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'''Galanin''' is a ] that is encoded by the ''GAL'' ],<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 ].<ref name="receptors"/> '''Galanin''' is a ] encoded by the ''GAL'' ],<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 ].<ref name="receptors"/>


The functional role of galanin, along with most other neuropeptides, remains largely unknown; however, galanin is predominately involved in the modulation and inhibition of ] in ]. Galanin has been implicated in many biologically diverse functions, including: ], 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 ].<ref name="pmid18500643"/> Galanin is linked to a number of diseases including ], ] as well as ], ] and ].<ref name="pmid16052044"/><ref name="pmid15944034"/> Galanin appears to have ] activity as its biosynthesis is increased 2-10 fold upon ] in the ] as well as when seizure activity occurs in the brain. It may also promote ].<ref name="receptors"/> Much of galanin's functional role is still undiscovered. Galanin is closely involved in the modulation and inhibition of ] in ]. Galanin has been implicated in many biologically diverse functions, including: ], 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 ].<ref name="pmid18500643"/> Galanin neurons in the ] of the hypothalamus may govern parental behaviour.<ref name="pmid24828191"/> Galanin is linked to a number of diseases including ], ] as well as ], ], ], and ].<ref name="pmid16052044"/><ref name="pmid15944034"/> Galanin appears to have ] activity as its biosynthesis is increased 2-10 fold upon ] in the ] as well as when seizure activity occurs in the brain. It may also promote ].<ref name="receptors"/>


Galanin is predominantly an inhibitory, ] neuropeptide<ref name="neuroscience"/> and as such inhibits ] release. Galanin is often co-localized with classical neurotransmitters such as ], ] and ] and also with other neuromodulators such as ], ] and ].<ref name = "acnp"/> Galanin is predominantly an inhibitory, ] neuropeptide<ref name="neuroscience"/> and as such inhibits ] release. Galanin is often co-localized with classical neurotransmitters such as ], ], and ], and also with other neuromodulators such as ], ], and ].<ref name = "acnp"/>


==Discovery== ==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 its ] of galanin was cloned from rat anterior pituitary library in 1987.<ref name = "MFN"/> Galanin was first identified from porcine intestinal extracts in 1978 by Professor Viktor Mutt and colleagues at the ], 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 ] of galanin was cloned from a rat ] library in 1987.<ref name = "MFN"/>


