Galanin

2017-11-18T02:22:54Z

171.66.209.9: /* Parental role in mice */

{{Infobox_gene}}
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'''Galanin''' is a [[neuropeptide]] encoded by the ''GAL'' [[gene]],<ref name="pmid7508413"/> that is widely expressed in the brain, spinal cord, and gut of humans as well as other mammals. Galanin signaling occurs through three [[G protein-coupled receptors]].<ref name="receptors"/>

The functional role of galanin remains largely unknown; however, galanin is predominantly involved in the modulation and inhibition of [[action potentials]] in [[neurons]]. Galanin has been implicated in many biologically diverse functions, including: [[nociception]], waking and sleep regulation, cognition, feeding, regulation of mood, regulation of blood pressure, it also has roles in development as well as acting as a [[trophic factor]].<ref name="pmid18500643"/> Galanin neurons in the medial preoptic area of the hypothalamus may govern parental behaviour.<ref name="pmid24828191">{{cite journal | vauthors = Wu Z, Autry AE, Bergan JF, Watabe-Uchida M, Dulac CG | title = Galanin neurons in the medial preoptic area govern parental behaviour | journal = Nature | volume = 509 | issue = 7500 | pages = 325–30 | date = May 2014 | pmid = 24828191 | pmc = 4105201 | doi = 10.1038/nature13307 }}</ref> Galanin is linked to a number of diseases including [[Alzheimer's disease]], [[epilepsy]] as well as [[Major depressive disorder|depression]], [[eating disorders]] and [[cancer]].<ref name="pmid16052044"/><ref name="pmid15944034"/> Galanin appears to have [[neuroprotection|neuroprotective]] activity as its biosynthesis is increased 2-10 fold upon [[axotomy]] in the [[peripheral nervous system]] as well as when seizure activity occurs in the brain. It may also promote [[neurogenesis]].<ref name="receptors"/>

Galanin is predominantly an inhibitory, [[Hyperpolarization (biology)|hyperpolarizing]] neuropeptide<ref name="neuroscience"/> and as such inhibits [[neurotransmitter]] release. Galanin is often co-localized with classical neurotransmitters such as [[acetylcholine]], [[serotonin]], and [[norepinephrine]], and also with other neuromodulators such as [[neuropeptide Y]], [[substance P]], and [[vasoactive intestinal peptide]].<ref name = "acnp"/>

==Discovery==
Galanin was first identified from porcine intestinal extracts in 1978 by Professor Viktor Mutt and colleagues at the Karolinska Institute, Sweden<ref name="MFN"/> using a chemical assay technique that detects peptides according to its C-terminal alanine amide structure. Galanin is so-called because it contains an N-terminal glycine residue and a C-terminal alanine.<ref name="History"/> The structure of galanin was determined in 1983 by the same team, and the [[cDNA]] of galanin was cloned from a rat anterior pituitary library in 1987.<ref name = "MFN"/>

==Tissue distribution==
Galanin is located predominantly in the central nervous system and gastrointestinal tract. Within the central nervous system, highest concentrations are found in the [[hypothalamus]], with lower levels in the [[cerebral cortex|cortex]] and [[brainstem]]. Gastrointestinal galanin is most abundant in the [[duodenum]], with lower concentrations in the stomach, small intestine, and colon.<ref name="pmid2448788"/>

==Structure==

{| class="wikitable" border="1" align="right"
|+Endogenously occurring galanin sequences
|-
! Species !! 1 !! 6 !! 11 !! 16 !! 21 !! 26 !
|-
! Pig
| G W T L N || S A G Y L || L G P H A || I D N H R || S F H D K || Y G L A *
|-
! Human
| G W T L N || S A G Y L || L G P H A || '''V''' '''G''' N H R || S F '''S''' D K || '''N''' G L '''T''' S **
|-
! Cow
| G W T L N || S A G Y L || L G P H A || '''L''' D '''S''' H R || S F '''Q''' D K || '''H''' G L A *
|-
! Rat
| G W T L N || S A G Y L || L G P H A || I D N H R || S F '''S''' D K || '''H''' G L '''T'''*
|-
| colspan=7 align=center| <small>* C-terminal amide ** C-terminal free acid</small>
|}

