Diagram of pituitary and pineal glands in the human brain
|Precursor||Neural ectoderm, roof of diencephalon|
|Artery||posterior cerebral artery|
|Anatomical terms of neuroanatomy|
The pineal gland, conarium, or epiphysis cerebri, is a small endocrine gland in the brain of most vertebrates. The pineal gland produces melatonin, a serotonin-derived hormone which modulates sleep patterns in both circadian and seasonal cycles. The shape of the gland resembles a pine cone from which it derived its name.The pineal gland is located in the epithalamus, near the center of the brain, between the two hemispheres, tucked in a groove where the two halves of the thalamus join. The pineal gland is one of the neuroendocrine secretory circumventricular organs in which there does not exist the blood–brain barrier at the capillary level.
Nearly all vertebrate species possess a pineal gland. The most important exception is a primitive vertebrate, the hagfish. Even in the hagfish, however, there may be a "pineal equivalent" structure in the dorsal diencephalon.The lancelet Branchiostoma lanceolatum , the nearest existing relative to vertebrates, also lacks a recognizable pineal gland. The lamprey (another primitive vertebrate), however, does possess one. A few more developed vertebrates have lost pineal glands over the course of their evolution.
The results of various scientific research in evolutionary biology, comparative neuroanatomy and neurophysiology have explained the evolutionary history (phylogeny) of the pineal gland in different vertebrate species. From the point of view of biological evolution, the pineal gland represents a kind of atrophied photoreceptor. In the epithalamus of some species of amphibians and reptiles, it is linked to a light-sensing organ, known as the parietal eye, which is also called the pineal eye or third eye.
René Descartes believed the human pineal gland to be the "principal seat of the soul". Academic philosophy among his contemporaries considered the pineal gland as a neuroanatomical structure without special metaphysical qualities; science studied it as one endocrine gland among many.
The word pineal, from Latin pinea (pine-cone), was first used in the late 17th century to refer to the cone shape of the brain gland.
The pineal gland is a midline brain structure that is unpaired. It takes its name from its pine-cone shape. mm) in humans. The pineal gland, also called the pineal body, is part of the epithalamus, and lies between the laterally positioned thalamic bodies and behind the habenular commissure. It is located in the quadrigeminal cistern near to the corpora quadrigemina. It is also located behind the third ventricle and is bathed in cerebrospinal fluid supplied through a small pineal recess of the third ventricle which projects into the stalk of the gland.The gland is reddish-gray and about the size of a grain of rice (5–8
Unlike most of the mammalian brain, the pineal gland is not isolated from the body by the blood–brain barrier system;it has profuse blood flow, second only to the kidney, supplied from the choroidal branches of the posterior cerebral artery.
The pineal gland receives a sympathetic innervation from the superior cervical ganglion. A parasympathetic innervation from the pterygopalatine and otic ganglia is also present.Further, some nerve fibers penetrate into the pineal gland via the pineal stalk (central innervation). Also, neurons in the trigeminal ganglion innervate the gland with nerve fibers containing the neuropeptide PACAP.
The pineal body consists in humans of a lobular parenchyma of pinealocytes surrounded by connective tissue spaces. The gland's surface is covered by a pial capsule.
The pineal gland consists mainly of pinealocytes, but four other cell types have been identified. As it is quite cellular (in relation to the cortex and white matter), it may be mistaken for a neoplasm.
|Pinealocytes||The pinealocytes consist of a cell body with 4–6 processes emerging. They produce and secrete melatonin. The pinealocytes can be stained by special silver impregnation methods. Their cytoplasm is lightly basophilic. With special stains, pinealocytes exhibit lengthy, branched cytoplasmic processes that extend to the connective septa and its blood vessels.|
|Interstitial cells||Interstitial cells are located between the pinealocytes. They have elongated nuclei and a cytoplasm that is stained darker than that of the pinealocytes.|
|Perivascular phagocyte||Many capillaries are present in the gland, and perivascular phagocytes are located close to these blood vessels. The perivascular phagocytes are antigen presenting cells.|
|Pineal neurons||In higher vertebrates neurons are usually located in the pineal gland. However, this is not the case in rodents.|
|Peptidergic neuron-like cells||In some species, neuronal-like peptidergic cells are present. These cells might have a paracrine regulatory function.|
The human pineal gland grows in size until about 1–2 years of age, remaining stable thereafter,although its weight increases gradually from puberty onwards. The abundant melatonin levels in children are believed to inhibit sexual development, and pineal tumors have been linked with precocious puberty. When puberty arrives, melatonin production is reduced.