==Tissue distribution== ==Tissue distribution==
Galanin is located predominantly in the central nervous system and ]. Within the central nervous system, highest concentrations are found in the ], with lower levels in the ] and ]. In the hypothalamus, it is for example found in the ] where it has sleep-promoting function. Within the brain, galanin has also been found in the ventral ] and ].<ref name="Kuteeva_2010">{{cite journal | vauthors = Kuteeva E, Hökfelt T, Wardi T, Ogren SO | journal = Experientia Supplementum (2012) | series = Experientia Supplementum | title = Galanin, galanin receptor subtypes and depression-like behaviour | volume = 102 | pages = 163–181 | date = 2010 | pmid = 21299068 | doi = 10.1007/978-3-0346-0228-0_12 | publisher = Springer | isbn = 978-3-0346-0227-3 | veditors = Hökfelt T }}</ref> Along with this, the immune reaction of galanin in the brain is centered in the hypothalamopituitary.<ref>{{cite journal | vauthors = Ch'ng JL, Christofides ND, Anand P, Gibson SJ, Allen YS, Su HC, Tatemoto K, Morrison JF, Polak JM, Bloom SR | display-authors = 6 | title = Distribution of galanin immunoreactivity in the central nervous system and the responses of galanin-containing neuronal pathways to injury | journal = Neuroscience | volume = 16 | issue = 2 | pages = 343–354 | date = October 1985 | pmid = 2417156 | doi = 10.1016/0306-4522(85)90007-7 | s2cid = 32774212 }}</ref> Gastrointestinal galanin is most abundant in the ], with lower concentrations in the stomach, small intestine, and colon.<ref name="pmid2448788"/> Galanin is also expressed in the skin where is serves anti-inflammatory functions.<ref name="Bauer_2010">{{cite journal | vauthors = Bauer JW, Lang R, Jakab M, Kofler B | journal = Experientia Supplementum (2012) | series = Experientia Supplementum | title = Galanin family of peptides in skin function | volume = 102 | pages = 51–59 | date = 2010 | pmid = 21299061 | doi = 10.1007/978-3-0346-0228-0_5 | publisher = Springer | isbn = 978-3-0346-0227-3 | veditors = Hökfelt T }}</ref> Specifically, it has been found in ]s, ]s, and around blood vessels.<ref name="Bauer_2010" /> Galanin has been found in ]s.<ref name="Mitsukawa_2010">{{cite journal | vauthors = Mitsukawa K, Lu X, Bartfai T | journal = Experientia Supplementum (2012) | series = Experientia Supplementum | title = Galanin, galanin receptors, and drug targets | volume = 102 | pages = 7–23 | date = 2010 | pmid = 21299058 | doi = 10.1007/978-3-0346-0228-0_2 | publisher = Springer | isbn = 978-3-0346-0227-3 | veditors = Hökfelt T }}</ref> Within gastric cancer cells, galanin has been found to have a ] role, but hypermethylation has been shown to stop its tumor suppressive properties.<ref>{{cite journal | vauthors = Yoon D, Bae K, Lee MK, Kim JH, Yoon KA | title = Galanin is an epigenetically silenced tumor suppressor gene in gastric cancer cells | journal = PLOS ONE | volume = 13 | issue = 2 | pages = e0193275 | date = 2018-02-20 | pmid = 29462183 | pmc = 5819827 | doi = 10.1371/journal.pone.0193275 | veditors = Suzuki H | doi-access = free | bibcode = 2018PLoSO..1393275Y }}</ref>
Galanin is located predominantly in the central nervous system and gastrointestinal tract. Within the central nervous system, highest concentrations are found in the ], with lower levels in the ] and ]. Gastrointestinal galanin is most abundant in the ], with lower concentrations in the stomach, small intestine, and colon.<ref name="pmid2448788"/>


==Structure== == Structure ==


{| class="wikitable" border="1" align="left" {| class="wikitable" border="1" align="right"
|+Endogenously occurring galanin sequences |+Endogenously occurring galanin sequences
|- |-
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| 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 * | 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 *
|- |-
! Man ! 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 ** | 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 **
|- |-
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| 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'''* | 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> | colspan=7 align=center| <small>* C-terminal amide ** C-terminal free acid</small>
|} |}


Galanin is a ] consisting of a chain of 29 ] (30 amino acids in humans) produced from the cleavage of a 123 amino acid known as preprogalanin, which is encoded by the ''GAL'' gene.<ref name="pmid7508413"/> The sequence of this gene is highly conserved amongst mammals, showing over 85% ] between rat, mouse, porcine, bovine and human sequences.<ref name = "acnp"/> In these animal forms, the first 15 amino acids from the ] are identical but amino acids differ at several positions on the ] end of the protein. Galanin is a ] consisting of a chain of 29 ] (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% ] between rat, mouse, porcine, bovine, and human sequences.<ref name = "acnp"/> In these animal forms, the first 15 amino acids from the ] are identical, but amino acids differ at several positions on the ] 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 ] 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"/> These slight differences in protein structure have far-reaching implications on their function. For example, porcine and rat galanin inhibit glucose-induced ] 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 preprogalanin.<ref name="History"/> The other two peptides, ] (GALP) and alarin, were identified relatively recently and are both encoded for in the same gene, the preproGALP gene. GALP and alarin are produced by different post-] ] of this gene.<ref name="overview"/> 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, ] (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 ] of this gene.<ref name="overview"/>