Galanin is a [[peptide]] consisting of a chain of 29 [[amino acids]] (30 amino acids in humans) produced from the cleavage of a 123-amino acid protein known as prepro galanin, which is encoded by the ''GAL'' gene.<ref name="pmid7508413"/> The sequence of this gene is highly conserved among mammals, showing over 85% [[homology (biology)|homology]] between rat, mouse, porcine, bovine, and human sequences.<ref name = "acnp"/> In these animal forms, the first 15 amino acids from the [[N-terminus]] are identical, but amino acids differ at several positions on the [[C-terminal]] end of the protein.

These slight differences in protein structure have far-reaching implications on their function. For example, porcine and rat galanin inhibit glucose-induced [[insulin]] secretion in rats and dogs but have no effect on insulin secretion in humans. This demonstrates that it is essential to study the effects of galanin and other regulatory peptides in their autologous species.<ref name="pmid1710578"/>

The galanin family of protein consists of four proteins, of which GAL was the first to be identified. The second was galanin message-associated protein (GMAP), a 59- or 60-amino acid peptide also formed from the cleavage of prepro galanin.<ref name="History"/> The other two peptides, [[galanin-like peptide]] (GALP) and alarin, were identified relatively recently and are both encoded for in the same gene, the prepro GALP gene. GALP and alarin are produced by different post-transcriptional [[RNA splicing|splicing]] of this gene.<ref name="overview"/>

{| align="center"
|{{Infobox protein family
| Symbol = Galanin
| Name = Galanin
| image =
| width =
| caption =
| Pfam = PF01296
| InterPro = IPR008174
| SMART =
| PROSITE = PDOC00673
| SCOP =
| TCDB =
| OPM family =
| OPM protein =
| PDB =
}}
|
{{Infobox protein family
| Symbol = GMAP
| Name = Galanin message associated peptide (GMAP)
| image =
| width =
| caption =
| Pfam = PF06540
| Pfam_clan =
| InterPro = IPR013068
| SMART =
| PROSITE =
| MEROPS =
| SCOP =
| TCDB =
| OPM family =
| OPM protein =
| CAZy =
| CDD =
}}
|}

==Receptors==

Galanin signalling occurs through three classes of receptors, [[GALR1]], [[GALR2]], and [[GALR3]], which are all part of the [[G protein-coupled receptor]] (GPCR) superfamily. Galanin receptors are expressed in the [[central nervous system]], in the [[pancreas]], and on [[solid tumour]]s. The level of expression of the different receptors varies at each location, and this distribution changes after injury to neurons.<ref name="receptors"/> Experiments into the function of the receptor subtypes involve mostly [[gene knockout|genetic knockout]] mice. The location of the receptor and the combination of receptors that are inhibited or stimulated heavily affect the outcome of galanin signalling.<ref name="receptors"/>

==Clinical characteristics==

===Alzheimer's disease===
One of the [[pathology|pathological]] features of the brain in the later stages of [[Alzheimer's disease]] is the presence of overgrown GAL-containing fibres innervating the surviving [[cholinergic]] neurons.<ref name="pmid18500641"/> Another feature is an increase in the expression of GAL and GAL receptors, in which increases of up to 200% have been observed in postmortem brains of Alzheimer's patients.<ref name="receptors"/><ref name="overview"/> The cause and role of this increase is poorly understood.<ref name="pmid18500641"/><ref name="pmid14993421"/>

It has been suggested that the hyper-innervation acts to promote the death of these neurons and that the inhibitory effect of galanin on cholinergic neurons worsened the degeneration of [[cognition|cognitive]] function in patients by decreasing the amount of [[acetylcholine]] available to these neurons.<ref name= "receptors"/><ref name="pmid18500641"/>

A second hypothesis has been generated based on data that suggest GAL is involved in protecting the hippocampus from [[excitotoxic]] damage and the neurons in the cholinergic basal forebrain from [[amyloid]] toxicity.<ref name="pmid16246567"/> It is interesting to note that studies of gene expression of CBF tissue suggests that the hyperinnervation of cholinergic neurons by GAL up regulates the [[Transcription (genetics)|transcription]] of factors that promote neuron function and survival. It is still unclear as to whether galanin acts to protect cholinergic neurons and promote their firing or whether it worsens the symptoms of this disease.