In the zebrafish the pineal gland does not straddle the midline but shows a left-sided bias. In humans, functional cerebral dominance is accompanied by subtle anatomical asymmetry.
The primary function of the pineal gland is to produce melatonin. Melatonin has various functions in the central nervous system, the most important of which is to help modulate sleep patterns. Melatonin production is stimulated by darkness and inhibited by light.Light sensitive nerve cells in the retina detect light and send this signal to the suprachiasmatic nucleus (SCN), synchronizing the SCN to the day-night cycle. Nerve fibers then relay the daylight information from the SCN to the paraventricular nuclei (PVN), then to the spinal cord and via the sympathetic system to superior cervical ganglia (SCG), and from there into the pineal gland.
The compound pinoline is also claimed to be produced in the pineal gland; it is one of the beta-carbolines.This claim is subject to some controversy.
Studies on rodents suggest that the pineal gland influences the pituitary gland's secretion of the sex hormones, follicle-stimulating hormone (FSH), and luteinizing hormone (LH). Pinealectomy performed on rodents produced no change in pituitary weight, but caused an increase in the concentration of FSH and LH within the gland.Administration of melatonin did not return the concentrations of FSH to normal levels, suggesting that the pineal gland influences pituitary gland secretion of FSH and LH through an undescribed transmitting molecule.
The pineal gland contains receptors for the regulatory neuropeptide, endothelin-1,which, when injected in picomolar quantities into the lateral cerebral ventricle, causes a calcium-mediated increase in pineal glucose metabolism.
Studies in mice suggest that the pineal-derived melatonin regulates new bone deposition. Pineal-derived melatonin mediates its action on the bone cells through MT2 receptors. This pathway could be a potential new target for osteoporosis treatment as the study shows the curative effect of oral melatonin treatment in a postmenopausal osteoporosis mouse model.
Calcification of the pineal gland is typical in young adults, and has been observed in children as young as two years of age.The internal secretions of the pineal gland inhibit the development of the reproductive glands because when it is severely damaged in children, development of the sexual organs and the skeleton are accelerated. Pineal gland calcification is detrimental to its ability to synthesize melatonin but has not been shown to cause sleep problems.
The calcified gland is often seen in skull x-rays.Calcification rates vary widely by country and correlate with an increase in age, with calcification occurring in an estimated 40% of Americans by age seventeen. Calcification of the pineal gland is associated with corpora arenacea, also known as "brain sand".
Tumors of the pineal gland are called pinealomas. These tumors are rare and 50% to 70% are germinomas that arise from sequestered embryonic germ cells. Histologically they are similar to testicular seminomas and ovarian dysgerminomas.
A pineal tumor can compress the superior colliculi and pretectal area of the dorsal midbrain, producing Parinaud's syndrome. Pineal tumors also can cause compression of the cerebral aqueduct, resulting in a noncommunicating hydrocephalus. Other manifestations are the consequence of their pressure effects and consist of visual disturbances, headache, mental deterioration, and sometimes dementia-like behaviour.
These neoplasms are divided into three categories, pineoblastomas, pineocytomas, and mixed tumors, based on their level of differentiation, which, in turn, correlates with their neoplastic aggressiveness.The clinical course of patients with pineocytomas is prolonged, averaging up to several years. The position of these tumors makes them difficult to remove surgically.
The morphology of the pineal gland differs markedly in different pathological conditions. For instance, it is known that its volume is reduced both in obese patients as well as patients with primary insomnia.
Most living vertebrates have pineal glands. It is likely that the common ancestor of all vertebrates had a pair of photosensory organs on the top of its head, similar to the arrangement in modern lampreys.Some extinct Devonian fishes have two parietal foramina in their skulls, suggesting an ancestral bilaterality of parietal eyes. The parietal eye and the pineal gland of living tetrapods are probably the descendants of the left and right parts of this organ, respectively.
During embryonic development, the parietal eye and the pineal organ of modern lizardsand tuataras form together from a pocket formed in the brain ectoderm. The loss of parietal eyes in many living tetrapods is supported by developmental formation of a paired structure that subsequently fuses into a single pineal gland in developing embryos of turtles, snakes, birds, and mammals.