{| align="center"
{|
|{{Infobox protein family |{{Infobox protein family
| Symbol = Galanin | Symbol = Galanin
Line 89: Line 98:
| OPM protein = | OPM protein =
| PDB = | 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 =
}} }}
|} |}
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==Receptors== ==Receptors==


Galanin signalling occurs through three classes of receptors, ], ] and ], which are all part of the ] (GPCR) super family. Galanin receptors are expressed in the ], in the ] as well as on ]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 mostly involve ] mice. The location of the receptor and the combination of receptors that are inhibited or stimulated heavily affects the outcome of galanin signalling.<ref name="receptors"/> Galanin signalling occurs through three classes of receptors, ], ], and ], which are all part of the ] (GPCR) superfamily. Galanin receptors are expressed in the ], in the ], and on ]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 ] 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== ==Clinical characteristics==

===Alzheimer’s Disease===
===Appetite===
One of the ] features of the brain in the later stages of ] is the presence of overgrown GAL-containing fibres innervating the surviving ] 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 post-mortem 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"/>
Injections of galanin into the ] or directly into the hypothalamus creates the urge to feed, with a preference for eating fats.<ref name="Mitsukawa_2010" /> Galanin also regulates glucose metabolism and can potentially alleviate symptoms of ] due to its interaction with insulin resistance.<ref name="Fang_2020">{{cite journal | vauthors = Fang P, Yu M, Shi M, Bo P, Zhang Z | title = Galanin peptide family regulation of glucose metabolism | journal = Frontiers in Neuroendocrinology | volume = 56 | pages = 100801 | date = January 2020 | pmid = 31705911 | doi = 10.1016/j.yfrne.2019.100801 | doi-access = free }}</ref> Galanin is an inhibitor of pancreatic secretion of insulin.<ref name="Mitsukawa_2010" />

=== Addiction ===
Galanin plays a role in addiction regulation.<ref name="Genders_2020">{{cite journal | vauthors = Genders SG, Scheller KJ, Djouma E | title = Neuropeptide modulation of addiction: Focus on galanin | journal = Neuroscience and Biobehavioral Reviews | volume = 110 | pages = 133–149 | date = March 2020 | pmid = 29949733 | doi = 10.1016/j.neubiorev.2018.06.021 | s2cid = 49486365 }}</ref> It is involved in repeated alcohol intake.<ref name="Mitsukawa_2010" /> Along with addiction to alcohol, galanin has been shown to play a role in addiction to ] and opiates.<ref name="Genders_2020" />

===Alzheimer's disease===
One of the ] features of the brain in the later stages of ] is the presence of overgrown GAL-containing fibres innervating the surviving ] 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 ] function in patients by decreasing the amount of ] available to these neurons.<ref name= "receptors"/><ref name="pmid18500641"/> 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 ] function in patients by decreasing the amount of ] 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 ] damage and the neurons in the cholinergic basal forebrain from ] toxicity.<ref name="pmid16246567"/> Interestingly, studies of gene expression of CBF tissue suggests that the hyperinnervation of cholinergic neurons by GAL up regulates the ] of factors that promote neuron function and survival. It is still unclear whether galanin acts to protect cholinergic neurons and promote their firing or whether it worsens the symptoms of this disease. A second hypothesis has been generated based on data that suggest GAL is involved in protecting the hippocampus from ] damage and the neurons in the cholinergic ] from ] toxicity.<ref name="pmid16246567"/>

=== Cognitive performance ===
Galanin participates in cognitive performance and has been shown to weaken learning and cognition.<ref name="Mitsukawa_2010" />

