Category Archives: Physiology

Recent Articles | Physiology | The Scientist Magazine

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Running releases an enzyme that is associated with memory function in mice and humans.

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By Chris Tachibana | February 1, 2016

Is the public dissection of zoo animals a boon to research and education, a PR nightmare, or both?

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By The Scientist Staff | February 1, 2016

Watch footage from the public dissection of Marius, the young giraffe at the Copenhagen Zoo who was ultimately fed to predators at the facility. (CAUTION: GRAPHIC IMAGES)

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By Bob Grant | January 25, 2016

Researchers confirm the unprecedented endothermic abilities of a South American reptile.

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By Kerry Grens | November 16, 2015

In mice lacking intestinal microbiota, white fat turns brown and obesity is prevented.

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By Kerry Grens | November 5, 2015

Scientists describe molecular underpinnings of salmon size and of fishes ability to navigate murky environments in separate studies.

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By Tracy Vence | November 1, 2015

The accumulation of fat within skeletal muscle, as happens with obesity, diminishes muscle performance.

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The latest analysis on GDF11, a proposed antiaging protein, blames discrepancies in the literature on misreported doses and misinterpretation.

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Researchers derive trophoblast stem cells from mouse fibroblasts, paving the way for cell therapy for placental dysfunction diseases.

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By Kerry Grens | October 8, 2015

Scientists know how to turn on these fat-combusting cells. Can these energy burners be used to combat obesity?

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Recent Articles | Physiology | The Scientist Magazine

Physiology – Superpower Wiki – Wikia

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What is physiology? | Physiological Society

Physiology is the study of how molecules, cells and organs interact to form a whole being. The work of Physiological Society Members, advancing our knowledge of biological systems, is essential to the development of new treatments for disease.Since The Society's foundation in 1876, our membership has included more than 20 Nobel Prize winners from Ivan Pavlov to Andrew Huxley. The scientists who make up The Society have made many key discoveries, ranging from how our nervous system works,how our cells divide andthe way in which our reflexes alter our behaviour. These have advanced our knowledge of biological systems and helped in the treatment of diseases such as cancer, cystic fibrosis and heart disease.

As well as supporting those active in physiology research, we also work to inspire the next generations of physiologists; working with teachers, lecturers and students to help highlight physiology as a relevant, engaging career choice. The Society's presence at public events such as science fairs and media promotion of the research it publishes in its journals, also aim to raise the profile of physiology and highlight its relevance to everyday life.

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What is physiology? | Physiological Society

Physiology – Wikipedia

Physiology (; from Ancient Greek (physis), meaning "nature, origin", and - (-logia), meaning "study of"[1]) is the scientific study of the normal function in living systems.[2] A sub-discipline of biology, its focus is in how organisms, organ systems, organs, cells, and biomolecules carry out the chemical or physical functions that exist in a living system.[3] Given the size of the field, it is divided into, among others, animal physiology (including that of humans), plant physiology, cellular physiology, microbial physiology (microbial metabolism), bacterial physiology, and viral physiology.[3] The Nobel Prize in Physiology or Medicine is awarded to those who make significant achievements in this discipline by the Royal Swedish Academy of Sciences. In medicine, a physiologic state is one occurring from normal body function, rather than pathologically, which is centered on the abnormalities that occur in animal diseases, including humans.[4]

Physiological studies date back to ancient civilizations of India[5][6] and Egypt alongside anatomical studies, but did not utilize dissection or vivisection.[7]

The study of human physiology as a medical field dates back to at least 420BC to the time of Hippocrates, also known as the "father of medicine."[8] Hippocrates incorporated his belief system called the theory of humours, which consisted of four basic substance: earth, water, air and fire. Each substance is known for having a corresponding humour: black bile, phlegm, blood and yellow bile, respectively. Hippocrates also noted some emotional connections to the four humours, which Claudis Galenus would later expand on. The critical thinking of Aristotle and his emphasis on the relationship between structure and function marked the beginning of physiology in Ancient Greece. Like Hippocrates, Aristotle took to the humoral theory of disease, which also consisted of four primary qualities in life: hot, cold, wet and dry.[9] Claudius Galenus (c. ~130200AD), known as Galen of Pergamum, was the first to use experiments to probe the functions of the body. Unlike Hippocrates though, Galen argued that humoral imbalances can be located in specific organs, including the entire body.[10] His modification of this theory better equipped doctors to make more precise diagnoses. Galen also played off of Hippocrates idea that emotions were also tied to the humours, and added the notion of temperaments: sanguine corresponds with blood; phlegmatic is tied to phlegm; yellow bile is connected to choleric; and black bile corresponds with melancholy. Galen also saw the human body consisting of three connected systems: the brain and nerves, which are responsible for thoughts and sensations; the heart and arteries, which give life; and the liver and veins, which can be attributed to nutrition and growth.[10] Galen was also the founder of experimental physiology.[11] And for the next 1,400 years, Galenic physiology was a powerful and influential tool in medicine.[10]