===Epilepsy===
Galanin in the [[hippocampus]] is an inhibitor of [[glutamate]] but not of [[GABA]]. This means that galanin is capable of increasing the [[seizure threshold]] <ref name= "receptors"/> and, therefore, is expected to act as an [[anticonvulsant]]. To be specific, GalR1 has been linked to the suppression of spontaneous seizures.<ref name="pmid15350653"/><ref name="pmid19199479">{{cite journal | vauthors = Zhang L, Robertson CR, Green BR, Pruess TH, White HS, Bulaj G | title = Structural requirements for a lipoamino acid in modulating the anticonvulsant activities of systemically active galanin analogues | journal = Journal of Medicinal Chemistry | volume = 52 | issue = 5 | pages = 1310–6 | date = Mar 2009 | pmid = 19199479 | pmc = 2765488 | doi = 10.1021/jm801397w }}</ref> An agonist antiepileptic drug candidate is NAX 5055.<ref name="pmid19053761">{{cite journal | vauthors = Bulaj G, Green BR, Lee HK, Robertson CR, White K, Zhang L, Sochanska M, Flynn SP, Scholl EA, Pruess TH, Smith MD, White HS | title = Design, synthesis, and characterization of high-affinity, systemically-active galanin analogues with potent anticonvulsant activities | journal = Journal of Medicinal Chemistry | volume = 51 | issue = 24 | pages = 8038–47 | date = Dec 2008 | pmid = 19053761 | doi = 10.1021/jm801088x }}</ref><ref name="pmid19332332">{{cite journal | vauthors = White HS, Scholl EA, Klein BD, Flynn SP, Pruess TH, Green BR, Zhang L, Bulaj G | title = Developing novel antiepileptic drugs: characterization of NAX 5055, a systemically-active galanin analog, in epilepsy models | journal = Neurotherapeutics | volume = 6 | issue = 2 | pages = 372–80 | date = Apr 2009 | pmid = 19332332 | pmc = 4402707 | doi = 10.1016/j.nurt.2009.01.001 }}</ref>

===In development===
It has been shown that galanin plays a role in the control of the early post-natal [[neural development]] of the [[dorsal root ganglion]] (DRG).<ref name="MFN"/> Galanin-mutant animals show a 13% decrease in the number of adult DRG cells as well as a 24% decrease in the percentage of cells expressing [[substance P]]. This suggests that the cell loss by [[apoptosis]] that usually occurs in the developing DRG is regulated by galanin and that the absence of galanin results in an increase in the number of cells that die.

===After injury===
''In vitro'' experiments show that DRG cells removed from galanin mutants have impaired abilities to extend [[neurites]] in culture, in that the number of cells producing neurites is decreased by a third and the mean length of these processes was halved when compared to wild-type controls. ''In vivo'', many of the actions of galanin in the brain after an injury are similar to those observed in the developing DRG. Adult mutant animals have been shown to be 35% less capable of regenerating the [[sciatic nerve]] after crush injury, which is linked to long-term functional problems.