=== Depression ===
] and ], two neurotransmitters involved in depression, are both co-expressed and modulated by galanin, suggesting that galanin plays a role in the regulation of depression.<ref name="Kuteeva_2010" /> Stimulation of the Gal1 and Gal3 receptors result in depression-like behaviors, whereas stimulation of the Gal2 receptor results in reduced depression-like behaviors.<ref name="Kuteeva_2010" /> Currently, one of the potential mechanisms for this is that galanin stimulates the hypothalamus-pituitary-adrenal axis, which leads to an increase in ] secretion.<ref name="Kuteeva_2010" /> Increased levels of glucocorticoid hormones is common in those who suffer from depression.<ref>{{cite journal | vauthors = Anacker C, Zunszain PA, Carvalho LA, Pariante CM | title = The glucocorticoid receptor: pivot of depression and of antidepressant treatment? | journal = Psychoneuroendocrinology | volume = 36 | issue = 3 | pages = 415–425 | date = April 2011 | pmid = 20399565 | pmc = 3513407 | doi = 10.1016/j.psyneuen.2010.03.007 }}</ref>

===Endocrine===
Galanin inhibits the secretion of ] and ] and stimulates the secretion of ], ], ], ], ], foliculotropin, ], hypothalamic ], and ].<ref>{{cite journal | vauthors = Zdrojewicz Z, Sowińska E, Sztuka-Pietkiewicz A | title = | journal = Endokrynologia, Diabetologia I Choroby Przemiany Materii Wieku Rozwojowego | volume = 6 | issue = 2 | pages = 129–134 | date = 2000 | pmid = 12818074 | url = https://pubmed.ncbi.nlm.nih.gov/12818074/ | access-date = 28 January 2023 }}</ref>


===Epilepsy=== ===Epilepsy===
Galanin in the ] is an inhibitor of ] but not ] and as such is capable of increasing the ] <ref name= "receptors"/> and therefore is expected to act as an ]. Specifically, GalR1 has been linked to the suppression of spontaneous seizures.<ref name="pmid15350653"/><ref name="pmid19199479">{{cite journal |author=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|year=2009 |month=February |pmid=19199479 |doi=10.1021/jm801397w |url= |pmc=2765488}}</ref> Agonist antiepileptic drug candidate NAX 5055.<ref name="pmid19053761">{{cite journal |author=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 |year=2008 |month=December |pmid=19053761 |doi=10.1021/jm801088x |url=}}</ref><ref name="pmid19332332">{{cite journal |author=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 : the Journal of the American Society for Experimental NeuroTherapeutics |volume=6 |issue=2 |pages=372–80 |year=2009 |month=April |pmid=19332332 |doi=10.1016/j.nurt.2009.01.001 |url=}}</ref> Galanin in the ] is an inhibitor of ] but not of ]. This means that galanin is capable of increasing the ]<ref name= "receptors"/> and, therefore, is expected to act as an ]. To be specific, GalR1 has been linked to the suppression of spontaneous seizures.<ref name="pmid15350653"/><ref name="pmid19199479"/> An agonist antiepileptic drug candidate is NAX 5055.<ref name="pmid19053761"/><ref name="pmid19332332"/>


===In development=== ===In development===
It has been shown that galanin plays a role in the control of the early post-natal ] of the ] (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 ]. This suggests that the cell loss by ] 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. It has been shown that galanin plays a role in the control of the early post-natal ] of the ] (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 ]. This suggests that the cell loss by ] 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.


=== Pain and neuroprotection ===
===After injury===
Galanin plays an inhibitory role in pain processing,<ref name="Hobson_2010">{{cite journal | vauthors = Hobson SA, Bacon A, Elliot-Hunt CR, Holmes FE, Kerr NC, Pope R, Vanderplank P, Wynick D | journal = Experientia Supplementum (2012) | series = Experientia Supplementum | display-authors = 6 | title = Galanin acts as a trophic factor to the central and peripheral nervous systems | volume = 102 | pages = 25–38 | date = 2010 | pmid = 21299059 | doi = 10.1007/978-3-0346-0228-0_3 | publisher = Springer | isbn = 978-3-0346-0227-3 | veditors = Hökfelt T }}</ref> with high doses having been shown to reduce pain.<ref name="Mitsukawa_2010" /> When galanin is added to the spinal cord, ] is reduced.<ref name="Xu_2010">{{cite journal | vauthors = Xu XJ, Hökfelt T, Wiesenfeld-Hallin Z | journal = Experientia Supplementum (2012) | series = Experientia Supplementum | title = Galanin and spinal pain mechanisms: past, present, and future | volume = 102 | pages = 39–50 | date = 2010 | pmid = 21299060 | doi = 10.1007/978-3-0346-0228-0_4 | publisher = Springer | isbn = 978-3-0346-0227-3 | veditors = Hökfelt T }}</ref> Along with this, galanin is believed to be effective in reducing spinal hyperexcitability.<ref name="Xu_2010" /> Sensory neurons increasingly release galanin when they are damaged.<ref name="Xu_2010" /> An increase in the concentrations of galanin are also believed to be for ] reasons and lead to promoted ].<ref name="Mitsukawa_2010" /> GalR2 activation is believed to mediate the survival role galanin plays in the ].<ref name="Hobson_2010" />
''In vitro'' experiments show that DRG cells removed from galanin mutants have impaired abilities to extend ] 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 ] after crush injury which is linked to long-term functional problems.