Jean Fernel (14971558), a French physician, introduced the term "physiology".[12]

In 1858, Joseph Lister studied the cause of blood coagulation and inflammation that resulted after previous injuries and surgical wounds. He later discovered and implemented antiseptics in the operating room, and as a result decreases death rate from surgery by a substantial amount.[4][13]

In 1891, Ivan Pavlov performed research on "conditional reflexes" that involved dogs' saliva production in response to a plethora of sounds and visual stimuli.[13]

In the 19th century, physiological knowledge began to accumulate at a rapid rate, in particular with the 1838 appearance of the Cell theory of Matthias Schleiden and Theodor Schwann. It radically stated that organisms are made up of units called cells. Claude Bernard's (18131878) further discoveries ultimately led to his concept of milieu interieur (internal environment), which would later be taken up and championed as "homeostasis" by American physiologist Walter B. Cannon in 1929. By homeostasis, Cannon meant "the maintenance of steady states in the body and the physiological processes through which they are regulated."[14] In other words, the body's ability to regulate its internal environment. It should be noted that, William Beaumont was the first American to utilize the practical application of physiology.

The Physiological Society was founded in London in 1876 as a dining club The American Physiological Society (APS) is a nonprofit devoted to fostering education, scientific research, and dissemination of information in the physiological sciences. The Society was founded in 1887 with 28 members.

In the 20th century, biologists became interested in how organisms other than human beings function, eventually spawning the fields of comparative physiology and ecophysiology.[15] Major figures in these fields include Knut Schmidt-Nielsen and George Bartholomew. Most recently, evolutionary physiology has become a distinct subdiscipline.[16]

In 1920, August Krogh won the Nobel Prize for discovering how, in capillaries, blood flow is regulated.[13]

In 1954, Andrew Huxley and Hugh Huxley, alongside their research team, discovered the sliding filaments in skeletal muscle, known today as the sliding filament theory.[13]

Initially, women were largely excluded from official involvement in any physiological society. The American Physiological Society, for example, was founded in 1887 and included only men in its ranks.[citation needed] In 1902, the American Physiological Society elected Ida Hyde as the first female member of the society.[citation needed] Hyde, a representative of the American Association of University Women and a global advocate for gender equality in education,[17] attempted to promote gender equality in every aspect of science and medicine.

Soon thereafter, in 1913, J.S. Haldane proposed that women be allowed to formally join The Physiological Society, which had been founded in 1876.[citation needed] On 3 July 1915, six women were officially admitted: Florence Buchanan, Winifred Cullis, Ruth C. Skelton, Sarah C. M. Sowton, Constance Leetham Terry, and Enid M. Tribe.[18] The centenary of the election of women was celebrated in 2015 with the publication of a book "Women physiologists: centenary celebrations and beyond for The Physiological Society ISBN 978-0-9933410-0-7.

Prominent women physiologists include:

There are many ways to categorize the subdiscplines of physiology:[27]

Human physiology seeks to understand the mechanisms that work to keep the human body alive and functioning,[3] through scientific enquiry into the nature of mechanical, physical, and biochemical functions of humans, their organs, and the cells of which they are composed. The principal level of focus of physiology is at the level of organs and systems within systems. The endocrine and nervous systems play major roles in the reception and transmission of signals that integrate function in animals. Homeostasis is a major aspect with regard to such interactions within plants as well as animals. The biological basis of the study of physiology, integration refers to the overlap of many functions of the systems of the human body, as well as its accompanied form. It is achieved through communication that occurs in a variety of ways, both electrical and chemical.[citation needed]

Much of the foundation of knowledge in human physiology was provided by animal experimentation. Physiology is the study of function and is closely related to anatomy which is the study of form and structure. Due to the frequent connection between form and function, physiology and anatomy are intrinsically linked and are studied in tandem as part of a medical curriculum.[28]

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Physiology - Wikipedia

What is Physiology | Human Physiology | Understanding Life …

Physiology is the science of life. It is a broad science which aims to understand the mechanisms of living, from the molecular basis of cell function to the integrated behaviour of the whole body.

