Liver

Liver
The liver is a major metabolic organ only

found in vertebrate animals, which
performs many essential biological
functions such as detoxification of the
organism, and the synthesis of proteins
and biochemicals necessary for
digestion and growth.[2][3][4] In humans, it
is located in the right upper quadrant of
the abdomen, below the diaphragm and
mostly shielded by the lower right rib
cage. Its other metabolic roles include
carbohydrate metabolism, the production
of hormones, conversion and storage of
nutrients such as glucose and glycogen,
and the decomposition of red blood
cells.[4]
Liver
The human liver is located in the upper right

abdomen
Location of human liver (in red) shown on a

male body
Details
Precursor Foregut
System Digestive system

Artery Hepatic artery
Vein Hepatic vein and
hepatic portal vein
Nerve Celiac ganglia and

vagus nerve[1]
Identifiers
Latin jecur, iecur
Greek hepar (ἧπαρ)

root hepat- (ἡπατ-)
MeSH D008099 (https://me

shb.nlm.nih.gov/reco
rd/ui?ui=D008099)
TA98 A05.8.01.001 (http

s://ifaa.unifr.ch/Publi
c/EntryPage/TA98%2
0Tree/Entity%20TA9
8%20EN/05.8.01.00
1%20Entity%20TA9
TA2 8%20EN.htm)
3023 (https://ta2view
er.openanatomy.or
g/?id=3023)
FMA 7197 (https://bioport

al.bioontology.org/on
tologies/FMA/?p=cla
sses&conceptid=htt
p%3A%2F%2Fpurl.or
g%2Fsig%2Font%2Ff
ma%2Ffma7197)
Anatomical terminology
The liver is also an accessory digestive

organ that produces bile, an alkaline fluid
containing cholesterol and bile acids,
which emulsifies and aids the breakdown
of dietary fat. The gallbladder, a small
hollow pouch that sits just under the
right lobe of liver, stores and
concentrates the bile produced by the
liver, which is later excreted to the
duodenum to help with digestion.[5] The
liver's highly specialized tissue,
consisting mostly of hepatocytes,
regulates a wide variety of high-volume
biochemical reactions, including the
synthesis and breakdown of small and
complex organic molecules, many of
which are necessary for normal vital
functions.[6] Estimates regarding the
organ's total number of functions vary,
but is generally cited as being around
500.[7]
It is not known how to compensate for
the absence of liver function in the long
term, although liver dialysis techniques
can be used in the short term. Artificial
livers have not been developed to
promote long-term replacement in the
absence of the liver. As of 2018,[8] liver
transplantation is the only option for
complete liver failure.
Structure
The liver, viewed from above, showing

the left and right lobes separated by
the falciform ligament
The liver is a dark reddish brown, wedge-
shaped organ with two lobes of unequal
size and shape. A human liver normally
weighs approximately 1.5 kilograms (3.3
pounds)[9] and has a width of about 15
centimetres (6 inches).[10] There is
considerable size variation between
individuals, with the standard reference
range for men being 970–1,860 grams
(2.14–4.10 lb)[11] and for women 600–
1,770 g (1.32–3.90 lb).[12] It is both the
heaviest internal organ and the largest
gland in the human body. It is located in
the right upper quadrant of the
abdominal cavity, resting just below the
diaphragm, to the right of the stomach,
and overlying 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 via the celiac
trunk, whereas the portal vein carries
blood rich in digested nutrients from the
entire gastrointestinal tract and also
from the spleen and pancreas.[8] These
blood vessels subdivide into small
capillaries known as liver sinusoids,
which then lead to hepatic lobules.
Hepatic lobules are the functional units

of the liver. Each lobule is made up of
millions of hepatic cells (hepatocytes),
which are the basic metabolic cells. The
lobules are held together by a fine, dense,
irregular, fibroelastic connective tissue
layer extending from the fibrous capsule
covering the entire liver known as
Glisson's capsule after British doctor
Francis Glisson.[4] This tissue extends
into the structure of the liver by
accompanying the blood vessels, ducts,
and nerves at the hepatic hilum. The
whole surface of the liver, except for the
bare area, is covered in a serous coat
derived from the peritoneum, and this
firmly adheres to the inner Glisson's
capsule.
Gross anatomy
Terminology related to the liver often
starts in hepat- from ἡπατο-, from the
Greek word for liver.[13]
Lobes
The liver, viewed from below, surface

showing four lobes and the
impressions
The liver is grossly divided into two parts

when viewed from above – a right and a
left lobe – and four parts when viewed
from below (left, right, caudate, and
quadrate lobes).[14]
The falciform ligament makes a

superficial division of the liver into a left
and right lobe. From below, the two
additional lobes are located between the
right and left lobes, one in front of the
other. A line can be imagined running
from the left of the vena cava and all the
way forward to divide the liver and
gallbladder into two halves.[15] This line is
called Cantlie's line.[16]
Other anatomical landmarks include the

ligamentum venosum and the round
ligament of the liver, which further divide
the left side of the liver in two sections.
An important anatomical landmark, the
porta hepatis, divides this left portion
into four segments, which can be
numbered starting at the caudate lobe as
I in an anticlockwise manner. From this
parietal view, seven segments can be
seen, because the eighth segment is only
visible in the visceral view.[17]
Surfaces
On the diaphragmatic surface, apart from

a triangular bare area where it connects
to the diaphragm, the liver is covered by a
thin, double-layered membrane, the
peritoneum, that helps to reduce friction
against other organs.[18] This surface
covers the convex shape of the two lobes
where it accommodates the shape of the
diaphragm. The peritoneum folds back
on itself to form the falciform ligament
and the right and left triangular
ligaments.[19]
These peritoneal ligaments are not

related to the anatomic ligaments in
joints, and the right and left triangular
ligaments 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]
The fossa of gallbladder lies to the right
of the quadrate lobe, occupied by the
gallbladder with its cystic duct close to
the right end of porta hepatis.
Impressions
Impressions of the liver
Several impressions on the surface of