===Parental role in mice===
A report has indicated that Galanin-expressing neurons in the medial preoptic area of the brain are responsible for regulating aggression towards pups by male mice. <ref>http://phys.org/news/2014-05-chemicals-neurons-responsible-parental-mice.html</ref>

== See also ==
* [[Galanin receptor]]

== References ==
{{Reflist|2|refs=

<ref name="pmid7508413">{{cite journal | vauthors = Evans H, Baumgartner M, Shine J, Herzog H | title = Genomic organization and localization of the gene encoding human preprogalanin | journal = Genomics | volume = 18 | issue = 3 | pages = 473–7 |date=December 1993 | pmid = 7508413 | doi =10.1016/S0888-7543(11)80002-9 | url = | issn = }}</ref>

<ref name="receptors">{{cite journal | vauthors = Mitsukawa K, Lu X, Bartfai T | title = Galanin, galanin receptors and drug targets | journal = Cell. Mol. Life Sci. | volume = 65 | issue = 12 | pages = 1796–805 |date=June 2008 | pmid = 18500647 | doi = 10.1007/s00018-008-8153-8 | url = | issn = }}</ref>

<ref name="neuroscience">{{cite journal | vauthors = Ito M | title = Functional roles of neuropeptides in cerebellar circuits | journal = Neuroscience | volume = 162 | issue = 3 | pages = 666–72 |date=September 2009 | pmid = 19361475 | doi = 10.1016/j.neuroscience.2009.01.019 | url = | issn = }}</ref>

<ref name="acnp">{{cite web|author=Bartfai, T., |title=Galanin – A neuropeptide with important central nervous system actions |year=2000 |url=http://www.acnp.org/g4/GN401000054/CH054.html |accessdate=November 19, 2009 |deadurl=yes |archiveurl=https://web.archive.org/web/20101202155619/http://www.acnp.org/g4/GN401000054/CH054.html |archivedate=December 2, 2010 |df= }}</ref>

<ref name="MFN">{{cite journal | vauthors = Wynick D, Thompson SW, McMahon SB | title = The role of galanin as a multi-functional neuropeptide in the nervous system | journal = Current Opinion in Pharmacology | volume = 1 | issue = 1 | pages = 73–7 |date=February 2001 | pmid = 11712539 | doi = 10.1016/S1471-4892(01)00006-6| url = | issn = }}</ref>

<ref name="History">{{cite journal | vauthors = Hökfelt T, Tatemoto K | title = Galanin--25 years with a multitalented neuropeptide | journal = Cell. Mol. Life Sci. | volume = 65 | issue = 12 | pages = 1793–5 |date=June 2008 | pmid = 18500648 | doi = 10.1007/s00018-008-8152-9 | url = | issn = }}</ref>

<ref name="pmid1710578">{{cite journal | vauthors = Bersani M, Johnsen AH, Højrup P, Dunning BE, Andreasen JJ, Holst JJ | title = Human galanin: primary structure and identification of two molecular forms | journal = FEBS Lett. | volume = 283 | issue = 2 | pages = 189–94 |date=June 1991 | pmid = 1710578 | doi = 10.1016/0014-5793(91)80585-Q| url = | issn = }}</ref>

<ref name="overview">{{cite journal | vauthors = Lang R, Gundlach AL, Kofler B | title = The galanin peptide family: receptor pharmacology, pleiotropic biological actions, and implications in health and disease | journal = Pharmacol. Ther. | volume = 115 | issue = 2 | pages = 177–207 |date=August 2007 | pmid = 17604107 | doi = 10.1016/j.pharmthera.2007.05.009 | url = | issn = }}</ref>

<ref name="pmid14993421">{{cite journal | vauthors = Counts SE, Perez SE, Ginsberg SD, De Lacalle S, Mufson EJ | title = Galanin in Alzheimer disease | journal = Mol. Interv. | volume = 3 | issue = 3 | pages = 137–56 |date=May 2003 | pmid = 14993421 | doi = 10.1124/mi.3.3.137 | url = | issn = }}</ref>

<ref name="pmid18500641">{{cite journal | vauthors = Counts SE, Perez SE, Mufson EJ | title = Galanin in Alzheimer's disease: neuroinhibitory or neuroprotective? | journal = Cell. Mol. Life Sci. | volume = 65 | issue = 12 | pages = 1842–53 |date=June 2008 | pmid = 18500641 | pmc = 2911017 | doi = 10.1007/s00018-008-8159-2 | url = | issn = }}</ref>

