2104_Three_Major_
Capillary_Types.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/1.
6/, Jun 19, 2013. Original artist: OpenStax College
File:2423_Microscopic_Anatomy_of_Liver.jpg Source: https://upload.wikimedia.org/wikipedia/commons/b/b7/2423_Microscopic_
Anatomy_of_Liver.jpg License: CC BY 3.0 Contributors: Anatomy & Physiology, Connexions Web site. http://cnx.org/content/col11496/
1.6/, Jun 19, 2013. Original artist: OpenStax College
File:Anatomy_Abdomen_Tiesworks.jpg Source: https://upload.wikimedia.org/wikipedia/commons/1/15/Anatomy_Abdomen_
Tiesworks.jpg License: Public domain Contributors: Own work Original artist: Tvanbr
File:Anterior_MIP_image_of_anomalous_hepatic_veins.jpg Source: https://upload.wikimedia.org/wikipedia/en/c/c3/Anterior_
MIP_image_of_anomalous_hepatic_veins.jpg License: CC-BY-SA-3.0 Contributors:
clinical work
Original artist:
Jto410
File:Axial_CT_anomalous_hepatic_veins.jpg Source: https://upload.wikimedia.org/wikipedia/en/2/28/Axial_CT_anomalous_
hepatic_veins.jpg License: CC-BY-SA-3.0 Contributors:
clinical work
Original artist:
Jto410
File:Big_Liver_Tumor.JPG Source: https://upload.wikimedia.org/wikipedia/commons/9/9b/Big_Liver_Tumor.JPG License: GFDL
Contributors: haitham alfalah Original artist: haitham alfalah
File:CTscanofmyLiver.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/6a/CTscanofmyLiver.jpg License: CC BY-SA
3.0 Contributors: Own work Original artist: RGshredfox
File:Commons-logo.svg Source: https://upload.wikimedia.org/wikipedia/en/4/4a/Commons-logo.svg License: PD Contributors: ? Origi-
nal artist: ?
File:Diagram_showing_the_position_of_the_perihilar_bile_ducts_CRUK_357.svg Source: https://upload.wikimedia.org/
wikipedia/commons/1/1d/Diagram_showing_the_position_of_the_perihilar_bile_ducts_CRUK_357.svg License: CC BY-SA 4.0
Contributors: Original email from CRUK Original artist: Cancer Research UK
File:Gray1085.png Source: https://upload.wikimedia.org/wikipedia/commons/b/ba/Gray1085.png License: Public domain Contributors:
Henry Gray (1918) Anatomy of the Human Body (See Book section below)
Original artist: Henry Vandyke Carter
File:Gray1087-liver.png Source: https://upload.wikimedia.org/wikipedia/commons/a/a3/Gray1087-liver.png License: Public domain
Contributors: Henry Gray (1918) Anatomy of the Human Body (See Book section below)
Original artist: Henry Vandyke Carter
File:Hepatic_structure2.svg Source: https://upload.wikimedia.org/wikipedia/commons/9/92/Hepatic_structure2.svg License: CC BY
2.5 Contributors: Based on the research article Intravital Observation of Plasmodium berghei Sporozoite Infection of the Liver, PLoS
Biology, doi:10.1371/journal.pbio.0030192.g011 Original artist: Originally by Frevert U, Engelmann S, Zougbd S, Stange J, Ng B, et al.
Converted to SVG by Viacheslav Vtyurin who was hired to do so by User:Eug.
16 11 TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

File:LDLTA.jpg Source: https://upload.wikimedia.org/wikipedia/en/9/9c/LDLTA.jpg License: CC-BY-3.0 Contributors:


Dr. I-Chen Tsai in Taichung Veterans General Hospital, Taiwan. The picture is released under GNU FDL.
Original artist:
Dr. I-Chen Tsai
File:LDLTP.jpg Source: https://upload.wikimedia.org/wikipedia/en/f/f2/LDLTP.jpg License: CC-BY-3.0 Contributors:
Dr. I-Chen Tsai. Department of Radiology, Taichung Veterans General Hospital, Taichung, Taiwan.
Original artist:
Dr. I-Chen Tsai.
File:LDLT_volume_measure.jpg Source: https://upload.wikimedia.org/wikipedia/en/1/17/LDLT_volume_measure.jpg License: CC-
BY-3.0 Contributors:
Created by Dr. I-Chen Tsai, Taichung Veterans General Hospital, Taichung, Taiwan.
Original artist:
Dr. I-Chen Tsai, Taichung Veterans General Hospital, Taichung, Taiwan.