the liver accommodate the various
adjacent structures and organs.
Underneath the right lobe and to the right
of the gallbladder fossa are two
impressions, one behind the other and
separated by a ridge. The one in front is a
shallow colic impression, formed by the
hepatic flexure and the one behind is a
deeper renal impression accommodating
part of the right kidney and part of the
suprarenal gland.[20]
The suprarenal impression is a small,

triangular, depressed area on the liver. It
is located close to the right of the fossa,
between the bare area and the caudate
lobe, and immediately above the renal
impression. The greater part of the
suprarenal impression is devoid of
peritoneum and it lodges the right
suprarenal gland.[21]
Medial to the renal impression is a third

and slightly marked impression, lying
between it and the neck of the gall
bladder. This is caused by the
descending portion of the duodenum,
and is known as the duodenal
impression.[21]
The inferior surface of the left lobe of the

liver presents behind and to the left of
the gastric impression.[21] This is
moulded over the upper front surface of
the stomach, and to the right of this is a
rounded eminence, the tuber omentale,
which fits into the concavity of the lesser
curvature of the stomach and lies in front
of the anterior layer of the lesser
omentum.
Microscopic anatomy
Cells, ducts, and blood vessels
Microscopically, each liver lobe is seen to

be made up of hepatic lobules. The
lobules are roughly hexagonal, and
consist of plates of hepatocytes, and
sinusoids radiating from a central vein
towards an imaginary perimeter of
interlobular portal triads.[22] The central
vein joins to the hepatic vein to carry
blood out from the liver. A distinctive
component of a lobule is the portal triad,
which can be found running along each
of the lobule's corners. The portal triad
consists of the hepatic artery, the portal
vein, and the common bile duct.[23] The
triad may be seen on a liver ultrasound,
as a Mickey Mouse sign with the portal
vein as the head, and the hepatic artery,
and the common bile duct as the ears.[24]
Histology, the study of microscopic

anatomy, shows two major types of liver
cell: parenchymal cells and
nonparenchymal cells. About 70–85% of
the liver volume is occupied by
parenchymal hepatocytes.
Nonparenchymal cells constitute 40% of
the total number of liver cells but only
6.5% of its volume.[25] The liver sinusoids
are lined with two types of cell,
sinusoidal endothelial cells, and
phagocytic Kupffer cells.[26] Hepatic
stellate cells are nonparenchymal cells
found in the perisinusoidal space,
between a sinusoid and a hepatocyte.[25]
Additionally, intrahepatic lymphocytes
are often present in the sinusoidal
lumen.[25]
Microscopic anatomy of the liver
Types of capillaries–sinusoid on right

3D Medical Animation Still Shot Depicting
parts of liver
Functional anatomy
Hilum of the liver, circled in yellow
The central area or hepatic hilum,

includes the opening known as the porta
hepatis which carries the common bile
duct and common hepatic artery, and the
opening for the portal vein. The duct,
vein, and artery divide into left and right
branches, and the areas of the liver
supplied by these branches constitute
the functional left and right lobes. The
functional lobes are separated by the
imaginary plane, Cantlie's line, joining the
gallbladder fossa to the inferior vena
cava. The plane separates the liver into
the true right and left lobes. The middle
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 left
hepatic vein.
The hilum of the liver is described in
terms of three plates that contain the bile
ducts and blood vessels. The contents of
the whole plate system are surrounded
by a sheath.[27] The three plates are the
hilar plate, the cystic plate and the
umbilical plate and the plate system is
the site of the many anatomical
variations to be found in the liver.[27]
Couinaud classification system
Shape of human liver in animation,

with eight Couinaud segments
labelled
In the widely used Couinaud system, the
functional lobes are further divided into a
total of eight subsegments based on a
transverse plane through the bifurcation
of the main portal vein.[28] The caudate
lobe is a separate structure that receives
blood flow from both the right- and left-
sided vascular branches.[29][30] The
Couinaud classification divides the liver
into eight functionally independent liver
segments. Each segment has its own
vascular inflow, outflow and biliary
drainage. In the centre of each segment
are branches of the portal vein, hepatic
artery, and bile duct. In the periphery of
each segment is vascular outflow
through the hepatic veins.[31] The
classification system uses the vascular
supply in the liver to separate the
functional units (numbered I to VIII) with
unit 1, the caudate lobe, receiving its
supply from both the right and the left
branches of the portal vein. It contains
one or more hepatic veins which drain
directly into the inferior vena cava.[28] The
remainder of the units (II to VIII) are
numbered in a clockwise fashion:[31]
Gene and protein expression
About 20,000 protein coding genes are

expressed in human cells and 60% of
these genes are expressed in a normal,
adult liver.[32][33] Over 400 genes are more
specifically expressed in the liver, with
some 150 genes highly specific for liver
tissue. A large fraction of the
corresponding liver-specific proteins are
mainly expressed in hepatocytes and
secreted into the blood and constitute
plasma proteins and hepatokines. Other
liver-specific proteins are certain liver
enzymes such as HAO1 and RDH16,
proteins involved in bile synthesis such
as BAAT and SLC27A5, and transporter
proteins involved in the metabolism of
drugs, such as ABCB11 and SLC2A2.
Examples of highly liver-specific proteins
include apolipoprotein A II, coagulation
factors F2 and F9, complement factor
related proteins, and the fibrinogen beta
chain protein.[34]
Development
CT scan showing an adult liver in the

