Human liver

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This article is about the Human liver. For Liver in general, see Liver.
Liver
Surface projections of the organs of the trunk.png
Surface projections of the organs of the trunk, showing liver in center
Latin Jecur, iecur
Greek Hepar (ἧπαρ)
root hepat- (ἡπατ-)
Gray's p.1188
System Digestive system
Artery hepatic artery
Vein Hepatic vein and hepatic portal vein
Nerve Celiac ganglia and vagus nerve[1]
Precursor Foregut
MeSH Liver
Anatomical terminology

The liver is a vital organ normally present in humans, in the upper right quadrant of the abdomen. The liver has a wide range of functions, including detoxification of various metabolites, protein synthesis, and the production of biochemicals necessary for digestion. The liver is necessary for survival, and there is currently no way to compensate for the absence of liver function in the long term, although new liver dialysis techniques can be used in the short term.

The liver is a gland and plays a major role in metabolism with numerous functions in the human body, including regulation of glycogen storage, decomposition of red blood cells, plasma protein synthesis, hormone production, and detoxification. The liver lies below the diaphragm in the abdominal-pelvic region of the abdomen. It produces bile, an alkaline compound which aids in digestion via the emulsification of lipids. The liver's highly specialized tissue consisting of mostly hepatocytes regulates a wide variety of high-volume biochemical reactions, including the synthesis and breakdown of small and complex molecules, many of which are necessary for normal vital functions.[2]

Terminology related to the liver often starts in hepar- or hepat- from the Greek word for liver, hēpar (ἧπαρ, root hepat-, ἡπατ-).[3][4]

Structure[edit]

The body of the liver is a reddish brown triangular organ with four lobes of unequal size and shape. A human liver normally weighs 1.44–1.66 kg (3.2–3.7 lb).[5]It is both the largest internal organ (the skin being the largest organ overall) 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 overlying the gallbladder. It is connected to two large blood vessels, the hepatic artery and the portal vein. The hepatic artery carries blood from the aorta, whereas the portal vein carries blood containing digested nutrients from the entire gastrointestinal tract and also from the spleen and pancreas. These blood vessels subdivide into capillaries, which then lead to a lobule. Each lobule is made up of millions of hepatic cells (hepatocytes) which are the basic metabolic cells. Lobules are the functional units of the liver.

Surfaces[edit]

On the diaphragmatic surface, apart from a large triangular bare area, where it connects to the diaphragm, the liver is covered by a thin double-layered membrane, the peritoneum, that reduces friction against other organs. 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.

These "peritoneal ligaments" are not related to the true anatomic ligaments in joints, and the right and left triangular ligaments have no known functional importance, but they serve as recognizable surface landmarks. The falciform ligament however, does function 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 gall bladder and the porta hepatis.

Lobes[edit]

Traditional gross anatomy divided the liver into two lobes (left and right), if viewed from the parietal surface; but if observed on the visceral surface it is divided into four lobes with the addition of the caudate and quadrate lobe.

The falciform ligament is visible on the front (anterior side) of the liver. This divides the liver into a left anatomical lobe, and a right anatomical lobe.

If the liver is flipped over, to look at it from below (the visceral surface), there are two additional lobes between the right and left. These are the caudate lobe (the more superior) and the quadrate lobe (the more inferior).

On the visceral surface a functional anatomy dictates how the liver is organized. One must view an imaginary line that passes to the left of the vena cava all the way forward and sections the gallblader into two halfs. This line is called Cantlie's Line. This line divides the liver into left and right. Other anatomical landmarks exist, such as the ligamentum venosum (ligamentum of Arancio) and the round ligament (ligamentum Teres) that further divide the left side of the liver in two sections. Now an important anatomical landmark, the transverse fissure of the liver divides this left portion of the liver into four segments which will be numbered starting at the caudate lobule as I in an anti-clock manner. From this visceral view we can see 7 segments because the 8th segment is only visible in the parietal view. Each of the lobes is made up of lobules; a vein goes from the centre, which then joins to the hepatic vein to carry blood out from the liver.

On the surface of the lobules, there are ducts, veins and arteries that carry fluids to and from them.