<ref name="pmid16246567">{{cite journal | vauthors = Ding X, MacTavish D, Kar S, Jhamandas JH | title = Galanin attenuates beta-amyloid (Abeta) toxicity in rat cholinergic [[basal forebrain]] neurons | journal = Neurobiol. Dis. | volume = 21 | issue = 2 | pages = 413–20 |date=February 2006 | pmid = 16246567 | doi = 10.1016/j.nbd.2005.08.016 | url = | issn = }}</ref>

<ref name="pmid15350653">{{cite journal | vauthors = Mazarati A, Lu X, Shinmei S, Badie-Mahdavi H, Bartfai T | title = Patterns of seizures, hippocampal injury and neurogenesis in three models of status epilepticus in galanin receptor type 1 (GalR1) knockout mice | journal = Neuroscience | volume = 128 | issue = 2 | pages = 431–41 | year = 2004 | pmid = 15350653 | pmc = 1360211 | doi = 10.1016/j.neuroscience.2004.06.052 | url = | issn = }}</ref>

<ref name="pmid15944034">{{cite journal | vauthors = Berger A, Santic R, Hauser-Kronberger C, Schilling FH, Kogner P, Ratschek M, Gamper A, Jones N, Sperl W, Kofler B | title = Galanin and galanin receptors in human cancers | journal = Neuropeptides | volume = 39 | issue = 3 | pages = 353–9 |date=June 2005 | pmid = 15944034 | doi = 10.1016/j.npep.2004.12.016 | url = | issn = }}</ref>

<ref name="pmid16052044">{{cite journal | vauthors = Lundström L, Elmquist A, Bartfai T, Langel U | title = Galanin and its receptors in neurological disorders | journal = Neuromolecular Med. | volume = 7 | issue = 1–2 | pages = 157–80 | year = 2005 | pmid = 16052044 | doi = 10.1385/NMM:7:1-2:157 | url = | issn = }}</ref>

<ref name="pmid18500643">{{cite journal | vauthors = Mechenthaler I | title = Galanin and the neuroendocrine axes | journal = Cell. Mol. Life Sci. | volume = 65 | issue = 12 | pages = 1826–35 |date=June 2008 | pmid = 18500643 | doi = 10.1007/s00018-008-8157-4 | url = | issn = }}</ref>

<ref name="pmid2448788">{{cite journal | vauthors = Kaplan LM, Spindel ER, Isselbacher KJ, Chin WW | title = Tissue-specific expression of the rat galanin gene | journal = Proc. Natl. Acad. Sci. U.S.A. | volume = 85 | issue = 4 | pages = 1065–9 |date=February 1988 | pmid = 2448788 | pmc = 279702 | doi = 10.1073/pnas.85.4.1065| url = | issn = }}</ref>