Research in physiology helps us to understand how the body works; it also helps us to realise what goes wrong in disease and to identify new treatments for disease.

Physiology forms an integral part of pre- and post-16 biology education, and can also be studied at university either as a stand-alone discipline or as part of an integrated degree, such as biomedical sciences. For more information about career paths in physiology, please visit the careers section of this website.

Pre-16, the study of physiology focuses primarily on how the body moves, and the structure and function of some of the major organ systems (including the cardiovascular and respiratory systems). Post-16, the study of physiology leans more towards the understanding of physiological processes such as homeostasis and excretion.

A degree in physiology will build on the knowledge and understanding developed at school/college: it will explore selected topics in greater detail and provide a holistic view of how the different cells, tissues, organs and systems of the body are integrated. Physiologists - scientists who have chosen to explore physiology as a career will continue to build on the knowledge they have gained during their degree and advance the science of life within an area of particular interest to them. It is important to highlight, however, that physiologists do not work in isolation: the sharing of information between scientists around the world is essential to continue developing our understanding of how the body works.

Physiology is an experimental science that underpins the biological and clinical sciences; it is key to the detection, prevention and treatment of disease. Without an understanding of basic physiology, progress made in other areas such as the sequencing of the human genome is limited because every biological advance must ultimately be related to the behaviour of the whole organism.

The Physiological Society recognises the importance of using animals in research in order to gain further knowledge of disease mechanisms in both animal and human diseases. We appreciate that this can be a difficult topic to understand and teach and have therefore developed supporting resources designed to address this area specifically.

To hear what physiology means to our members, listen to the podcasts available in our resources section.

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Physiology jobs – Science Careers

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York, Pennsylvania Competitive salary with benefits York College of Pennsylvania

York College is seeking an Animal Physiologist for a three year visiting assistant professor position to teach lecture and laboratory courses.

Kln W3 Uniklinik Kln

The University of Cologne, Faculty of Medicine, posts the position of a Professorship (W3) for vegetative Physiology as per the requirements outlin...

La Crosse, WI Academic year salary is competitive and commensurate with experience. University of Wisconsin-La Crosse Biology Department

The Department of Biology in the College of Science and Health at the University of Wisconsin - La Crosse (UWL) invites applications for an academi...

Durham, North Carolina, USA NIH scale Rawls Lab

Postdoctoral position at Duke University investigating the mechanisms by which microbiomes regulate host metabolism in adolescents with obesity

Saint Louis, Missouri (US) Salary is in keeping with NIH guidelines egantm@slu.edu

We seek highly motivated applicants to study native and recombinant ligand-gated ion channels using electrophysiology and fluorescence microscopy

Atlanta, Georgia, USA Competitive salaries at the corresponding faculty rank EMORY UNIVERSITY - Physiology search

Applications are invited for one Tenure Track Faculty position at the level of Associate or Full Professor in the Department of Physiology at Emory...

Saint Paul, Minnesota 9-month Appointment University of Minnesota

Are you a creative microbiologist who excels at working collaboratively and pushing disciplinary boundaries? Then we want to hear from you. The Col...

San Marcos, California Commensurate with experience California State University San Marcos-Biology

The Department of Biological Sciences at California State University San Marcos (CSUSM) is pleased to be seeking applications for a tenure-track As...

Providence, Rhode Island (US) Salary and benefits will be commensurate with NIH guidelines. Brown University, Dept. of Pathology and Laboratory Medicine

Postdoctoral Research Associate: Biology of Metals An NIH-funded postdoctoral position is available in the laboratory of Dr. Tom Bartnikas in the D...

St. Paul, Minnesota (US) Undisclosed College of Biological Sciences at the University of Minnesota

We seek outstanding microbiologists who utilize various approaches to understand microorganisms ...

Woods Hole, Massachusetts Undisclosed Marine Biological Laboratory

designed to allow exceptional scientists early in their careers to establish their own research programs...

Paris, France (FR) Highly attractive packages to match the experience of the candidate will be provided Institut Pasteur

... establish new independent research groups in the cutting edge interdisciplinary environment ...