axial plane
Organogenesis, the development of the

organs, takes place from the third to the
eighth week during embryogenesis. The
origins of the liver lie in both the ventral
portion of the foregut endoderm
(endoderm being one of the three
embryonic germ layers) and the
constituents of the adjacent septum
transversum mesenchyme. In the human
embryo, the hepatic diverticulum is the
tube of endoderm that extends out from
the foregut into the surrounding
mesenchyme. The mesenchyme of
septum transversum induces this
endoderm to proliferate, to branch, and to
form the glandular epithelium of the liver.
A portion of the hepatic diverticulum
(that region closest to the digestive tube)
continues to function as the drainage
duct of the liver, and a branch from this
duct produces the gallbladder.[35]
Besides signals from the septum
transversum mesenchyme, fibroblast
growth factor from the developing heart
also contributes to hepatic competence,
along with retinoic acid emanating from
the lateral plate mesoderm. The hepatic
endodermal cells undergo a
morphological transition from columnar
to pseudostratified resulting in thickening
into the early liver bud. Their expansion
forms a population of the bipotential
hepatoblasts.[36] Hepatic stellate cells
are derived from mesenchyme.[37]
After migration of hepatoblasts into the

septum transversum mesenchyme, the
hepatic architecture begins to be
established, with liver sinusoids and bile
canaliculi appearing. The liver bud
separates into the lobes. The left
umbilical vein becomes the ductus
venosus and the right vitelline vein
becomes the portal vein. The expanding
liver bud is colonized by hematopoietic
cells. The bipotential hepatoblasts begin
differentiating into biliary epithelial cells
and hepatocytes. The biliary epithelial
cells differentiate from hepatoblasts
around portal veins, first producing a
monolayer, and then a bilayer of cuboidal
cells. In ductal plate, focal dilations
emerge at points in the bilayer, become
surrounded by portal mesenchyme, and
undergo tubulogenesis into intrahepatic
bile ducts. Hepatoblasts not adjacent to
portal veins instead differentiate into
hepatocytes and arrange into cords lined
by sinusoidal epithelial cells and bile
canaliculi. Once hepatoblasts are
specified into hepatocytes and undergo
further expansion, they begin acquiring
the functions of a mature hepatocyte,
and eventually mature hepatocytes
appear as highly polarized epithelial cells
with abundant glycogen accumulation. In
the adult liver, hepatocytes are not
equivalent, with position along the
portocentrovenular axis within a liver
lobule dictating expression of metabolic
genes involved in drug metabolism,
carbohydrate metabolism, ammonia
detoxification, and bile production and
secretion. WNT/β-catenin has now been
identified to be playing a key role in this
phenomenon.[36]
Adult ultrasound showing the right

lobe of the liver and right kidney
At birth, the liver comprises roughly 4% of

body weight and weighs on average
about 120 g (4 oz). Over the course of
further development, it will increase to
1.4–1.6 kg (3.1–3.5 lb) but will only take
up 2.5–3.5% of body weight.[38]
Hepatosomatic index (HSI) is the ratio of

liver weight to body weight.[39]
Fetal blood supply

pp y
In the growing fetus, a major source of

blood to the liver is the umbilical vein,
which supplies nutrients to the growing
fetus. The umbilical vein enters the
abdomen at the umbilicus and passes
upward along the free margin of the
falciform ligament of the liver to the
inferior surface of the liver. There, it joins
with the left branch of the portal vein.
The ductus venosus carries blood from
the left portal vein to the left hepatic vein
and then to the inferior vena cava,
allowing placental blood to bypass the
liver. In the fetus, the liver does not
perform the normal digestive processes
and filtration of the infant liver because
nutrients are received directly from the
mother via the 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 cells shifts to
the red bone marrow. After 2–5 days, the
umbilical vein and ductus venosus are
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.
Functions
The various functions of the liver are
carried out by the liver cells or
hepatocytes. The liver is thought to be
responsible for up to 500 separate
functions, usually in combination with
other systems and organs. Currently, no
artificial organ or device is capable of
reproducing all the functions of the liver.
Some functions can be carried out by
liver dialysis, an experimental treatment
for liver failure. The liver also accounts
for about 20% of resting total body
oxygen consumption.
Blood supply
The liver receives a dual blood supply
from the hepatic portal vein and hepatic
arteries. The hepatic portal vein delivers
around 75% of the liver's blood supply
and carries venous blood drained from
the spleen, gastrointestinal tract, and its
associated organs. The hepatic arteries
supply arterial blood to the liver,
accounting for the remaining quarter of
its blood flow. Oxygen is provided from
both sources; about half of the liver's
oxygen demand is met by the hepatic
portal vein, and half is met by the hepatic
arteries.[40] The hepatic artery also has
both alpha- and beta-adrenergic
receptors; therefore, flow through the
artery is controlled, in part, by the
splanchnic nerves of the autonomic
nervous system.
Blood flows through the liver sinusoids

and empties into the central vein of each
lobule. The central veins coalesce into
hepatic veins, which leave the liver and
drain into the inferior vena cava.[41]
The liver and its veins

Diagram of liver, lobule, and portal tract and
their inter-relations
Biliary flow
Biliary tract
The biliary tract is derived from the

branches of the bile ducts. The biliary
tract, also known as the biliary tree, is the
path by which bile is secreted by the liver
then transported to the first part of the
small intestine, the duodenum. The bile
produced in the liver is collected in bile
canaliculi, small grooves between the
faces of adjacent hepatocytes. The
canaliculi radiate to the edge of the liver
lobule, where they merge to form bile
ducts. Within the liver, these ducts are
termed intrahepatic bile ducts, and once
they exit the liver, they are considered
extrahepatic. The intrahepatic ducts
eventually drain into the right and left
hepatic ducts, which exit the liver at the
transverse fissure, and merge to form the
common hepatic duct. The cystic duct
from the gallbladder joins with the
common hepatic duct to form the
common bile duct.[41] The biliary system
and connective tissue is supplied by the
hepatic artery alone.
Bile either drains directly into the

duodenum via the common bile duct, or
is temporarily stored in the gallbladder
via the cystic duct. The common bile
duct and the pancreatic duct enter the
second part of the duodenum together at
the hepatopancreatic ampulla, also
known as the ampulla of Vater.
Metabolism
The liver plays a major role in