The pineal organs of mammals fall into one of three categories based on shape. Rodents have more structurally complex pineal glands than other mammals.
Crocodilians and some tropical lineages of mammals (some xenarthrans (sloths), pangolins, sirenians (manatees & dugongs), and some marsupials (sugar gliders) have lost both their parietal eye and their pineal organ.Polar mammals, such as walruses and some seals, possess unusually large pineal glands.
All amphibians have a pineal organ, but some frogs and toads also have what is called a "frontal organ", which is essentially a parietal eye.
Pinealocytes in many non-mammalian vertebrates have a strong resemblance to the photoreceptor cells of the eye. Evidence from morphology and developmental biology suggests that pineal cells possess a common evolutionary ancestor with retinal cells.
Pineal cytostructure seems to have evolutionary similarities to the retinal cells of the lateral eyes.Modern birds and reptiles express the phototransducing pigment melanopsin in the pineal gland. Avian pineal glands are thought to act like the suprachiasmatic nucleus in mammals. The structure of the pineal eye in modern lizards and tuatara is analogous to the cornea, lens, and retina of the lateral eyes of vertebrates.
In most vertebrates, exposure to light sets off a chain reaction of enzymatic events within the pineal gland that regulates circadian rhythms.In humans and other mammals, the light signals necessary to set circadian rhythms are sent from the eye through the retinohypothalamic system to the suprachiasmatic nuclei (SCN) and the pineal gland.
The fossilized skulls of many extinct vertebrates have a pineal foramen (opening), which in some cases is larger than that of any living vertebrate. million years old, shows a relatively large parietal eye and pineal gland.Although fossils seldom preserve deep-brain soft anatomy, the brain of the Russian fossil bird Cerebavis cenomanica from Melovatka, about 90
Seventeenth-century philosopher and scientist René Descartes was highly interested in anatomy and physiology. He discussed the pineal gland both in his first book, the Treatise of Man (written before 1637, but only published posthumously 1662/1664), and in his last book, The Passions of the Soul (1649) and he regarded it as "the principal seat of the soul and the place in which all our thoughts are formed."In the Treatise of Man, Descartes described conceptual models of man, namely creatures created by God, which consist of two ingredients, a body and a soul. In the Passions, Descartes split man up into a body and a soul and emphasized that the soul is joined to the whole body by "a certain very small gland situated in the middle of the brain's substance and suspended above the passage through which the spirits in the brain's anterior cavities communicate with those in its posterior cavities". Descartes attached significance to the gland because he believed it to be the only section of the brain to exist as a single part rather than one-half of a pair. Most of Descartes's basic anatomical and physiological assumptions were totally mistaken, not only by modern standards, but also in light of what was already known in his time.
The notion of a "pineal-eye" is central to the philosophy of the French writer Georges Bataille, which is analyzed at length by literary scholar Denis Hollier in his study Against Architecture. In this work Hollier discusses how Bataille uses the concept of a "pineal-eye" as a reference to a blind-spot in Western rationality, and an organ of excess and delirium.This conceptual device is explicit in his surrealist texts, The Jesuve and The Pineal Eye.
In the late 19th century Madame Blavatsky (who founded theosophy) identified the pineal gland with the Hindu concept of the third eye, or the Ajna chakra. This association is still popular today.
Rick Strassman, an author and Clinical Associate Professor of Psychiatry at the University of New Mexico School of Medicine, has theorised that the human pineal gland is capable of producing the hallucinogen N,N-Dimethyltryptamine (DMT) under certain circumstances.In 2013 he and other researchers first reported DMT in the pineal gland microdialysate of rodents.
In the short story "From Beyond" by H. P. Lovecraft, a scientist creates an electronic device that emits a resonance wave, which stimulates an affected person's pineal gland, thereby allowing her or him to perceive planes of existence outside the scope of accepted reality, a translucent, alien environment that overlaps our own recognized reality. It was adapted as a film of the same name in 1986. The 2013 horror film Banshee Chapter is heavily influenced by this short story.
The secretory activity of the pineal gland is only partially understood. Its location deep in the brain suggested to philosophers throughout history that it possesses particular importance. This combination led to its being regarded as a "mystery" gland with mystical, metaphysical, and occult theories surrounding its perceived functions.