Paris, France (FR) Undisclosed Institut Pasteur

... launched an international call for a junior* research group studying the gut host and microbiome homeostasis ...

Manhattan, Kansas Associate Level: includes competitive salary and start-up funding The Department of Biochemistry and Molecular Biophysics at Kansas State University

We seek individuals who will develop a strong research program and excel in teaching a diverse population of undergraduate and graduate students

Ypsilanti, Michigan Competitive salary and benefits Eastern Michigan University

EASTERN MICHIGAN UNIVERSITY Plant Ecologist The Department of Biology invites applications for a tenure-track position in Plant Ecology at the leve...

Stevens Point, Wisconsin Negotiable. With full health and pension benefits. University of Wisconsin - Stevens Point

The Department of Biology at the University of Wisconsin-Stevens Point is offering a tenure-track, nine-month faculty position in Human Physiology.

New York City, New York (US) TBD Columbia University

The Department of Pharmacology of Columbia University seeks one or more Postdoctoral Research Scientists to join our team.

Toronto (Region), Ontario (CA) N/A Department of Biology, York University

The Department of Biology, York University invites applications from emerging world-class leaders in neurophysiology to be nominated for a Tier 2 CRC.

Taiwan (TW) none Institute of Cellular and Organismica Biology, Academia Sinica

The Institute of Cellular and Organismic Biology of the Academia Sinica, Taiwan seeks exceptional, creative scientists to join its faculty.

Chapel Hill, North Carolina (US) Dependent on Qualifications UNC Chapel Hill Cell Biology and Physiology

The Dept. of Cell Biology & Physiology at UNC Chapel Hill invites applications for several tenure track positions at the level of Assistant Professor

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Physiology jobs - Science Careers

Physiology | Animal Science

General areas of research in physiology include factors affecting gonadal development (ovarian and testis) and sex differentiation, factors affecting embryo development and reproductive hormone research, and genes regulating egg and sperm function. Undergraduate employment opportunities include maintaining mice and rate colonies including daily care, dissections of rats and mice, in vitro fertilization of mice, genotyping mice to determine if they carry the trans-gene, jobs in pig physiology, general lab work, and aiding with blood collection in cattle, artificial insemination, ultrasonography, palpation, and estrus detection. Recent and ongoing research includes:

Faculty members at Animal Science will be glad to help you with any questions you might have.

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Physiology | Animal Science

Careers in Physiology (Physiology Majors) – Monash University

The study of physiology prepares you for a wide range of careers. You can apply your knowledge of physiology directly, choosing a career in the biomedical sphere. Physiology graduates can also use the general skills and knowledge they have acquired to pursue a career in a variety of workplaces. Just some of the careers in which Physiology graduates are currently employed are shown below with relevant links where available.

Please note: An additional qualification may be required for some of the careers shown below.

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Careers in Physiology (Physiology Majors) - Monash University

Liver – Wikipedia, the free encyclopedia

The liver is a vital organ of vertebrates and some other animals.[2] In the human, it is located in the upper right quadrant of the abdomen, below the diaphragm. The liver has a wide range of functions, including detoxification of various metabolites, protein synthesis, and the production of biochemicals necessary for digestion.[3]

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.[3] It is an accessory digestive gland and produces bile, an alkaline compound which aids in digestion via the emulsification of lipids. The gallbladder, a small pouch that sits just under the liver, stores bile produced by the liver.[4] 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.[5] Estimates regarding the organ's total number of functions vary, but textbooks generally cite it being around 500.[6]

Terminology related to the liver often starts in hepat- from the Greek word for liver.[7]

There is currently no way 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 are yet to be developed to promote long term replacement in the absence of the liver. As of now,[8]liver transplantation is the only option for complete liver failure.

The liver is a reddish brown wedge-shaped organ with four lobes of unequal size and shape. A human liver normally weighs 1.441.66kg (3.23.7lb).[9] 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.[4]

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.[8] These blood vessels subdivide into small capillaries known as liver sinusoids, which then lead to a lobule.

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 fine areolar tissue which extends into the structure of the liver, by accompanying the vessels (veins and arteries) ducts and nerves through the hepatic portal, as a fibrous capsule called Glisson's capsule.[10] The whole surface of the liver is covered in a serous coat derived from peritoneum and this has an inner fibrous coat (Glisson's capsule) to which it is firmly adhered. The fibrous coat is of areolar tissue and follows the vessels and ducts to support them.