carbohydrate, protein, amino acid, and
lipid metabolism.
Carbohydrate metabolism
The liver performs several roles in

carbohydrate metabolism.
The liver synthesizes and stores

around 100g of glycogen via
glycogenesis, the formation of
glycogen from glucose.
When needed, the liver releases
glucose into the blood by performing
glycogenolysis, the breakdown of
glycogen into glucose.[42]
The liver is also responsible for
gluconeogenesis, which is the
synthesis of glucose from certain
amino acids, lactate, or glycerol.
Adipose and liver cells produce
glycerol by breakdown of fat, which the
liver uses for gluconeogenesis.[42]
Liver also does glyconeogenesis
which is synthesis of glycogen from
lactic acid.[43]
Protein metabolism
The liver is responsible for the mainstay

of protein metabolism, synthesis as well
as degradation. All plasma proteins
except Gamma-globulins are synthesised
in the liver.[44] It is also responsible for a
large part of amino acid synthesis. The
liver plays a role in the production of
clotting factors, as well as red blood cell
production. Some of the proteins
synthesized by the liver include
coagulation factors I (fibrinogen), II
(prothrombin), V, VII, VIII, IX, X, XI, XII, XIII,
as well as protein C, protein S and
antithrombin. The liver is a major site of
production for thrombopoietin, a
glycoprotein hormone that regulates the
production of platelets by the bone
marrow.[45]
Lipid metabolism
The liver plays several roles in lipid

metabolism: it performs cholesterol
synthesis, lipogenesis, and the
production of triglycerides, and a bulk of
the body's lipoproteins are synthesized in
the liver. The liver plays a key role in
digestion, as it produces and excretes
bile (a yellowish liquid) required for
emulsifying fats and help the absorption
of vitamin K from the diet. Some of the
bile drains directly into the duodenum,
and some is stored in the gallbladder.
The liver produces insulin-like growth
factor 1, a polypeptide protein hormone
that plays an important role in childhood
growth and continues to have anabolic
effects in adults.
Breakdown
The liver is responsible for the
breakdown of insulin and other
hormones. The liver breaks down
bilirubin via glucuronidation, facilitating
its excretion into bile. The 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. Preferably, the
toxins are conjugated to avail excretion in
bile or urine. The liver converts ammonia
into urea as part of the ornithine cycle or
the urea cycle, and the urea is excreted in
the urine.[46]
Blood reservoir
Because the liver is an expandable organ,

large quantities of blood can be stored in
its blood vessels. Its normal blood
volume, including both that in the hepatic
veins and that in the hepatic sinuses, is
about 450 milliliters, or almost 10
percent of the body's total blood volume.
When high pressure in the right atrium
causes backpressure in the liver, the liver
expands, and 0.5 to 1 liter of extra blood
is occasionally stored in the hepatic
veins and sinuses. This occurs especially
in cardiac failure with peripheral
congestion. Thus, in effect, the liver is a
large, expandable, venous organ capable
of acting as a valuable blood reservoir in
times of excess blood volume and
capable of supplying extra blood in times
of diminished blood volume.[47]
Lymph production
Because the pores in the hepatic

sinusoids are very permeable and allow
ready passage of both fluid and proteins
into the perisinusoidal space, the lymph
draining from the liver usually has a
protein concentration of about 6 g/dl,
which is only slightly less than the
protein concentration of plasma. Also,
the high permeability of the liver sinusoid
epithelium allows large quantities of
lymph to form. Therefore, about half of
all the lymph formed in the body under
resting conditions arises in the liver.
Other
The liver stores a multitude of

substances, including vitamin A (1–2
years' supply), vitamin D (1–4 months'
supply),[48] vitamin B12 (3–5 years'
supply),[49] vitamin K, vitamin E, iron,
copper, zinc, cobalt, molybdenum, etc.
Haemopoiesis - The formation of
blood cells is called haemopoiesis. In
embryonic stage RBC and WBC are
formed by liver. In the first trimester
fetus, the liver is the main site of red
blood cell production. By the 32nd
week of gestation, the bone marrow
has almost completely taken over that
task.[50]
The liver helps in the purification of
blood. The Kupffer cells of liver are
phagocytic cells, helps in phagocytosis
of dead blood cells and bacteria from
the blood.[51]
The liver is responsible for
immunological effects – the
mononuclear phagocyte system of the
liver contains many immunologically
active cells, acting as a 'sieve' for
antigens carried to it via the portal
system.
The liver produces albumin, the most
abundant protein in blood serum. It is
essential in the maintenance of
oncotic pressure, and acts as a
transport for fatty acids and steroid
hormones.
The liver synthesizes angiotensinogen,
a hormone that is responsible for
raising the blood pressure when
activated by renin, an enzyme that is
released when the kidney senses low
blood pressure.
The liver produces the enzyme
catalase to break down hydrogen
peroxide, a toxic oxidising agent, into
water and oxygen.
With aging
The oxidative capacity of the liver

decreases with aging, and therefore any
medications that require oxidation (for
instance, benzodiazepines) are more
likely to accumulate to toxic levels.
However, medications with shorter half-
lives, such as lorazepam and oxazepam,
are preferred in most cases when
benzodiazepines are required in regard to
geriatric medicine.
Clinical significance
Disease
Left lobe liver tumor
The liver is a vital organ and supports

almost every other organ in the body.
Because of its strategic location and
multidimensional functions, the liver is
prone to many diseases.[52] The bare
area of the liver is a site that is vulnerable
to the passing of infection from the
abdominal cavity to the thoracic cavity.
Liver diseases may be diagnosed by liver
function tests–blood tests that can
identify various markers. For example,
acute-phase reactants are produced by
the liver in response to injury or
inflammation.
The most common chronic liver disease