The pineal gland was originally believed to be a "vestigial remnant" of a larger organ. In 1917, it was known that extract of cow pineals lightened frog skin. Dermatology professor Aaron B. Lerner and colleagues at Yale University, hoping that a substance from the pineal might be useful in treating skin diseases, isolated and named the hormone melatonin in 1958.The substance did not prove to be helpful as intended, but its discovery helped solve several mysteries such as why removing the rat's pineal accelerated ovary growth, why keeping rats in constant light decreased the weight of their pineals, and why pinealectomy and constant light affect ovary growth to an equal extent; this knowledge gave a boost to the then new field of chronobiology.
The pineal body is labeled in these images.
Melatonin is a hormone that regulates the sleep–wake cycle. It is primarily released by the pineal gland. As a dietary supplement, it is often used for the short-term treatment of trouble sleeping such as from jet lag or shift work. Evidence of benefit, however, is unclear. One review found onset of sleep occurred six minutes faster with use but found no change in total time asleep. It may work as well as the medication ramelteon. It is typically taken by mouth.
A major organ of the endocrine system, the anterior pituitary is the glandular, anterior lobe that together with the posterior lobe makes up the pituitary gland (hypophysis). The anterior pituitary regulates several physiological processes, including stress, growth, reproduction, and lactation. Proper functioning of the anterior pituitary and of the organs it regulates can often be ascertained via blood tests that measure hormone levels.
The third ventricle is one of four connected fluid-filled cavities comprising the ventricular system within the mammalian brain. It is a median cleft in the diencephalon between the two thalami, and is filled with cerebrospinal fluid (CSF).
The suprachiasmatic nucleus or nuclei (SCN) is a tiny region of the brain in the hypothalamus, situated directly above the optic chiasm. It is responsible for controlling circadian rhythms. The neuronal and hormonal activities it generates regulate many different body functions in a 24-hour cycle. The mouse SCN contains approximately 20,000 neurons.
Pinealocytes are the main cells contained in the pineal gland, located behind the third ventricle and between the two hemispheres of the brain. The primary function of the pinealocytes is the secretion of the hormone melatonin, important in the regulation of circadian rhythms. In humans, the suprachiasmatic nucleus of the hypothalamus communicates the message of darkness to the pinealocytes, and as a result, controls the day and night cycle. It has been suggested that pinealocytes are derived from photoreceptor cells. Research has also shown the decline in the number of pinealocytes by way of apoptosis as the age of the organism increases. There are two different types of pinealocytes, type I and type II, which have been classified based on certain properties including shape, presence or absence of infolding of the nuclear envelope, and composition of the cytoplasm.
Melanopsin is a type of photopigment belonging to a larger family of light-sensitive retinal proteins called opsins and encoded by the gene Opn4. In the mammalian retina, there are two additional categories of opsins, both involved in the formation of visual images: rhodopsin and photopsin in the rod and cone photoreceptor cells, respectively.
The epithalamus is a (dorsal) posterior segment of the diencephalon. The diencephalon is a part of the forebrain that also contains the thalamus, the hypothalamus and pituitary gland. The epithalamus includes the habenular nuclei and their interconnecting fibers, the habenular commissure, the stria medullaris and the pineal gland.
Corpora arenacea are calcified structures in the pineal gland and other areas of the brain such as the choroid plexus. Older organisms have numerous corpora arenacea, whose function, if any, is unknown. Concentrations of "brain sand" increase with age, so the pineal gland becomes increasingly visible on X-rays over time, usually by the third or fourth decade. They are sometimes used as anatomical landmarks in radiological examinations.
A pinealoma is a tumor of the pineal gland, a part of the brain that produces melatonin. If a pinealoma destroys the cells of the pineal gland in a child, it can cause precocious puberty.
Circumventricular organs (CVOs) are structures in the brain characterized by their extensive and highly permeable capillaries, unlike those in the rest of the brain where there exists a blood–brain barrier (BBB) at the capillary level. Although the term "circumventricular organs" was originally proposed in 1958 by Austrian anatomist Helmut O. Hofer concerning structures around the brain ventricular system, the penetration of blood-borne dyes into small specific CVO regions was discovered in the early 20th century. The permeable CVOs enabling rapid neurohumoral exchange include the subfornical organ (SFO), the area postrema (AP), the vascular organ of lamina terminalis (VOLT), the median eminence, the pituitary neural lobe, and the pineal gland.
