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{{details|topic=the basal ganglia|Basal ganglia}}
[[File:Basal ganglia and related structures (2).svg|thumb|right|Basal ganglia (red) and related structures (blue) shown within the brain]]
The [[dorsolateral]] striatum is associated with the acquisition of habits and is the main neuronal cell nucleus linked to procedural memory. Connecting excitatory [[afferent nerve fiber]]s help in the regulation of activity in the basal ganglia circuit. Essentially, two parallel information processing pathways diverge from the striatum. Both acting in opposition to each other in the control of movement, they allow for association with other needed functional structures<ref>{{cite journal | last1 = Alexander | first1 = GE | last2 = Crutcher | first2 = MD | year = 1990 | title = Functional architecture of basal ganglia circuits; neural substrates of parallel processing | journal = Trends Neurosci | volume = 13 | issue = 7| pages = 266–271 | doi=10.1016/0166-2236(90)90107-l | pmid=1695401| s2cid = 3990601 }}</ref> One pathway is direct while the other is indirect and all pathways work together to allow for a functional neural feedback loop. Many looping circuits connect back at the striatum from other areas of the brain; including those from the emotion-center linked limbic cortex, the reward-center linked [[ventral striatum]] and other important motor regions related to movement.<ref>{{cite journal | last1 = Haber | first1 = SN | last2 = Fudge | first2 = JL | last3 = McFarland | first3 = NR | year = 2000 | title = Striatonigrostriatal pathways in primates form an ascending spiral from the shell to the dorsolateral striatum | journal = J. Neurosci. | volume = 20 | issue = 6| pages = 2369–2382 | doi = 10.1523/JNEUROSCI.20-06-02369.2000 | pmid = 10704511 | pmc = 6772499 | doi-access = free }}</ref> The main looping circuit involved in the motor skill part of procedural memory is usually called the cortex-basal ganglia-thalamus-cortex loop.<ref>{{cite journal | last1 = Parent | first1 = A | year = 1990 | title = Extrinsic connections of the basal ganglia | journal = Trends Neurosci | volume = 13 | issue = 7| pages = 254–258 | doi=10.1016/0166-2236(90)90105-j| pmid = 1695399 | s2cid = 3995498 }}</ref>
The striatum is unique because it lacks the [[glutamate]]-related neurons found throughout most of the brain. Instead, it is categorized by a high concentration of a special type of [[GABA]] related inhibiting cell known as the [[medium spiny neuron]].<ref>{{cite journal | last1 = Smith | first1 = Y. | last2 = Raju | first2 = D. V. | last3 = Pare | first3 = J. F. | last4 = Sidibe | first4 = M. | year = 2004 | title = The thalamostriatal system: a highly specific network of the basal ganglia circuitry | url = https://www.semanticscholar.org/paper/99588f5770f5388d0c260eb6b70b9c88ebff0171| journal = Trends Neurosci | volume = 27 | issue = 9| pages = 520–527 | doi=10.1016/j.tins.2004.07.004| pmid = 15331233 | s2cid = 22202019 }}</ref> The two parallel pathways previously mentioned travel to and from the striatum and are made up of these same special medium spiny neurons. These neurons are all sensitive to different neurotransmitters and contain a variety of corresponding receptors including dopamine receptors ([[DRD1]], [[DRD2]]), [[muscarinic receptors]] (M4) and [[adenosine receptors]] (A2A). Separate interneurons are known to communicate with striatal spiny neurons in the presence of the [[somatic nervous system]] neurotransmitter [[acetylcholine]].<ref>{{cite journal | last1 = Zhou | first1 = FM | last2 = Wilson | first2 = CJ | last3 = Dani | first3 = JA | year = 2002 | title = Cholinergic Interneuron characteristics and nicotinic properties in the striatum | journal = J. Neurobiol. | volume = 53 | issue = 4| pages = 590–605 | doi=10.1002/neu.10150 | pmid=12436423| doi-access = free }}</ref>
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===Limbic system===
{{details|topic=the limbic system|Limbic system}}