is nonalcoholic fatty liver disease, which
affects an estimated one-third of the
world population.[53]
Hepatitis is a common condition of

inflammation of the 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.
Inflammation can also be caused by
other viruses in the family Herpesviridae
such as the herpes simplex virus.
Chronic (rather than acute) infection with
hepatitis B virus or hepatitis C virus is the
main cause of liver cancer.[54] Globally,
about 248 million individuals are
chronically infected with hepatitis B (with
843,724 in the U.S.),[55] and 142 million
are chronically infected with hepatitis
C[56] (with 2.7 million in the U.S.).[57]
Globally there are about 114 million and
20 million cases of hepatitis A[56] and
hepatitis E[58] respectively, but these
generally resolve and do not become
chronic. Hepatitis D virus is a "satellite"
of hepatitis B virus (it can only infect in
the presence of hepatitis B), and co-
infects nearly 20 million people with
hepatitis B, globally.[59]
Hepatic encephalopathy is caused by an

accumulation of toxins in the
bloodstream that are normally removed
by the liver. This condition can result in
coma and can prove fatal. Budd–Chiari
syndrome is a condition caused by
blockage of the hepatic veins (including
thrombosis) that drain the liver. It
presents with the classical triad of
abdominal pain, ascites and liver
enlargement.[60] 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.
Other disorders caused by excessive

alcohol consumption are grouped under
alcoholic liver diseases and these
include alcoholic hepatitis, fatty liver, and
cirrhosis. Factors contributing to the
development of alcoholic liver diseases
are not only the quantity and frequency
of alcohol consumption, but can also
include gender, genetics, and liver insult.
Liver damage can also be caused by
drugs, particularly paracetamol and
drugs used to treat cancer. A rupture of
the liver can be caused by a liver shot
used in combat sports.
Primary biliary cholangitis is an

autoimmune disease of the liver.[61][62] It
is marked by slow progressive
destruction of the small bile ducts of the
liver, with the intralobular ducts (Canals
of Hering) affected early in the
disease.[63] When these ducts are
damaged, bile and other toxins build up
in the liver (cholestasis) and over time
damages the liver tissue in combination
with ongoing immune related damage.
This can lead to scarring (fibrosis) and
cirrhosis. Cirrhosis increases the
resistance to blood flow in the liver, and
can result in portal hypertension.
Congested anastomoses between the
portal venous system and the systemic
circulation, can be a subsequent
condition.
There are also many pediatric liver

diseases, including biliary atresia, alpha-1
antitrypsin deficiency, 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. A
genetic disorder causing multiple cysts
to form in the liver tissue, usually in later
life, and usually asymptomatic, is
polycystic liver disease. 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
produces symptoms after extensive
damage.
Hepatomegaly refers to an enlarged liver

and can be due to many causes. It can be
palpated in a liver span measurement.
Symptoms
The classic symptoms of liver damage

include the following:
Pale stools occur when stercobilin, a
brown pigment, is absent from the
stool. Stercobilin is derived from
bilirubin metabolites produced in the
liver.
Dark urine occurs when bilirubin mixes
with urine
Jaundice (yellow skin and/or whites of
the eyes) This is where bilirubin
deposits in skin, causing an intense
itch. Itching is the most common
complaint by people who have liver
failure. Often this itch cannot be
relieved by drugs.
Swelling of the abdomen, and swelling
of the ankles and feet occurs because
the liver fails to make albumin.
Excessive fatigue occurs from a
generalized loss of nutrients, minerals
and vitamins.
Bruising and easy bleeding are other
features of liver disease. The liver
makes clotting factors, substances
which help prevent bleeding. When
liver damage occurs, these factors are
no longer present and severe bleeding
can occur.[64]
Pain in the upper right quadrant can
result from the stretching of Glisson's
capsule in conditions of hepatitis and
pre-eclampsia.
Diagnosis
The diagnosis of liver disease is made by
liver function tests, groups of blood tests,
that can readily show the extent of liver
damage. If infection is suspected, then
other serological tests will be carried out.
A physical examination of the liver can
only reveal its size and any tenderness,
and some form of imaging such as an
ultrasound or CT scan may also be
needed.
Sometimes a liver biopsy will be

necessary, and a tissue sample is taken
through a needle inserted into the skin
just below the rib cage. This procedure
may be helped by a sonographer
providing ultrasound guidance to an
interventional radiologist.[65]
Axial CT image showing anomalous

hepatic veins coursing on the liver's
subcapsular anterior surface[66]
Maximum intensity projection (MIP) CT
image as viewed anteriorly showing the
anomalous hepatic veins coursing on the
anterior surface of the liver
Lateral MIP view in the same patient as

previous image
A CT scan in which the liver and portal vein
are shown
Liver regeneration
The liver is the only human internal organ

capable of natural regeneration of lost
tissue; as little as 25% of a liver can
regenerate into a whole liver.[67] This is,
however, not true regeneration but rather
compensatory growth in mammals.[68]
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 regeneration
where both original function and form are
restored. In some other species, such as
zebrafish, the liver undergoes true
regeneration by restoring both shape and
size of the organ.[69] In the liver, large
areas of the tissues are formed but for
the formation of new cells there must be
sufficient amount of material so the
circulation of the blood becomes more
active.[70]
This is predominantly due to the

hepatocytes re-entering the cell cycle.
That is, the hepatocytes go from the
quiescent G0 phase to the G1 phase and
undergo mitosis. This process is
activated by the p75 receptors.[71] 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 in the canals of
Hering. These cells can differentiate into
either hepatocytes or cholangiocytes.
Cholangiocytes are the epithelial lining
cells of the bile ducts.[72] 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. Research is
being carried out on the use of stem cells
for the generation of an artificial liver.
Scientific and medical works about liver