Impressions[edit]

Impressions on liver (posterior view)

There are several impressions on the surface of the liver which accommodate the various adjacent structures and organs. The portion of the under surface of the right lobe to the right of the fossa for the gall-bladder presents two impressions, one situated behind the other, and separated by a ridge. The posterior of these, the renal impression, is deeper and is occupied by the upper part of the right kidney and lower part of the right suprarenal gland. The anterior of these two impressions, the colic impression, is shallow and is produced by the right colic flexure.

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.

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.

The inferior surface of the left lobe of the liver presents behind and to the left the gastric impression, moulded over the antero-superior surface of the stomach, and to the right of this 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.

Functional anatomy[edit]

Correspondence between anatomic lobes and Couinaud segments
Segment* Couinaud segments
Caudate 1
Lateral 2, 3
Medial 4a, 4b
Right 5, 6, 7, 8

* or lobe, in the case of the caudate lobe
Each number in the list corresponds to one in the table. 1. Caudate
2. Superior subsegment of the lateral segment
3. Inferior subsegment of the lateral segment
4a. Superior subsegment of the medial segment
4b. Inferior subsegment of the medial segment
5. Inferior subsegment of the anterior segment
6. Inferior subsegment of the posterior segment
7. Superior subsegment of the posterior segment
8. Superior subsegment of the anterior segment

The central area where the common bile duct, hepatic portal vein, and hepatic artery proper enter is the hilum or "porta hepatis". The duct, vein, and artery divide into left and right branches, and the portions of the liver supplied by these branches constitute the functional left and right lobes.

The functional lobes are separated by an imaginary plane (historically called 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 fissure for the ligamentum teres also separates the medial and lateral segments. The medial segment is also called the quadrate lobe. In the widely used Couinaud (or "French") 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. The caudate lobe is a separate structure which receives blood flow from both the right- and left-sided vascular branches.[6][7]

Physiology[edit]

The various functions of the liver are carried out by the liver cells or hepatocytes. Currently, there is no artificial organ or device capable of emulating all the functions of the liver. Some functions can be emulated by liver dialysis, an experimental treatment for liver failure. The liver is thought to be responsible for up to 500 separate functions, usually in combination with other systems and organs.

Cell types[edit]

Two major types of cells populate the liver lobes: parenchymal and non-parenchymal cells. 80% of the liver volume is occupied by parenchymal cells commonly referred to as hepatocytes. Non-parenchymal cells constitute 40% of the total number of liver cells but only 6.5% of its volume. Sinusoidal hepatic endothelial cells, Kupffer cells and hepatic stellate cells are some of the non-parenchymal cells that line the liver sinusoid.[8]

Blood supply[edit]

The liver gets a dual blood supply from the hepatic portal vein and hepatic arteries. Supplying approximately 75% of the liver's blood supply, the hepatic portal vein 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 remainder of its blood flow. Oxygen is provided from both sources; approximately half of the liver's oxygen demand is met by the hepatic portal vein, and half is met by the hepatic arteries.[9]
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.

Biliary flow[edit]

The biliary tree

The term biliary tree is derived from the arboreal branches of the bile ducts. The bile produced in the liver is collected in bile canaliculi, which merge to form bile ducts. Within the liver, these ducts are called intrahepatic (within the liver) bile ducts, and once they exit the liver they are considered extrahepatic (outside the liver). The intrahepatic ducts eventually drain into the right and left hepatic ducts, which 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.

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 ampulla of Vater.

Synthesis[edit]

Breakdown[edit]

Other functions[edit]

Relation to medicine and pharmacology[edit]

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 regards to geriatric medicine.

Clinical significance[edit]

Disease[edit]

Main article: Liver disease
Left lobe liver tumor

The liver supports almost every organ in the body and is vital for survival. Because of its strategic location and multidimensional functions, the liver is also prone to many diseases.[11]

The most common include: Infections such as hepatitis A, B, C, D, E, alcohol damage, fatty liver, cirrhosis, cancer, drug damage (particularly by acetaminophen (paracetamol) and cancer drugs).

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 including biliary atresia, alpha-1 antitrypsin deficiency, alagille syndrome, progressive familial intrahepatic cholestasis, and Langerhans cell histiocytosis, to name but a few.

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.

Liver diseases may be diagnosed by liver function tests, for example, by production of acute phase proteins.