===Parental role in mice===
==See also==
Galanin-expressing neurons in the medial preoptic area of the brain are responsible for regulating aggression towards pups by male mice.<ref name="pmid24828191"/>
* ]


Galanin-expressing neurons in the medial preoptic area are remodelled during pregnancy. Estrogen and progesterone genomic receptors in galanin (Gal)-expressing neurons control discrete aspected of plasticity.<ref>{{cite journal | vauthors = Ammari R, Monaca F, Cao M, Nassar E, Wai P, Del Grosso NA, Lee M, Borak N, Schneider-Luftman D, Kohl J | display-authors = 6 | title = Hormone-mediated neural remodeling orchestrates parenting onset during pregnancy | journal = Science | volume = 382 | issue = 6666 | pages = 76–81 | date = October 2023 | pmid = 37797007 | pmc = 7615220 | doi = 10.1126/science.adi0576 | bibcode = 2023Sci...382...76A }}</ref>
==References==
{{Reflist|2|refs=


== See also ==
<ref name="pmid7508413">{{cite journal | author = 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 | year = 1993 | month = December | pmid = 7508413 | doi = | url = | issn = }}</ref>
* ]


== References ==
<ref name="receptors">{{cite journal | author = 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 | year = 2008 | month = June | pmid = 18500647 | doi = 10.1007/s00018-008-8153-8 | url = | issn = }}</ref>
{{Reflist|refs=


<ref name="neuroscience">{{cite journal | author = Ito M | title = Functional roles of neuropeptides in cerebellar circuits | journal = Neuroscience | volume = 162 | issue = 3 | pages = 666–72 | year = 2009 | month = September | pmid = 19361475 | doi = 10.1016/j.neuroscience.2009.01.019 | url = | issn = }}</ref> <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 = 473–477 | date = December 1993 | pmid = 7508413 | doi = 10.1016/S0888-7543(11)80002-9 }}</ref>


<ref name = "acnp">{{cite web | author = Bartfai, T., | title = Galanin A neuropeptide with important central nervous system actions | date = 2000 | url = http://www.acnp.org/g4/GN401000054/CH054.html | accessdate = November 19, 2009}}</ref> <ref name="receptors">{{cite journal | vauthors = Mitsukawa K, Lu X, Bartfai T | title = Galanin, galanin receptors and drug targets | journal = Cellular and Molecular Life Sciences | volume = 65 | issue = 12 | pages = 1796–1805 | date = June 2008 | pmid = 18500647 | doi = 10.1007/s00018-008-8153-8 | s2cid = 263470319 | pmc = 11131746 }}</ref>


<ref name="MFN">{{cite journal | author = Wynick D, Thompson SW, McMahon SB | title = The role of galanin as a multi-functional neuropeptide in the nervous system | journal = Curr Opin Pharmacol | volume = 1 | issue = 1 | pages = 73–7 | year = 2001 | month = February | pmid = 11712539 | doi = 10.1016/S1471-4892(01)00006-6| 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 = 666–672 | date = September 2009 | pmid = 19361475 | doi = 10.1016/j.neuroscience.2009.01.019 | s2cid = 207245197 }}</ref>