The [[limbic system]] is a group of unique brain areas that work together in many interrelated processes involved in emotion, motivation, learning and memory. Current thinking indicates that the limbic system shares anatomy with a component of the neostriatum already credited with the major task of controlling procedural memory. Once thought to be functionally separate, this vital section of the brain found on the striatum's back border has only recently been linked to memory and is now being called the marginal division zone (MrD).<ref>{{cite journal | last1 = Shu | first1 = S.Y. | last2 = Bao | first2 = X.M. | last3 = Li | first3 = S.X. | last4 = Chan | first4 = W.Y. | last5 = Yew | first5 = D. | year = 2000 | title = A New Subdivision, Marginal Division, in the Neostriatum of the Monkey Brain | journal = Biomedical and Life Sciences | volume = 25 | issue = 2| pages = 231–7 | doi = 10.1023/a:1007523520251 | pmid = 10786707 | s2cid = 11876741 }}</ref> A special membrane protein associated with the limbic system is said to concentrate in related structures and to travel towards the basal nuclei. To put things simply, the activation of brain regions that work together during procedural memory can be followed because of this limbic system associated membrane protein and its application in molecular and [[immunohistochemistry]] research.<ref>{{cite journal | last1 = Yun Shu | first1 = Si | last2 = Min Bao | first2 = Xin | last3 = Ning | first3 = Qun | last4 = Ming Wu | first4 = Yong | last5 = Wang | first5 = Jun | last6 = Leonard | first6 = Brian E. | year = 2003 | title = New component of the limbic system; Marginal division of the neostriatum that links the limbic system to the basal nucleus of Meynert | journal = Journal of Neuroscience Research | volume = 71 | issue = 5| pages = 751–757 | doi=10.1002/jnr.10518| pmid = 12584733 | s2cid = 21343863 }}</ref>
==Physiology==
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===Human immunodeficiency virus (HIV)===
{{details|topic=human immunodeficiency virus|HIV}}
Neural systems used by procedural memory are commonly targeted by [[Human Immunodeficiency Virus]]; the striatum being the structure most notably affected.<ref>{{cite journal | last1 = Reger | first1 = M | last2 = Welsh | first2 = R | last3 = Razani | first3 = J | last4 = Martin | first4 = DJ | last5 = Boone | first5 = KB | year = 2002 | title = A meta-analysis of the neuropsychological sequelae of HIV infection | journal = Journal of the International Neuropsychological Society | volume = 8 | issue = 3| pages = 410–424 | doi=10.1017/s1355617702813212| pmid = 11939699 | s2cid = 30520253 }}</ref> MRI studies have even shown white matter irregularity and basal ganglia subcortical atrophy in these vital areas necessary for both procedural memory and motor-skill.<ref>{{cite journal | last1 = Chang | first1 = L | last2 = Lee | first2 = PL | last3 = Yiannoutsos | first3 = CT | last4 = Ernst | first4 = T | last5 = Marra | first5 = CM | last6 = Richards | first6 = T | display-authors = etal | year = 2004 | title = A multicenter in vivo proton-MRS study of HIV-associated dementia and its relationship to age | url = https://www.semanticscholar.org/paper/f773811cd3929a13639cb15fa8cedcf5f0e01212| journal = NeuroImage | volume = 23 | issue = 4| pages = 1336–1347 | doi=10.1016/j.neuroimage.2004.07.067 | pmid=15589098| s2cid = 2664814 }}</ref> Applied research using various procedural memory tasks such as the Rotary pursuit, Mirror star tracing and Weather prediction tasks have shown that HIV positive individuals perform worse than HIV negative participants suggesting that poorer overall performance on tasks is due to the specific changes in the brain caused by the disease.<ref>{{cite journal | last1 = Gonzalez | first1 = R | last2 = Jacobus | first2 = J | last3 = Amatya | first3 = AK | last4 = Quartana | first4 = PJ | last5 = Vassileva | first5 = J | last6 = Martin | first6 = EM | year = 2008 | title = Deficits in complex motor functions, despite no evidence of procedural learning deficits, among HIV+ individuals with history of substance dependence | journal = Neuropsychology | volume = 22 | issue = 6| pages = 776–86 | doi=10.1037/a0013404| pmid = 18999351 | pmc = 2630709 }}</ref>