Symptoms[edit]

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, 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 substances which help prevent bleeding. When liver damage occurs, these substances are no longer present and severe bleeding can occur.[12]

Diagnosis[edit]

The diagnosis of liver function is made by blood tests. Liver function tests can readily pinpoint the extent of liver damage. If infection is suspected, then other serological tests are done. Sometimes, one may require an ultrasound or a CT scan to produce an image of the liver.

Physical examination of the liver is not accurate in determining the extent of liver damage. It can only reveal presence of tenderness or the size of liver, but in all cases, some type of radiological study is required to examine it.[13]

Biopsy / scan[edit]

Damage to the liver is sometimes determined with a biopsy, particularly when the cause of liver damage is unknown. In the 21st century they were largely replaced by high-resolution radiographic scans. The latter do not require ultrasound guidance, lab involvement, microscopic analysis, organ damage, pain, or patient sedation; and the results are available immediately on a computer screen.

In a biopsy, a needle is inserted into the skin just below the rib cage and a tissue sample obtained. The tissue 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 ultrasound guidance.[14]

Regeneration[edit]

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.[15] This is, however, not true regeneration but rather compensatory growth.[16] 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 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.[17]

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.[18] 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, the latter being the cells that line the bile ducts.

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 knew about the liver’s remarkable capacity for self-repair, however, this claim is purely speculative.[19]

Liver transplantation[edit]

Main article: Liver transplantation

Human liver transplants were first performed by Thomas Starzl in the United States and Roy Calne in Cambridge, England in 1963 and 1965, 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, autoimmune hepatitis, and many others. Less commonly, liver transplantation is done for fulminant hepatic failure, in which liver failure occurs over days to 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 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 have been at least two donor deaths out of the first several hundred cases. A recent publication has addressed the problem of donor mortality, and at least 14 cases have been found.[20] 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.

Development[edit]

Organogenesis[edit]

The origins of the liver lie in both the ventral portion of the foregut endoderm (endoderm being one of the 3 embryonic germ cell layers) and the constituents of the adjacent septum transversum mesenchyme. In 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.[21] 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.[22] Hepatic stellate cells are derived from mesenchyme.[23]

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 sinudoidal 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.[22]

Fetal blood supply[edit]

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 develops throughout normal gestation, and does not perform the normal filtration of the infant liver. The liver does not perform digestive processes because the fetus does not consume meals directly, but receives nourishment from the mother via the placenta. The fetal liver releases some blood stem cells that migrate to the fetal thymus, so initially the lymphocytes, called T-cells, are created from fetal liver stem cells. Once the fetus is delivered, the formation of blood stem cells in infants shifts to the red bone marrow.

After birth, the umbilical vein and ductus venosus are completely obliterated in two to five days; the former becomes the ligamentum teres and the latter becomes the ligamentum venosum. In the disease state of cirrhosis and portal hypertension, the umbilical vein can open up again.

During childhood[edit]

At birth the liver comprises roughly 4% of body weight and is at average 120g. Over the course of developement it will increase to 1,4–1,6 kg but will only take up 2.5–3.5% of body weight.[24]

Society and culture[edit]

In Greek mythology, Prometheus was punished by the gods for revealing fire to humans, by being chained 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, 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.[25] 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 (جگر or जिगर 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!".[26] 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.

The legend of Liver-Eating Johnson says that he would 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 narrations have to be questioned.

Additional Images[edit]

See also[edit]

References[edit]