regeneration often refer to the Greek
Titan Prometheus who was chained to a
rock in the Caucasus where, each day, his
liver was devoured by an eagle, only to
grow back each night. The myth
suggests the ancient Greeks may have
known about the liver's remarkable
capacity for self-repair.[73]
Liver transplantation
Human liver transplants were first

performed by Thomas Starzl in the
United States and Roy Calne in
Cambridge, England in 1963 and 1967,
respectively.
After resection of left lobe liver tumor
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, and
autoimmune hepatitis. Less commonly,
liver transplantation is done for fulminant
hepatic failure, in which liver failure
occurs rapidly over a period of days or
weeks.
Liver allografts for transplant usually
come from donors who have died from
fatal brain injury. Living donor liver
transplantation is a technique in which a
portion of a living person's liver is
removed (hepatectomy) and used to
replace the entire liver of the recipient.
This was first performed in 1989 for
pediatric liver transplantation. Only 20
percent of an adult's 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
donor's right hepatic lobe, which
amounts to 60 percent of the liver. Due to
the ability of the liver to regenerate, both
the donor and recipient end up with
normal liver function if all goes well. This
procedure is more controversial, as it
entails performing a much larger
operation on the donor, and indeed there
were at least two donor deaths out of the
first several hundred cases. A 2006
publication addressed the problem of
donor mortality and found at least
fourteen cases.[74] The risk of
postoperative complications (and death)
is far greater in right-sided operations
than that in left-sided operations.
With the recent advances of noninvasive
imaging, living liver donors usually have
to undergo imaging examinations for
liver anatomy to decide if the anatomy is
feasible for donation. The evaluation is
usually performed by multidetector row
computed tomography (MDCT) and
magnetic resonance imaging (MRI).
MDCT is good in vascular anatomy and
volumetry. MRI is used for biliary tree
anatomy. Donors with very unusual
vascular anatomy, which makes them
unsuitable for donation, could be
screened out to avoid unnecessary
operations.
MDCT image. Arterial anatomy
contraindicated for liver donation
MDCT image. Portal venous anatomy

contraindicated for liver donation
MDCT image. 3D image created by MDCT
can clearly visualize the liver, measure the
liver volume, and plan the dissection plane
to facilitate the liver transplantation
procedure.
Phase contrast CT image. Contrast is
perfusing the right liver but not the left due
to a left portal vein thrombus.
Society and culture
Some cultures regard the liver as the

seat of the soul.[75] In Greek mythology,
the gods punished Prometheus for
revealing fire to humans by chaining him
to a rock where a vulture (or an eagle)
would peck out his liver, which would
regenerate overnight (the liver is the only
human internal organ that actually can
regenerate itself to a significant extent).
Many ancient peoples of the Near East
and Mediterranean areas practiced a
type of divination called haruspicy or
hepatomancy, where they tried to obtain
information by examining the livers of
sheep and other animals.
In Plato, and in later physiology, the liver

was thought to be the seat of the darkest
emotions (specifically wrath, jealousy
and greed) which drive men to action.[76]
The Talmud (tractate Berakhot 61b)
refers to the liver as the seat of anger,
with the gallbladder counteracting this.
The Persian, Urdu, and Hindi languages
(‫ ﺟﮕﺮ‬or जगर or jigar) refer to the liver in
figurative speech to indicate courage and
strong feelings, or "their best"; e.g., "This
Mecca has thrown to you the pieces of
its liver!".[77] 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 desirable, especially
women. In the Zulu language, the word
for liver (isibindi) is the same as the word
for courage. In English the term 'lily-
livered' is used to indicate cowardice
from the medieval belief that the liver
was the seat of courage. Spanish
hígados also means "courage".[78]
However the secondary meaning of
Basque gibel is "indolence".[79]
In biblical Hebrew, the word for liver, ‫כבד‬

(Kauved, stemmed KBD or KVD, similar to
Arabic ‫)اﻟﻜﺒﺪ‬, also means heavy and is
used to describe the rich ("heavy" with
possessions) and honor (presumably for
the same reason). In the Book of
Lamentations (2:11) it is used to
describe the physiological responses to
sadness by "my liver spilled to earth"
along with the flow of tears and the
overturning in bitterness of the
intestines.[80] On several occasions in the
book of Psalms (most notably 16:9), the
word is used to describe happiness in the
liver, along with the heart (which beats
rapidly) and the flesh (which appears red
under the skin). Further usage as the self
(similar to "your honor") is widely
available throughout the old testament,
sometimes compared to the breathing
soul (Genesis 49:6, Psalms 7:6, etc.). An
honorable hat was also referred to with
this word (Job 19:9, etc.) and under that
definition appears many times along with
‫ פאר‬Pe'er - grandeur.[81]
These four meanings were used in

preceding ancient Afro-Asiatic languages
such as Akkadian and Ancient Egyptian
preserved in classical Ethiopic Ge'ez
language.[82]
Food
Maksalaatikko, a Finnish liver

casserole
Humans commonly eat the livers of

mammals, fowl, and fish as food.
Domestic pig, ox, lamb, calf, chicken, and
goose livers are widely available from
butchers and supermarkets. In the
Romance languages, the anatomical
word for "liver" (French foie, Spanish
hígado, etc.) derives not from the Latin
anatomical term, jecur, but from the
culinary term ficatum, literally "stuffed
with figs," referring to the livers of geese
that had been fattened on figs.[83] Animal
livers are rich in iron, vitamin A and
vitamin B12; and cod liver oil is commonly
used as a dietary supplement.
Liver can be baked, boiled, broiled, fried,

stir-fried, or eaten raw (asbeh nayeh or
sawda naye in Lebanese cuisine, or liver
sashimi in Japanese cuisine). In many
preparations, pieces of liver are
combined with pieces of meat or
kidneys, as in the various forms of
Middle Eastern mixed grill (e.g. meurav
Yerushalmi). Well-known examples
include liver pâté, foie gras, chopped
liver, and leverpastej. Liver sausages,
such as Braunschweiger and liverwurst,
are also a valued meal. Liver sausages
may also be used as spreads. A
traditional South African delicacy,
skilpadjies, is made of minced lamb's
liver wrapped in netvet (caul fat), and
grilled over an open fire. Traditionally,
some fish livers were valued as food,
especially the stingray liver. It was used
to prepare delicacies, such as poached
skate liver on toast in England, as well as
the beignets de foie de raie and foie de
raie en croute in French cuisine.[84]
Giraffe liver
19th-century drinking scene in
Kordofan, home to the Humr tribe,
who made a drink from giraffe liver.
Plate from Le Désert et le Soudan by
Stanislas d'Escayrac de Lauture.
The Humr are one of the tribes in the