<ref name="History">{{cite journal | author = Hökfelt T, Tatemoto K | title = Galanin--25 years with a multitalented neuropeptide | journal = Cell. Mol. Life Sci. | volume = 65 | issue = 12 | pages = 1793–5 | year = 2008 | month = June | pmid = 18500648 | doi = 10.1007/s00018-008-8152-9 | url = | issn = }}</ref> <ref name="acnp">{{cite web| vauthors = Bartfai T |title=Galanin A neuropeptide with important central nervous system actions |year=2000 |url=http://www.acnp.org/g4/GN401000054/CH054.html |access-date=November 19, 2009 |url-status=dead |archive-url=https://web.archive.org/web/20101202155619/http://www.acnp.org/g4/GN401000054/CH054.html |archive-date=December 2, 2010 }}</ref>


<ref name="pmid1710578">{{cite journal | author = 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 | year = 1991 | month = June | pmid = 1710578 | doi = 10.1016/0014-5793(91)80585-Q| url = | issn = }}</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 = 73–77 | date = February 2001 | pmid = 11712539 | doi = 10.1016/S1471-4892(01)00006-6 }}</ref>


<ref name="overview">{{cite journal | author = 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 | year = 2007 | month = August | pmid = 17604107 | doi = 10.1016/j.pharmthera.2007.05.009 | url = | issn = }}</ref> <ref name="History">{{cite journal | vauthors = Hökfelt T, Tatemoto K | title = Galanin--25 years with a multitalented neuropeptide | journal = Cellular and Molecular Life Sciences | volume = 65 | issue = 12 | pages = 1793–1795 | date = June 2008 | pmid = 18500648 | doi = 10.1007/s00018-008-8152-9 | s2cid = 19878753 | pmc = 11131681 }}</ref>


<ref name="pmid14993421">{{cite journal | author = 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 | year = 2003 | month = May | pmid = 14993421 | doi = 10.1124/mi.3.3.137 | url = | issn = }}</ref> <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 = 325–330 | date = May 2014 | pmid = 24828191 | pmc = 4105201 | doi = 10.1038/nature13307 | bibcode = 2014Natur.509..325W }}</ref>


<ref name="pmid18500641">{{cite journal | author = 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 | year = 2008 | month = June | pmid = 18500641 | pmc = 2911017 | doi = 10.1007/s00018-008-8159-2 | 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 Letters | volume = 283 | issue = 2 | pages = 189–194 | date = June 1991 | pmid = 1710578 | doi = 10.1016/0014-5793(91)80585-Q | s2cid = 19148582 | doi-access = free | bibcode = 1991FEBSL.283..189B }}</ref>
<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 = 1310–1316 | date = March 2009 | pmid = 19199479 | pmc = 2765488 | doi = 10.1021/jm801397w }}</ref>


<ref name="pmid16246567">{{cite journal | author = Ding X, MacTavish D, Kar S, Jhamandas JH | title = Galanin attenuates beta-amyloid (Abeta) toxicity in rat cholinergic ] neurons | journal = Neurobiol. Dis. | volume = 21 | issue = 2 | pages = 413–20 | year = 2006 | month = February | pmid = 16246567 | doi = 10.1016/j.nbd.2005.08.016 | url = | issn = }}</ref> <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 | display-authors = 6 | 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 = 8038–8047 | date = December 2008 | pmid = 19053761 | doi = 10.1021/jm801088x }}</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 = Pharmacology & Therapeutics | volume = 115 | issue = 2 | pages = 177–207 | date = August 2007 | pmid = 17604107 | doi = 10.1016/j.pharmthera.2007.05.009 }}</ref>