Gross anatomy traditionally divided the liver into two portions a right and a left lobe, as viewed from the front (diaphragmatic) surface; but the underside (the visceral surface) shows it to be divided into four lobes and includes the caudate and quadrate lobes.[11]

The falciform ligament, visible on the front of the liver, divides the liver into a left and a much larger right lobe. From the visceral surface, 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.[12] This line is called Cantlie's line.[13]

Other anatomical landmarks exist, such as the ligamentum venosum and the round ligament of the liver (ligamentum teres), which further divide the left side of the liver in two sections. An important anatomical landmark, the porta hepatis, also known as the transverse fissure of the liver, divides this left portion into four segments, which can be numbered starting at the caudate lobe as I in an anticlockwise manner. From this visceral view, seven segments can be seen, because the eighth segment is only visible in the parietal view.[14]

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 help reduces friction against other organs.[15] 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.[16]

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.[16] 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.[15]

There are several impressions on the surface of the liver which 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.[17]

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.[18]

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.[18]

The inferior surface of the left lobe of the liver presents behind and to the left the gastric impression.[18] 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.

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 central vein.[19][pageneeded]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, misleadingly named, consists of five structures: a branch of the hepatic artery, a branch of the hepatic portal vein, and a bile duct, as well as lymphatic vessels and a branch of the vagus nerve.[20] Between the hepatocyte plates are liver sinusoids, which are enlarged capillaries through which blood from the hepatic portal vein and hepatic artery enters via the portal triads, then drains to the central vein.[19][pageneeded]

Histology, the study of microscopic anatomy, shows two major types of liver cell: parenchymal cells and non-parenchymal cells. 7085% of the liver volume is occupied by parenchymal hepatocytes. Non-parenchymal cells constitute 40% of the total number of liver cells but only 6.5% of its volume.[21] The liver sinusoids are lined with two types of cell, sinusoidal endothelial cells, and phagocytic Kupffer cells.[22]Hepatic stellate cells are non-parenchymal cells found in the perisinusoidal space, between a sinusoid and a hepatocyte.[21] Additionally, intrahepatic lymphocytes are often present in the sinusoidal lumen.[21]

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 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 fissure for the round ligament of the liver (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.[23] The caudate lobe is a separate structure which receives blood flow from both the right- and left-sided vascular branches.[24][25] The Couinaud classification of liver anatomy divides the liver into eight functionally independent segments. Each segment has its own vascular inflow, outflow and biliary drainage. In the centre of each segment there is a branch of the portal vein, hepatic artery and bile duct. In the periphery of each segment there is vascular outflow through the hepatic veins.[26] The division of the liver into independent units means that segments can be resected without damaging the remaining segments.[27] To preserve the viability of the liver following surgery, resections follow the vessels defining the peripheries of each segment. This means that resection lines parallel the hepatic veins, leaving the portal veins, bile ducts, and hepatic arteries intact.[23]

The classification system uses the vascular supply in the liver to separate the functional units (numbered I to VIII):

The remainder of the units (II to VIII) are numbered in a clockwise fashion:[26]

Units V to VIII make up the right part of the liver:[26]

Organogenesis, the development of the organs takes place from the third to the eighth week in human embryogenesis. The origins of the liver lie in both the ventral portion of the foregut endoderm (endoderm being one of the 3 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.[28] 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.[29]Hepatic stellate cells are derived from mesenchyme.[30]

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/-catenin has now been identified to be playing a key role in this phenomenon.[29]

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 two to five 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.

At birth the liver comprises roughly 4% of body weight and is at average 120 g. Over the course of development, it will increase to 1.41.6kg but will only take up 2.53.5% of body weight.[31]

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, there is no artificial organ or device 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 receives a dual blood supply from the hepatic portal vein and hepatic arteries. The hepatic portal vein delivers approximately 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; approximately half of the liver's oxygen demand is met by the hepatic portal vein, and half is met by the hepatic arteries.[32]

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.[20]

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.[20]

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.

The liver plays a major role in carbohydrate, protein, amino acid, and lipid metabolism.