Baggara ethnic group, native to
southwestern Kordofan in Sudan who
speak Shuwa (Chadian Arabic), make a
non-alcoholic drink from the liver and
bone marrow of the giraffe, which they
call umm nyolokh. They claim it is
intoxicating (Arabic ‫ ﺳﻜﺮان‬sakran),
causing dreams and even waking
hallucinations.[85] Anthropologist Ian
Cunnison accompanied the Humr on one
of their giraffe-hunting expeditions in the
late 1950s, and noted that:
It is said that a person, once he has

drunk umm nyolokh, will return to
giraffe again and again. Humr, being
Mahdists, are strict abstainers [from
alcohol] and a Humrawi is never drunk
(sakran) on liquor or beer. But he uses
this word to describe the effects which
umm nyolokh has upon him.[86]
Cunnison's remarkable account of an

apparently psychoactive mammal found
its way from a somewhat obscure
scientific paper into more mainstream
literature through a conversation
between W. James of the Institute of
Social and Cultural Anthropology at the
University of Oxford and specialist on the
use of hallucinogens and intoxicants in
society, and R. Rudgley, who discussed it
in a book on psychoactive drugs for
general readers.[85] He speculated that a
hallucinogenic compound N,N-
Dimethyltryptamine in the giraffe liver
might account for the intoxicating
properties claimed for umm nyolokh.[85]
Cunnison, on the other hand, writing in

1958 found it hard to believe in the literal
truth of the Humr's assertion that the
drink was intoxicating:
I can only assume that there is no

intoxicating substance in the drink, and
that the effect it produces is simply a
matter of convention, although it may
be brought about subconsciously.[86]
The study of entheogens in general –

including entheogens of animal origin
(e.g. hallucinogenic fish and toad venom)
– has, however, made considerable
progress in the sixty-odd years since
Cunnison's report; the idea that some
intoxicating substance might reside in
giraffe livers may no longer be as far-
fetched as it seemed to Cunnison.
However, to date, proof (or disproof) still
waits on detailed analyses of the organ
and the beverage made from it.[85]
Arrow/bullet poison
Arrow/bullet poison
Certain Tungusic peoples of northeast

Asia formerly prepared a type of arrow
poison from rotting animal livers, which
was, in later times, also applied to
bullets. Russian anthropologist S. M.
Shirokogoroff wrote that:
Formerly the using of poisoned arrows

was common. For instance, among the
Kumarčen, [a subgroup of the Oroqen]
even in recent times, a poison was
used which was prepared from
decaying liver.
[Note] This has been confirmed by the

Kumarčen. I am not competent to
judge as to the chemical conditions of
production of poison which is not
destroyed by the heat of explosion.
However, the Tungus themselves
compare this method [of poisoning
ammunition] with the poisoning of
arrows.[87]
Other animals
Sheep's liver
The liver is found in all vertebrates and is

typically the largest internal organ. The
internal structure of the liver is broadly
similar in all vertebrates, though its form
varies considerably in different species,
and is largely determined by the shape
and arrangement of the surrounding
organs. Nonetheless, in most species it
is divided into right and left lobes;
exceptions to this general rule include
snakes, where the shape of the body
necessitates a simple cigar-like form.[88]
An organ sometimes referred to as a liver