}}

== Further reading ==
{{Refbegin| 2}}
* {{cite journal | vauthors = Vrontakis ME | title = Galanin: a biologically active peptide | journal = Current Drug Targets. CNS and Neurological Disorders | volume = 1 | issue = 6 | pages = 531–41 | date = Dec 2002 | pmid = 12769595 | doi = 10.2174/1568007023338914 }}
* {{cite journal | vauthors = Mufson EJ, Counts SE, Perez SE, Binder L | title = Galanin plasticity in the cholinergic basal forebrain in Alzheimer's disease and transgenic mice | journal = Neuropeptides | volume = 39 | issue = 3 | pages = 233–7 | date = Jun 2005 | pmid = 15893372 | doi = 10.1016/j.npep.2004.12.005 }}
* {{cite journal | vauthors = Robinson JK, Bartfai T, Langel U | title = Galanin/GALP receptors and CNS homeostatic processes | journal = CNS & Neurological Disorders Drug Targets | volume = 5 | issue = 3 | pages = 327–34 | date = Jun 2006 | pmid = 16787232 | doi = 10.2174/187152706777452281 }}
* {{cite journal | vauthors = McKnight GL, Karlsen AE, Kowalyk S, Mathewes SL, Sheppard PO, O'Hara PJ, Taborsky GJ | title = Sequence of human galanin and its inhibition of glucose-stimulated insulin secretion from RIN cells | journal = Diabetes | volume = 41 | issue = 1 | pages = 82–7 | date = Jan 1992 | pmid = 1370155 | doi = 10.2337/diabetes.41.1.82 }}
* {{cite journal | vauthors = Gai WP, Geffen LB, Blessing WW | title = Galanin immunoreactive neurons in the human hypothalamus: colocalization with vasopressin-containing neurons | journal = The Journal of Comparative Neurology | volume = 298 | issue = 3 | pages = 265–80 | date = Aug 1990 | pmid = 1698834 | doi = 10.1002/cne.902980302 }}
* {{cite journal | vauthors = Burleigh DE, Furness JB | title = Distribution and actions of galanin and vasoactive intestinal peptide in the human colon | journal = Neuropeptides | volume = 16 | issue = 2 | pages = 77–82 | date = Jun 1990 | pmid = 1701228 | doi = 10.1016/0143-4179(90)90115-F }}
* {{cite journal | vauthors = Fried G, Meister B, Rådestad A | title = Peptide-containing nerves in the human pregnant uterine cervix: an immunohistochemical study exploring the effect of RU 486 (mifepristone) | journal = Human Reproduction | volume = 5 | issue = 7 | pages = 870–6 | date = Oct 1990 | pmid = 1702449 | doi = }}
* {{cite journal | vauthors = Hyde JF, Engle MG, Maley BE | title = Colocalization of galanin and prolactin within secretory granules of anterior pituitary cells in estrogen-treated Fischer 344 rats | journal = Endocrinology | volume = 129 | issue = 1 | pages = 270–6 | date = Jul 1991 | pmid = 1711463 | doi = 10.1210/endo-129-1-270 }}
* {{cite journal | vauthors = Bennet WM, Hill SF, Ghatei MA, Bloom SR | title = Galanin in the normal human pituitary and brain and in pituitary adenomas | journal = The Journal of Endocrinology | volume = 130 | issue = 3 | pages = 463–7 | date = Sep 1991 | pmid = 1719117 | doi = 10.1677/joe.0.1300463 }}
* {{cite journal | vauthors = Schmidt WE, Kratzin H, Eckart K, Drevs D, Mundkowski G, Clemens A, Katsoulis S, Schäfer H, Gallwitz B, Creutzfeldt W | title = Isolation and primary structure of pituitary human galanin, a 30-residue nonamidated neuropeptide | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 88 | issue = 24 | pages = 11435–9 | date = Dec 1991 | pmid = 1722333 | pmc = 53150 | doi = 10.1073/pnas.88.24.11435 }}
* {{cite journal | vauthors = Bauer FE, Christofides ND, Hacker GW, Blank MA, Polak JM, Bloom SR | title = Distribution of galanin immunoreactivity in the genitourinary tract of man and rat | journal = Peptides | volume = 7 | issue = 1 | pages = 5–10 | year = 1986 | pmid = 2423990 | doi = 10.1016/0196-9781(86)90052-5 }}
* {{cite journal | vauthors = Bauer FE, Adrian TE, Christofides ND, Ferri GL, Yanaihara N, Polak JM, Bloom SR | title = Distribution and molecular heterogeneity of galanin in human, pig, guinea pig, and rat gastrointestinal tracts | journal = Gastroenterology | volume = 91 | issue = 4 | pages = 877–83 | date = Oct 1986 | pmid = 2427385 | doi = }}
* {{cite journal | vauthors = Tainio H, Vaalasti A, Rechardt L | title = The distribution of substance P-, CGRP-, galanin- and ANP-like immunoreactive nerves in human sweat glands | journal = The Histochemical Journal | volume = 19 | issue = 6–7 | pages = 375–80 | year = 1987 | pmid = 2444569 | doi = 10.1007/BF01680455 }}
* {{cite journal | vauthors = Maggi CA, Santicioli P, Patacchini R, Turini D, Barbanti G, Beneforti P, Giuliani S, Meli A | title = Galanin: a potent modulator of excitatory neurotransmission in the human urinary bladder | journal = European Journal of Pharmacology | volume = 143 | issue = 1 | pages = 135–7 | date = Nov 1987 | pmid = 2446889 | doi = 10.1016/0014-2999(87)90744-8 }}
* {{cite journal | vauthors = Marti E, Gibson SJ, Polak JM, Facer P, Springall DR, Van Aswegen G, Aitchison M, Koltzenburg M | title = Ontogeny of peptide- and amine-containing neurones in motor, sensory, and autonomic regions of rat and human spinal cord, dorsal root ganglia, and rat skin | journal = The Journal of Comparative Neurology | volume = 266 | issue = 3 | pages = 332–59 | date = Dec 1987 | pmid = 2447134 | doi = 10.1002/cne.902660304 }}
* {{cite journal | vauthors = Beal MF, Clevens RA, Chattha GK, MacGarvey UM, Mazurek MF, Gabriel SM | title = Galanin-like immunoreactivity is unchanged in Alzheimer's disease and Parkinson's disease dementia cerebral cortex | journal = Journal of Neurochemistry | volume = 51 | issue = 6 | pages = 1935–41 | date = Dec 1988 | pmid = 2460590 | doi = 10.1111/j.1471-4159.1988.tb01181.x }}
* {{cite journal | vauthors = Berrettini WH, Kaye WH, Sunderland T, May C, Gwirtsman HE, Mellow A, Albright A | title = Galanin immunoreactivity in human CSF: studies in eating disorders and Alzheimer's disease | journal = Neuropsychobiology | volume = 19 | issue = 2 | pages = 64–8 | year = 1989 | pmid = 2465504 | doi = 10.1159/000118436 }}
{{Refend}}