===Huntington's disease===
{{details|topic=Huntington's disease|Huntington's disease}}
[[File:Huntington.jpg|thumb|left|Coronal FSPGR through the brain of Huntington's patient]]
Despite being a disorder that directly affects striatal areas of the brain used in procedural memory, most individuals with [[Huntington's disease]] don't display the same memory problems as other people with striatum related brain diseases.<ref>{{cite journal | last1 = Sprengelmeyer | first1 = R | last2 = Canavan | first2 = AG | last3 = Lange | first3 = HW | last4 = Hömberg | first4 = V | date = Jan 1995 | title = Associative learning in degenerative neostriatal disorders: contrasts in explicit and implicit remembering between Parkinson's and Huntington's diseases | journal = Mov Disord | volume = 10 | issue = 1| pages = 51–65 | doi=10.1002/mds.870100110| pmid = 7885356 | s2cid = 38578307 }}</ref> In more advanced stages of the disease, however, procedural memory is affected by damage to the important brain pathways that help the inner subcortical and prefrontal cortex parts of the brain to communicate.<ref>Saint-Cyr JA, Taylor AE, Lang AE. (1988) "Procedural learning and neostriatal dysfunction in man" ''Brain'' 1988 Aug;111 ( Pt 4):941-59.</ref>
===Obsessive compulsive disorder===
{{details|topic=obsessive–compulsive disorder|OCD}}
Neuroimaging studies show that [[OCD]] patients perform considerably better on procedural memory tasks because of noticeable over-activation of the striatum brain structures, specifically the frontostriatal circuit. These studies suggest that procedural memory in OCD patients is unusually improved in the early learning stages of procedural memory.<ref>{{cite journal | last1 = Roth | first1 = RM | last2 = Baribeau | first2 = J | last3 = Milovan | first3 = D | last4 = O'Connor | first4 = K | last5 = Todorov | first5 = C | date = Sep 2004 | title = Procedural and declarative memory in obsessive-compulsive disorder | journal = J Int Neuropsychol Soc | volume = 10 | issue = 5| pages = 647–54 | doi=10.1017/s1355617704105018| pmid = 15327712 | s2cid = 29064519 }}</ref> Individuals with OCD do not perform significantly different on procedural working memory tasks than healthy controls.<ref name="Shahar 197–204"/>
===Parkinson's disease===
{{details|topic=Parkinson's disease|Parkinson's disease}}
[[Parkinson's disease]] is known to affect selective areas in the frontal lobe area of the brain. Current scientific information suggests that the memory performance problems notably shown in patients are controlled by unusual frontostriatal circuits.<ref>{{cite journal | last1 = Sarazin | first1 = M | last2 = Deweer | first2 = B | last3 = Pillon | first3 = B | last4 = Merkl | first4 = A | last5 = Dubois | first5 = B | date = Dec 2001 | title = Procedural learning and Parkinson disease: implication of striato-frontal loops | journal = Rev Neurol | volume = 157 | issue = 12| pages = 1513–8 | pmid = 11924447 }}</ref> Parkinson's patients often have difficulty with the sequence-specific knowledge that is needed in the acquisition step of procedural memory.<ref>{{cite journal | last1 = Muslimovic | first1 = D | last2 = Post | first2 = B | last3 = Speelman | first3 = JD | last4 = Schmand | first4 = B | date = Nov 2007 | title = Motor procedural learning in Parkinson's disease | journal = Brain | volume = 130 | issue = 11| pages = 2887–97 | doi=10.1093/brain/awm211| pmid = 17855374 | doi-access = free }}</ref> Further evidence suggests that the frontal lobe networks relate to executive function and only act when specific tasks are presented to the patient. This tells us that the frontostriatal circuits are independent but able to work collaboratively with other areas of the brain to help with various things such as paying attention or focusing.<ref>{{cite journal | last1 = Sarazin | first1 = M | last2 = Deweer | first2 = B | last3 = Merkl | first3 = A | last4 = Von Poser | first4 = N | last5 = Pillon | first5 = B | last6 = Dubois | first6 = B | date = Mar 2002 | title = Procedural learning and striatofrontal dysfunction in Parkinson's disease | journal = Mov Disord | volume = 17 | issue = 2| pages = 265–73 | doi=10.1002/mds.10018| pmid = 11921111 | s2cid = 32165795 }}</ref>