<ref name="pmid15350653">{{cite journal | author = 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="pmid19332332">{{cite journal | vauthors = White HS, Scholl EA, Klein BD, Flynn SP, Pruess TH, Green BR, Zhang L, Bulaj G | display-authors = 6 | title = Developing novel antiepileptic drugs: characterization of NAX 5055, a systemically-active galanin analog, in epilepsy models | journal = Neurotherapeutics | volume = 6 | issue = 2 | pages = 372–380 | date = April 2009 | pmid = 19332332 | pmc = 4402707 | doi = 10.1016/j.nurt.2009.01.001 }}</ref>


<ref name="pmid15944034">{{cite journal | author = 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 | year = 2005 | month = June | pmid = 15944034 | doi = 10.1016/j.npep.2004.12.016 | 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 = Molecular Interventions | volume = 3 | issue = 3 | pages = 137–156 | date = May 2003 | pmid = 14993421 | doi = 10.1124/mi.3.3.137 }}</ref>


<ref name="pmid16052044">{{cite journal | author = 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="pmid18500641">{{cite journal | vauthors = Counts SE, Perez SE, Mufson EJ | title = Galanin in Alzheimer's disease: neuroinhibitory or neuroprotective? | journal = Cellular and Molecular Life Sciences | volume = 65 | issue = 12 | pages = 1842–1853 | date = June 2008 | pmid = 18500641 | pmc = 2911017 | doi = 10.1007/s00018-008-8159-2 }}</ref>


<ref name="pmid18500643">{{cite journal | author = Mechenthaler I | title = Galanin and the neuroendocrine axes | journal = Cell. Mol. Life Sci. | volume = 65 | issue = 12 | pages = 1826–35 | year = 2008 | month = June | pmid = 18500643 | doi = 10.1007/s00018-008-8157-4 | 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 = Neurobiology of Disease | volume = 21 | issue = 2 | pages = 413–420 | date = February 2006 | pmid = 16246567 | doi = 10.1016/j.nbd.2005.08.016 | s2cid = 53192040 }}</ref>


<ref name="pmid2448788">{{cite journal | author = 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 | year = 1988 | month = February | pmid = 2448788 | pmc = 279702 | doi = 10.1073/pnas.85.4.1065| 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 = 431–441 | year = 2004 | pmid = 15350653 | pmc = 1360211 | doi = 10.1016/j.neuroscience.2004.06.052 }}</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 | display-authors = 6 | title = Galanin and galanin receptors in human cancers | journal = Neuropeptides | volume = 39 | issue = 3 | pages = 353–359 | date = June 2005 | pmid = 15944034 | doi = 10.1016/j.npep.2004.12.016 | s2cid = 1108702 }}</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 Medicine | volume = 7 | issue = 1–2 | pages = 157–180 | year = 2005 | pmid = 16052044 | doi = 10.1385/NMM:7:1-2:157 | s2cid = 19729607 }}</ref>

<ref name="pmid18500643">{{cite journal | vauthors = Mechenthaler I | title = Galanin and the neuroendocrine axes | journal = Cellular and Molecular Life Sciences | volume = 65 | issue = 12 | pages = 1826–1835 | date = June 2008 | pmid = 18500643 | doi = 10.1007/s00018-008-8157-4 | s2cid = 8754964 | pmc = 11131683 }}</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 = Proceedings of the National Academy of Sciences of the United States of America | volume = 85 | issue = 4 | pages = 1065–1069 | date = February 1988 | pmid = 2448788 | pmc = 279702 | doi = 10.1073/pnas.85.4.1065 | doi-access = free | bibcode = 1988PNAS...85.1065K }}</ref>