== External links ==
* {{MeshName|Galanin}}

{{Neuropeptides}}
{{Neuropeptidergics}}

[[Category:Neuropeptides]]

ARGLU1

2017-11-02T16:58:21Z

171.66.209.9:

{{Infobox_gene}}
'''Arginine and glutamate-rich protein 1''' is a [[protein]] that in humans is encoded by the ''ARGLU1'' [[gene]] located at [[chromosome 13|13q33.3]].<ref name="entrez">{{cite web | title = Entrez Gene: ARGLU1 arginine and glutamate rich 1| url = https://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=55082| accessdate = }}</ref>

The protein product of this gene has been proposed as a [[MED1]]-interacting protein required for [[estrogen]]-dependent [[gene transcription]] and [[breast cancer]] [[cell growth]].<ref name="pmid21454576">{{cite journal |vauthors=Zhang D, Jiang P, Xu Q, Zhang X | title = Arginine and glutamate-rich 1 (ARGLU1) interacts with mediator subunit 1 (MED1) and is required for estrogen receptor-mediated gene transcription and breast cancer cell growth | journal = J. Biol. Chem. | volume = 286 | issue = 20 | pages = 17746–54 |date=May 2011 | pmid = 21454576 | pmc = 3093850 | doi = 10.1074/jbc.M110.206029 }}</ref>

The ARGLU1 gene expresses at least three distinct RNA splice isoforms - a fully spliced isoform coding for the protein, an isoform containing a retained intron that is detained in the nucleus, and an isoform containing an alternative exon that targets the transcript for nonsense mediated decay. Furthermore, ARGLU1 contains a long, highly evolutionarily conserved sequence known as an [[Ultra-conserved_element|Ultraconserved Element (UCE)]] that is within the retained intron and overlaps the alternative exon. <ref name="pmid27899669">{{cite journal | title = An Ultraconserved Element (UCE) controls homeostatic splicing of ARGLU1 mRNA. | pmid = 27899669 | pmc = 5389617 | doi = 10.1093/nar/gkw1140 }}</ref>

==References==
{{reflist}}

[[Category:Genes on human chromosome 13]]

==External links==
* {{UCSC gene info|ARGLU1}}

wikidoc - User contributions [en]

Galanin

ARGLU1