File:Labeled_Human_liver.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e2/Labeled_Human_liver.jpg License: CC
BY-SA 3.0 Contributors: Own work Original artist:
File:Lateral_MIP_CT_of_anomalous_surface_hepatic_veins.jpg Source: https://upload.wikimedia.org/wikipedia/en/6/65/Lateral_
MIP_CT_of_anomalous_surface_hepatic_veins.jpg License: CC-BY-SA-3.0 Contributors:
clinical work
Original artist:
Jto410
File:Leber_Schaf.jpg Source: https://upload.wikimedia.org/wikipedia/commons/6/64/Leber_Schaf.jpg License: CC-BY-SA-3.0 Con-
tributors: Uploaded on de:WP by de:Benutzer:Uwe Gille Original artist: de:Benutzer:Uwe Gille
File:Liver..JPG Source: https://upload.wikimedia.org/wikipedia/commons/3/3a/Liver..JPG License: GFDL Contributors: haitham alfalah
Original artist: haitham alfalah
File:Liver_(organ).png Source: https://upload.wikimedia.org/wikipedia/commons/5/58/Liver_%28organ%29.png License: CC BY-SA
4.0 Contributors: Own work Original artist: BruceBlaus
File:Liver_01_animation1.gif Source: https://upload.wikimedia.org/wikipedia/commons/8/88/Liver_01_animation1.gif License: CC
BY-SA 2.1 jp Contributors: Polygon data is from BodyParts3D[1] Original artist: Polygon data is generated by Database Center for Life
Science(DBCLS)[2]
File:Liver_04_Couinaud_classification.svg Source: https://upload.wikimedia.org/wikipedia/commons/e/e8/Liver_04_Couinaud_
classification.svg License: CC BY-SA 2.1 jp Contributors: Polygon data is from BodyParts3D[1] Original artist: Polygon data is generated
by Database Center for Life Science(DBCLS)[2]
File:Liver_04_Couinaud_classification_animation.gif Source: https://upload.wikimedia.org/wikipedia/commons/f/f1/Liver_04_
Couinaud_classification_animation.gif License: CC BY-SA 2.1 jp Contributors: Polygon data is from BodyParts3D[1] Original artist:
Polygon data is generated by Database Center for Life Science(DBCLS)[2]
File:Liver_veins.jpg Source: https://upload.wikimedia.org/wikipedia/commons/e/e5/Liver_veins.jpg License: CC-BY-SA-3.0 Contribu-
tors: http://commons.wikimedia.org/wiki/File:Galblaas_en_lever.jpg Original artist: Marcelo Reis
File:Lock-green.svg Source: https://upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg License: CC0 Contributors: en:File:
Free-to-read_lock_75.svg Original artist: User:Trappist the monk
File:R_vs_L_Liver_by_CT.PNG Source: https://upload.wikimedia.org/wikipedia/en/0/06/R_vs_L_Liver_by_CT.PNG License: PD
Contributors:
Naval Medical Center Portsmouth
Original artist:
Niels Olson
File:Slide2CHA.JPG Source: https://upload.wikimedia.org/wikipedia/commons/8/88/Slide2CHA.JPG License: CC BY-SA 3.0 Contrib-
utors: Own work Original artist: Anatomist90
File:Slide2CHAR.JPG Source: https://upload.wikimedia.org/wikipedia/commons/5/50/Slide2CHAR.JPG License: CC BY-SA 3.0 Con-
tributors: Own work Original artist: Anatomist90
File:Slide4CHA.JPG Source: https://upload.wikimedia.org/wikipedia/commons/5/51/Slide4CHA.JPG License: CC BY-SA 3.0 Contrib-
utors: Own work Original artist: Anatomist90
File:Sobo_1906_389.png Source: https://upload.wikimedia.org/wikipedia/commons/c/cc/Sobo_1906_389.png License: Public domain
Contributors: Sobottas Atlas and Text-book of Human Anatomy 1906 Original artist: Dr. Johannes Sobotta
File:Sobo_1906_393.png Source: https://upload.wikimedia.org/wikipedia/commons/1/1b/Sobo_1906_393.png License: Public domain
Contributors: Sobottas Atlas and Text-book of Human Anatomy 1906 Original artist: Dr. Johannes Sobotta
File:Sobo_1906_405.png Source: https://upload.wikimedia.org/wikipedia/commons/6/65/Sobo_1906_405.png License: Public domain
Contributors: Sobottas Atlas and Text-book of Human Anatomy 1906 Original artist: Dr. Johannes Sobotta
File:Sobo_1906_416.png Source: https://upload.wikimedia.org/wikipedia/commons/6/67/Sobo_1906_416.png License: Public domain
Contributors: Sobottas Atlas and Text-book of Human Anatomy 1906 Original artist: Dr. Johannes Sobotta
File:Sobo_1906_417.png Source: https://upload.wikimedia.org/wikipedia/commons/9/94/Sobo_1906_417.png License: Public domain
Contributors: Sobottas Atlas and Text-book of Human Anatomy 1906 Original artist: Dr. Johannes Sobotta
File:Wiktionary-logo-en-v2.svg Source: https://upload.wikimedia.org/wikipedia/commons/9/99/Wiktionary-logo-en-v2.svg License:
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