==Further reading==
{{Refbegin| 2}}
{{PBB_Further_reading
| citations =
*{{Cite journal | author=Vrontakis ME |title=Galanin: a biologically active peptide. |journal=Current drug targets. CNS and neurological disorders |volume=1 |issue= 6 |pages= 531–41 |year= 2003 |pmid= 12769595 |doi=10.2174/1568007023338914 }}
*{{Cite journal | author=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 |year= 2005 |pmid= 15893372 |doi= 10.1016/j.npep.2004.12.005 }}
*{{Cite journal | author=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 |year= 2006 |pmid= 16787232 |doi=10.2174/187152706777452281 }}
*{{Cite journal | author=McKnight GL, Karlsen AE, Kowalyk S, ''et al.'' |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 |year= 1992 |pmid= 1370155 |doi=10.2337/diabetes.41.1.82 }}
*{{Cite journal | author=Gai WP, Geffen LB, Blessing WW |title=Galanin immunoreactive neurons in the human hypothalamus: colocalization with vasopressin-containing neurons. |journal=J. Comp. Neurol. |volume=298 |issue= 3 |pages= 265–80 |year= 1990 |pmid= 1698834 |doi= 10.1002/cne.902980302 }}
*{{Cite journal | author=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 |year= 1991 |pmid= 1701228 |doi=10.1016/0143-4179(90)90115-F }}
*{{Cite journal | author=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=Hum. Reprod. |volume=5 |issue= 7 |pages= 870–6 |year= 1991 |pmid= 1702449 |doi= }}
*{{Cite journal | author=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 |year= 1991 |pmid= 1711463 |doi=10.1210/endo-129-1-270 }}
*{{Cite journal | author=Bennet WM, Hill SF, Ghatei MA, Bloom SR |title=Galanin in the normal human pituitary and brain and in pituitary adenomas. |journal=J. Endocrinol. |volume=130 |issue= 3 |pages= 463–7 |year= 1991 |pmid= 1719117 |doi=10.1677/joe.0.1300463 }}
*{{Cite journal | author=Schmidt WE, Kratzin H, Eckart K, ''et al.'' |title=Isolation and primary structure of pituitary human galanin, a 30-residue nonamidated neuropeptide. |journal=Proc. Natl. Acad. Sci. U.S.A. |volume=88 |issue= 24 |pages= 11435–9 |year= 1992 |pmid= 1722333 |doi=10.1073/pnas.88.24.11435 | pmc=53150 }}
*{{Cite journal | author=Bauer FE, Christofides ND, Hacker GW, ''et al.'' |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 | author=Bauer FE, Adrian TE, Christofides ND, ''et al.'' |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 |year= 1986 |pmid= 2427385 |doi= }}
*{{Cite journal | author=Tainio H, Vaalasti A, Rechardt L |title=The distribution of substance P-, CGRP-, galanin- and ANP-like immunoreactive nerves in human sweat glands. |journal=Histochem. J. |volume=19 |issue= 6-7 |pages= 375–80 |year= 1987 |pmid= 2444569 |doi=10.1007/BF01680455 }}
*{{Cite journal | author=Maggi CA, Santicioli P, Patacchini R, ''et al.'' |title=Galanin: a potent modulator of excitatory neurotransmission in the human urinary bladder. |journal=Eur. J. Pharmacol. |volume=143 |issue= 1 |pages= 135–7 |year= 1988 |pmid= 2446889 |doi=10.1016/0014-2999(87)90744-8 }}
*{{Cite journal | author=Marti E, Gibson SJ, Polak JM, ''et al.'' |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=J. Comp. Neurol. |volume=266 |issue= 3 |pages= 332–59 |year= 1988 |pmid= 2447134 |doi= 10.1002/cne.902660304 }}
*{{Cite journal | author=Beal MF, Clevens RA, Chattha GK, ''et al.'' |title=Galanin-like immunoreactivity is unchanged in Alzheimer's disease and Parkinson's disease dementia cerebral cortex. |journal=J. Neurochem. |volume=51 |issue= 6 |pages= 1935–41 |year= 1988 |pmid= 2460590 |doi=10.1111/j.1471-4159.1988.tb01181.x }}
*{{Cite journal | author=Berrettini WH, Kaye WH, Sunderland T, ''et al.'' |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== == External links ==
* {{MeshName|Galanin}} * {{MeshName|Galanin}}


{{Neuropeptides}} {{Neuropeptides}}

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