is found associated with the digestive
tract of the primitive chordate
Amphioxus. Although it performs many
functions of a liver, it is not considered a
"true" liver but rather a homolog of the
vertebrate liver.[89][90][91] The amphioxus
hepatic caecum produces the liver-
specific proteins vitellogenin,
antithrombin, plasminogen, alanine
aminotransferase, and insulin/Insulin-like
growth factor.[92]
See also
Porphyria
Johann Joseph Dömling (published Is
the liver a purifying organ in 1798)
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g/10.1016%2Fj.jhep.2010.02.032) .
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p. 281. ISBN 9781616404642. Retrieved
2018-09-16. "Now among people in a
primitive state of culture the soul is
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7 . Krishna, Gopi; Hillman, James (1970).
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0. ‫כלו בדמעות עיני חמרמרו מעי נשפך לארץ‬
‫כבדי על שבר בת עמי בעטף עולל ויונק‬
‫" ברחבות קריה‬My eyes terminated with
tears, my intestines overturned with
bitterness, my liver spilled to the earth on
the breaking of my nation's daughter
when a suckling baby lies in the town
squares" (Lamentations 2:11) this could
be interpreted to be read as my honor
spilled, or myself being spilled).
1. Kavod - Honor (https://www.ruvik.co.il/%D
7%A1%D7%99%D7%A4%D7%95%D7%A8%
D7%99-%D7%9E%D7%99%D7%9C%D7%9
9%D7%9D/%D7%9B/%D7%9B%D7%91%D
7%95%D7%93.aspx) (in Hebrew, Israeli
Linguist Ruvik Rosenthal's website).
Rosenthal hypothesized that the term's
usage to describe heaviness comes
perhaps from the liver being the heaviest
of all body parts in some farm animals or
in humans.
2. See Kabadu (https://www.assyrianlangua
ges.org/akkadian/dosearch.php?searchke
y=kab%C4%81tu&language=rawakkadia
n) in Akkadian, (From online dictionary at
Association Assyrophile de France
organization website)
3. "Foie" (https://larousse.fr/dictionnaires/fr
ancais/foie/34366?q=foie#34314) .
Larousse.fr. Archived (https://web.archiv
e.org/web/20180612002718/http://www.l
arousse.fr/dictionnaires/francais/foie/34
366?q=foie#34314) from the original on
2018-06-12. Retrieved 2019-04-16.
4. Schwabe, Calvin W. (1979).
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2015-06-27.
5. Rudgley, R. (1998). The Encyclopedia of
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. Romer, Alfred Sherwood; Parsons,
Thomas S. (1977). The Vertebrate Body.
Philadelphia, PA: Holt-Saunders
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9. Yuan, Shaochun; Ruan, Jie; Huang,
Shengfeng; Chen, Shangwu; Xu, Anlong
(2015). "Amphioxus as a model for
investigating evolution of the vertebrate
immune system" (https://web.archive.org/
web/20151222131815/http://mosas.sys
u.edu.cn/lab/refsys/uppdf/20146202151
1673499.pdf) (PDF). Developmental &
Comparative Immunology. 48 (2): 297–
305. doi:10.1016/j.dci.2014.05.004 (http
s://doi.org/10.1016%2Fj.dci.2014.05.00
i.nlm.nih.gov/24877655) . Archived from
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90. Yu, Jr-Kai Sky; Lecroisey, Claire; Le
Pétillon, Yann; Escriva, Hector; Lammert,
Eckhard; Laudet, Vincent (2015).
"Identification, evolution and expression
of an insulin-like peptide in the
cephalochordate Branchiostoma
lanceolatum" (https://www.ncbi.nlm.nih.g
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ONE. 10 (3): e0119461.
Bibcode:2015PLoSO..1019461L (https://u
i.adsabs.harvard.edu/abs/2015PLoSO..10
19461L) .
doi:10.1371/journal.pone.0119461 (http
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. PMID 25774519 (https://pubmed.ncbi.nl
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91. Escriva, Hector; Chao, Yeqing; Fan,
Chunxin; Liang, Yujun; Gao, Bei; Zhang,
Shicui (2012). "A novel serpin with
antithrombin-like activity in
Branchiostoma japonicum: Implications
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PLOS ONE. 7 (3): e32392.
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92. Guo, Bin; Zhang, Shicui; Wang, Shaohui;
Liang, Yujun (2009). "Expression,
mitogenic activity and regulation by
growth hormone of growth hormone /
insulin-like growth factor in
Branchiostoma belcheri" (https://www.res
earchgate.net/publication/26719830) .
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Works cited
Dorland's illustrated medical dictionary

(32nd ed.). Philadelphia, PA: Elsevier /
Saunders. 2012. ISBN 978-1-4557-
0985-4.
Young, Barbara; O'Dowd, Geraldine;
Woodford, Phillip (4 November 2013).
Wheater's Functional Histology: A text
and colour atlas (6th ed.). Philadelphia,
PA: Elsevier. ISBN 9780702047473.
External links
Look up liver in Wiktionary, the free

dictionary.
Wikimedia Commons has media
related to Livers.
Liver at the Human Protein Atlas (http
s://www.proteinatlas.org/humanproteo
me/liver)
VIRTUAL Liver – online learning
resource (http://pie.med.utoronto.ca/V
Liver/)
Liver enzymes (http://liverenzymes.ne
t)
"Liver" (https://en.wikisource.org/wiki/
1911_Encyclop%C3%A6dia_Britannica/
Liver) . Encyclopædia Britannica.
Vol. 16 (11th ed.). 1911. pp. 801–803.
with several diagrams.
Beaver tail liver
Retrieved from
"https://en.wikipedia.org/w/index.php?
title=Liver&oldid=1190037488"
This page was last edited on 15 December 2023,
at 15:41 (UTC). •
Content is available under CC BY-SA 4.0 unless
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The liver is a major metabolic organ only

The human liver is located in the upper right

Location of human liver (in red) shown on a

System Digestive system

Nerve Celiac ganglia and

Latin jecur, iecur

Greek hepar (ἧπαρ)

MeSH D008099 (https://me

TA98 A05.8.01.001 (http

FMA 7197 (https://bioport

The liver is also an accessory digestive

The liver, viewed from above, showing

Hepatic lobules are the functional units

The liver, viewed from below, surface

The liver is grossly divided into two parts

The falciform ligament makes a

Other anatomical landmarks include the

On the diaphragmatic surface, apart from

These peritoneal ligaments are not

The visceral surface or inferior surface is

Impressions of the liver

Several impressions on the surface of

The suprarenal impression is a small,

Medial to the renal impression is a third

The inferior surface of the left lobe of the

Microscopically, each liver lobe is seen to

Histology, the study of microscopic

Types of capillaries–sinusoid on right

Hilum of the liver, circled in yellow

The central area or hepatic hilum,

Couinaud classiﬁcation system

Shape of human liver in animation,

Gene and protein expression

About 20,000 protein coding genes are

CT scan showing an adult liver in the

Organogenesis, the development of the

After migration of hepatoblasts into the

Adult ultrasound showing the right

At birth, the liver comprises roughly 4% of

Hepatosomatic index (HSI) is the ratio of

Fetal blood supply

In the growing fetus, a major source of

Blood ﬂows through the liver sinusoids

The liver and its veins

The biliary tract is derived from the

Bile either drains directly into the

The liver plays a major role in

The liver performs several roles in

The liver synthesizes and stores

The liver is responsible for the mainstay

The liver plays several roles in lipid

Because the liver is an expandable organ,

Because the pores in the hepatic

The liver stores a multitude of

The oxidative capacity of the liver

Left lobe liver tumor

The liver is a vital organ and supports

The most common chronic liver disease

Hepatitis is a common condition of