===Schizophrenia===
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==Sleep==
Practice is clearly an important process for learning and perfecting a new skill. With over 40 years of research, it is well established in both humans and animals that the formation of all forms of memory are greatly enhanced during the brain-state of sleep. Furthermore, with humans, sleep has been consistently shown to aid in the development of procedural knowledge by the ongoing process of memory consolidation, especially when sleep soon follows the initial phase of memory acquisition.<ref>{{cite journal | last1 = Karni | first1 = A. | last2 = Tanne | first2 = D. | last3 = Rubenstein | first3 = B.S. | last4 = Askenasy | first4 = J.J. | last5 = Sagi | first5 = D. | year = 1994 | title = Dependence on REM sleep of overnight improvement of a perceptual skill | journal = Science | volume = 265 | issue = 5172| pages = 679–682 | doi=10.1126/science.8036518| pmid = 8036518 | bibcode = 1994Sci...265..679K }}</ref><ref>{{cite journal | last1 = Gais | first1 = S. | last2 = Plihal | first2 = W. | last3 = Wagner | first3 = U. | last4 = Born | first4 = J. | year = 2000 | title = Early sleep triggers memory for early visual discrimination skills
Whether a skill is learned explicitly (with [[attention]]) or implicitly, each plays a role in the offline consolidation effect. Research suggests that explicit awareness and understanding of the skill being learned during the acquisition process greatly improves the consolidation of procedural memories during sleep.<ref>{{cite journal | last1 = Robertson | first1 = E.M. | display-authors = etal | year = 2004 | title = Awareness modifies skill-learning benefits of sleep | journal = Curr. Biol. | volume = 14 | issue = 3| pages = 208–212 | doi=10.1016/s0960-9822(04)00039-9| pmid = 14761652 | doi-access = free }}</ref> This finding is not surprising, as it is widely accepted that intention and awareness at time of learning enhances the acquisition of most forms of memory.
==Language==
Language works because of the brain’s ability to retrieve pieces of information from memory and then combine those pieces into a larger, more complex unit based on context. The latter part of this process is called unification.<ref>{{cite journal | last1 = Hagoort | first1 = Peter | title = MUC (Memory, Unification, Control) and beyond | journal = Frontiers in Psychology | volume = 4 | year = 2013 | pages = 416 | doi = 10.3389/fpsyg.2013.00416| pmid = 23874313 | pmc = 3709422 | doi-access = free }}</ref> Results of several studies provide evidence that suggests procedural memory is not only responsible for sequential unification, but for syntactic priming and grammatical processing as well.
One study used patients with [[Korsakoff’s syndrome]] to show that procedural memory subserves [[syntactic priming]]. Although Korsakoff’s patients have deficits in declarative memory, their nondeclarative memory is preserved, allowing them to successfully complete syntactic priming tasks, as in the study. This result proves syntactic priming is a nondeclarative memory function. These patients were also capable of forming proper grammatical sentences, suggesting that procedural memory is responsible for grammatical processing in addition to syntactic priming.<ref>{{cite journal|last1=Heyselaar|first1=Evelien|last2=Segaert|first2=Katrien|last3=Walvoort|first3=Serge J.W.|last4=Kessels|first4=Roy P.C.|last5=Hagoort|first5=Peter|year=2017|title=The role of nondeclarative memory in the skill for language: Evidence from syntactic priming in patients with amnesia|url=http://pure-oai.bham.ac.uk/ws/files/40798978/HSWKH_Neuropsychologia_revised_2.pdf|journal=Neuropsychologia|volume=101|pages=97–105|doi=10.1016/j.neuropsychologia.2017.04.033|pmid=28465069|hdl=11858/00-001M-0000-002D-4D0D-1|s2cid=4109634|hdl-access=free}}<!--http://pure-oai.bham.ac.uk/ws/files/40798978/HSWKH_Neuropsychologia_revised_2.pdf--></ref>
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