The liver performs several roles in carbohydrate metabolism: The liver synthesizes and stores approximately 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.[33] 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.[33]

The liver is responsible for the mainstay of protein metabolism, synthesis as well as degradation. 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, XIII, as well as protein C, protein S and antithrombin. 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. The liver is a major site of production for thrombopoietin, a glycoprotein hormone that regulates the production of platelets by the bone marrow.[34]

The liver plays several roles in lipid metabolism: it performs cholesterol synthesis, lipogenesis, 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 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 effects in adults.

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 breaks down ammonia into urea as part of the urea cycle, and the urea is excreted in the urine.[19]

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.

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 also prone to many diseases.[36] 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.

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 Herpesviridae family such as the herpes simplex virus. Infection with hepatitis B virus or hepatitis C virus is the main cause of liver cancer.[37]

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.

Other disorders caused by excessive alcohol consumption are grouped under alcoholic liver diseases and these include alcoholic hepatitis, fatty liver, and cirrhosis. Liver damage can also be caused by drugs, particularly paracetamol and drugs used to treat cancer.

BuddChiari 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.[38]

Primary biliary cirrhosis is an autoimmune disease of the liver.[39][40] 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.[41] 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.

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, 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 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.

Liver diseases may be diagnosed by liver function testsblood tests that can identify various markers. For example, acute-phase reactants are produced by the liver in response to injury or inflammation.

The classic symptoms of liver damage include the following:

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. Sometimes, an ultrasound or a CT scan is needed to produce an image of the liver.

Physical examination of the liver can only reveal its size and any tenderness, and some form of imaging will also be needed.[43]

Axial CT image showing anomalous hepatic veins coursing on the subcapsular anterior surface of the liver.[44]

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

A CT scan in which the liver and portal vein are shown.

Damage to the liver is sometimes determined with a biopsy, particularly when the cause of liver damage is unknown. In the 21st century they have been 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.[citation needed]

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.[45]

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.[46] This is, however, not true regeneration but rather compensatory growth in mammals.[47] 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 fish, the liver undergoes true regeneration by restoring both shape and size of the organ.[48] 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.[49]

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.[50] 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.[51] 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.

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 livers remarkable capacity for self-repair.[52]

Human liver transplants were first performed by Thomas Starzl in the United States and Roy Calne in Cambridge, England in 1963 and 1967, respectively.

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.[53] 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.

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.[54] 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 ( or 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!".[55] 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 has to be questioned.

On November 26, 1987, the city of Ferrol, Spain, inaugurated what is believed to be the only monument to the liver in the world. The then - major, 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".[56]

The liver of mammals, fowl, and fish are commonly eaten as food by humans. Domestic pig, ox, lamb, calf, chicken, and goose livers are widely available from butchers and supermarkets.

Liver can be baked, boiled, broiled, fried, stir-fried, or eaten raw (asbeh nayeh or sawda naye in Lebanese cuisine, liver sashimi). In many preparations, pieces of liver are combined with pieces of meat or kidneys, like in the various forms of Middle Eastern mixed grill (e.g. meurav Yerushalmi). Liver is often made into spreads. Well-known examples include liver pt, 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, namely skilpadjies, is made of minced lamb's liver wrapped in netvet (caul fat), and grilled over an open fire.

Animal livers are rich in iron and vitamin A, and cod liver oil is commonly used as a dietary supplement. 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.[57]

The liver is found in all vertebrates, and is typically the largest visceral (internal) organ. 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. The internal structure of the liver is broadly similar in all vertebrates.[58]

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 a homolog of the vertebrate liver.[59][60][61] The amphioxus hepatic caecum produces the liver-specific proteins vitellogenin, antithrombin, plasminogen, alanine aminotransferase, and insulin/Insulin-like growth factor (IGF)[62]

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

Liver lifted to show gall bladder and stomach in situ.

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

Cross section of an inferior portion of the liver, showing gallbladder and various structures.

Human liver. Visceral surface of liver.

Human liver. Horizontal section to newborn

Showing ligaments and bare area

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Nobel Prizes in Physiology or Medicine have been awarded between 1901 and 2015.

Medicine Prizes have been given to one Laureate only.

women have been awarded the Medicine Prize so far.

years was the age of the youngest Medicine Laureate ever, Frederick G. Banting, who was awarded the 1923 Medicine Prize for the discovery of insulin.

years was the age of the oldest Medicine Laureate ever, Peyton Rous, when he was awarded the Medicine Prize in 1966 for his discovery of tumour-inducing viruses.

All facts and figures about the Nobel Prize in Physiology or Medicine

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