  1. ^ Physiology at MCG 6/6ch2/s6ch2_30
  2. ^ Maton, Anthea; Jean Hopkins; Charles William McLaughlin; Susan Johnson; Maryanna Quon Warner; David LaHart; Jill D. Wright (1993). Human Biology and Health. Englewood Cliffs, New Jersey, USA: Prentice Hall. ISBN 0-13-981176-1. OCLC 32308337. 
  3. ^ "Etymology online hepatic". Retrieved December 12, 2013. 
  4. ^ "Etymology online hepatos". Retrieved December 12, 2013. 
  5. ^ Cotran, Ramzi S.; Kumar, Vinay; Fausto, Nelson; Nelso Fausto; Robbins, Stanley L.; Abbas, Abul K. (2005). Robbins and Cotran pathologic basis of disease (7th ed.). St. Louis, MO: Elsevier Saunders. p. 878. ISBN 0-7216-0187-1. 
  6. ^ "Three-dimensional Anatomy of the Couinaud Liver Segments". Retrieved 2009-02-17. 
  7. ^ Strunk, Holger. "Limitations and Pitfalls of Couinaud`s Segmentation of the Liver in Transaxial Imaging". Retrieved 2009-02-17. 
  8. ^ Kmieć Z (2001). "Cooperation of liver cells in health and disease". Adv Anat Embryol Cell Biol 161: III–XIII, 1–151. PMID 11729749. 
  9. ^ Shneider, Benjamin L. ; Sherman, Philip M. (2008). Pediatric Gastrointestinal Disease. Connecticut: PMPH-USA. p. 751. ISBN 1-55009-364-9. 
  10. ^ Sheporaitis, L; Freeny, PC (1998). "Hepatic and portal surface veins: A new anatomic variant revealed during abdominal CT". AJR. American journal of roentgenology 171 (6): 1559–64. doi:10.2214/ajr.171.6.9843288. PMID 9843288. 
  11. ^ Cirrhosis Overview National Digestive Diseases Information Clearinghouse. Retrieved on 2010-01-22
  12. ^ Extraintestinal Complications: Liver Disease Crohn's & Colitis Foundation of America. Retrieved on 2010-01-22
  13. ^ Liver Information HealthLine. Retrieved on 2010-01-22
  14. ^ Liver.. The largest gland in the body MedicineNet. Retrieved on 2010-01-22
  15. ^ Dieter Häussinger, ed. (2011). Liver Regeneration. Berlin: De Gruyter. p. 1. ISBN 9783110250794. 
  16. ^ Robbins and Cotran Pathologic Basis of Disease (7th ed.). 1999. p. 101. ISBN 0-8089-2302-1. 
  17. ^ W. T. Councilman (1913). "Two". Disease and Its Causes. New York Henry Holt and Company London Williams and Norgate The University Press, Cambridge, U.S.A. 
  18. ^ Suzuki K, Tanaka M, Watanabe N, Saito S, Nonaka H, Miyajima A (2008). "p75 Neurotrophin receptor is a marker for precursors of stellate cells and portal fibroblasts in mouse fetal liver". Gastroenterology 135 (1): 270–281.e3. doi:10.1053/j.gastro.2008.03.075. PMID 18515089. 
  19. ^ An argument for the ancient Greek’s knowing about liver regeneration is provided by Chen, T. S.; Chen, P. S. (1994). "The myth of Prometheus and the liver". Journal of the Royal Society of Medicine 87 (12): 754–755. PMC 1294986. PMID 7853302.  edit Counterarguments are provided by Tiniakos, D. G.; Kandilis, A.; Geller, S. A. (2010). "Tityus: A forgotten myth of liver regeneration". Journal of Hepatology 53 (2): 357–361. doi:10.1016/j.jhep.2010.02.032. PMID 20472318.  edit and by Power, C.; Rasko, J. E. (2008). "Whither prometheus' liver? Greek myth and the science of regeneration". Annals of internal medicine 149 (6): 421–426. PMID 18794562.  edit
  20. ^ Bramstedt K (2006). "Living liver donor mortality: where do we stand?". Am. J. Gastrointestinal 101 (4): 755–9. doi:10.1111/j.1572-0241.2006.00421.x. PMID 16494593. 
  21. ^ Gilbert SF (2000). Developmental Biology (6th ed.). Sunderland (MA): Sinauer Associates. 
  22. ^ a b Lade AG, Monga SP (2011). "Beta-catenin signaling in hepatic development and progenitors: which way does the WNT blow?". Dev Dyn 240 (3): 486–500. doi:10.1002/dvdy.22522. PMID 21337461. 
  23. ^ Berg T, DeLanghe S, Al Alam D, Utley S, Estrada J, Wang KS (2010). "β-catenin regulates mesenchymal progenitor cell differentiation during hepatogenesis". J Surg Res 164 (2): 276–85. doi:10.1016/j.jss.2009.10.033. PMC 2904820. PMID 20381814. 
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  25. ^ Krishna, Gopi; Hillman, James (commentary) (1970). Kundalini – the evolutionary energy in man. London: Stuart & Watkins. p. 77. SBN 7224 0115 9. 
  26. ^ The Great Battle Of Badar (Yaum-E-Furqan). Shawuniversitymosque.org (2006-07-08). Retrieved on 2013-03-19.

External links[edit]