A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some species of fish, amphibian and reptile. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhythmicity and hormone production for thermoregulation.
Melatonin receptors are G protein-coupled receptors (GPCR) which bind melatonin. Three types of melatonin receptors have been cloned. The MT1 (or Mel1A or MTNR1A) and MT2 (or Mel1B or MTNR1B) receptor subtypes are present in humans and other mammals, while an additional melatonin receptor subtype MT3 (or Mel1C or MTNR1C) has been identified in amphibia and birds. The receptors are crucial in the signal cascade of melatonin. In the field of chronobiology, melatonin has been found to be a key player in the synchrony of biological clocks. Melatonin secretion by the pineal gland has circadian rhythmicity regulated by the suprachiasmatic nucleus (SCN) found in the brain. The SCN functions as the timing regulator for melatonin; melatonin then follows a feedback loop to decrease SCN neuronal firing. The receptors MT1 and MT2 control this process. Melatonin receptors are found throughout the body in places such as brain, retina, cardiovascular system, liver and gallbladder, colon, skin, kidney, and many others. In 2019, crystal structures of MT1 and MT2 were reported.
Aralkylamine N-acetyltransferase (AANAT), also known as arylalkylamine N-acetyltransferase or serotonin N-acetyltransferase (SNAT), is an enzyme that is involved in the day/night rhythmic production of melatonin, by modification of serotonin. It is in humans encoded by the ~2.5 kb AANAT gene containing four exons, located on chromosome 17q25. The gene is translated into a 23 kDa large enzyme. It is well conserved through evolution and the human form of the protein is 80% identical to sheep and rat AANAT. It is an acetyl-CoA-dependent enzyme of the GCN5-related family of N-acetyltransferases (GNATs). It may contribute to multifactorial genetic diseases such as altered behavior in sleep/wake cycle and research is on-going with the aim of developing drugs that regulate AANAT function.
N-Acetylserotonin O-methyltransferase, also known as ASMT, is an enzyme which catalyzes the final reaction in melatonin biosynthesis: converting Normelatonin to melatonin. This reaction is embedded in the more general tryptophan metabolism pathway. The enzyme also catalyzes a second reaction in tryptophan metabolism: the conversion of 5-hydroxy-indoleacetate to 5-methoxy-indoleacetate. The other enzyme which catalyzes this reaction is n-acetylserotonin-o-methyltransferase-like-protein.
Melatonin receptor type 1A is a protein that in humans is encoded by the MTNR1A gene.
Light effects on circadian rhythm are the effects that light has on circadian rhythm.
Steven M. Reppert is an American neuroscientist known for his contributions to the fields of chronobiology and neuroethology. His research has focused primarily on the physiological, cellular, and molecular basis of circadian rhythms in mammals and more recently on the navigational mechanisms of migratory monarch butterflies. He was the Higgins Family Professor of Neuroscience at the University of Massachusetts Medical School from 2001 to 2017, and from 2001 to 2013 was the founding chair of the Department of Neurobiology. Reppert stepped down as chair in 2014. He is currently distinguished professor emeritus of neurobiology.
Saniwa is an extinct genus of varanid lizard that lived about 48 million years ago during the Eocene epoch. It is known from well-preserved fossils found in the Bridger and Green River Formations of Wyoming, and evidence indicates Saniwa also lived in Europe. The type species S. ensidens was described in 1870 as the first fossil lizard known from North America. Several other species have since been added, but their validity is uncertain. It is a close relative of Varanus, the genus that includes monitor lizards.
A chronobiotic is an agent that can cause phase adjustment of the body clock. That is, it is a substance capable of therapeutically entraining or re-entraining long-term desynchronized or short-term dissociated circadian rhythms in mammals, or prophylactically preventing their disruption following an environmental insult such as is caused by rapid travel across several time zones. The most widely recognized chronobiotic is the hormone melatonin, secreted at night in both diurnal and nocturnal species.
Brain-specific Homeobox is a protein that in humans is encoded by the BSX gene.
Exogenous melatonin has acute sleepiness-inducing and temperature-lowering effects during 'biological daytime', and when suitably timed (it is most effective around dusk and dawn) it will shift the phase of the human circadian clock (sleep, endogenous melatonin, cortisol) to earlier (advance phase shift) or later (delay phase shift) times.
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