The Basal Ganglia and Chunking of Action Repertoires

“, in creating chunks) [20–24], the functional contribution of the hippocampus could be in the detection and formation of higher-order sequential associations [9,25–27] and the stabilization of the motor performance [7].”

“The classical concept of parallel loops (Alexander et al., 1986) has been modified by findings of considerable convergence in cortico-striate projections (Graybiel, 1998; Haber et al., 2006).”

“The behavioral fate of repetitive actions, such as compulsive alcohol consumption, seem to be instantiated in mesostriatocortical networks (Graybiel 1998; Volkow et al. 2013).”

“For example, the neurobiology of perseverative, ‘compulsive-like’ behavior can be rigorously studied in animals, an effort relevant to understanding OCD, given that perseverative behavior (compulsion) and perseverative thought (obsession) may involve similar neural circuitry [3,25].”

“This pattern of results with ACh output in the dorsomedial striatum is consistent with the idea proposed by Graybiel (1998) in which cholinergic interneurons may be important for facilitating the coordination between different striatal input–output modules, but are not essential for the expression…”

“Graybiel (1998) has shown that the BG is involved in the integration of motor and cognitive sequencing.”

“Although chunking in motor sequence learning has been extensively studied behaviorally, it has gained less attention in neuroimaging studies (see also Bor et al., 2003; Graybiel, 1998; Kennerley et al., 2004; Verwey et al., 2002).”

“This flow of information is likely involved in the initiation and sequencing of learned actions (Graybiel, 1998) and in habit formation through NMDARdependent corticostriatal plasticity (Kelley et al.”

“…an “error” signal which modulates corticostriatal synaptic transmission (Wickens 1990; Miller and Wickens 1991; Graybiel et al. 1994; Pennartz et al. 1994; Groves et al. 1995; Houk 1995; Houk et al. 1995; Kimura 1995; Pennartz 1995; Schultz et al. 1995a,b; Wickens and Kotter 1995; Graybiel 1998).”

“Activation of the latDS and NAcb contribute differently to the generation of behavior (Graybiel, 1998; Cardinal et al., 2002; Ghitza Correspondence: Dr F. Woodward Hopf, as above.”

“As a consequence, the general ability to identify and learn sequences is a widespread feature across species and throughout development (Terrace and Mcgonigle, 1994; Saffran et al., 1996; Graybiel, 1998; Dehaene et al., 2015), but the specific ways by which sequences are learned can show substantial differences.”

“…the basal ganglia are critical for both cognitive and motor aspects of sequential learning (e.g., Canales & Graybiel, 2000; Fujii & Graybiel, 2003; Graybiel, 1997; 1998; 2000; Jog, Kubota, Connolly, Hillegaart, & Graybiel, 1999; Saka, Goodrich, Harlan, Madras, & Graybiel, 2004) and likely also…”

“The dorsal circuit links the dorsolateral PFC (DLPFC) with the dorsal caudate and putamen, and plays a role in information integration and associative learning (27,31,39,40).”

“Finally, this framework can also be readily extended to sequences of actions, such that a chunked sequence of actions predictive of reward, later becomes repeated as a unit (Barnes et al., 2005; Dezfouli and Balleine, 2012; Graybiel, 1998).”

“Corticostriatal circuits are crucial for sequential behavior and especially its acquisition [1].”

“…is thought to mediate reinforcement learning (Bogacz and Larsen 2011; Costa 2007) and behavioral selection via connections to the basal ganglia (Graybiel 1998), the disturbance of which could explain why OCD patients choose inappropriate actions for specific circumstances, as a result of an…”

“…the basal ganglia have also been implicated in several cognitive processes, including language (Crosson, 1992), learning and memory (Crosson, 1992; Graybiel, 1998), working memory (Mattay et al., 2002), problem solving (Owen & Doyon, 1999), and executive functions (Hassler, 1978; Saint-Cyr,…”

“These language structures are holistically produced as a complete gesture, possibly benefiting from the motor organization and chunking processes of the basal ganglia (Graybiel, 1998).”

“As already mentioned, besides Broca ’ s area, the other structure commonly considered to be involved in chunking is the BG (Jin, Tecuapetla, & Costa, 2014; Wymbs et al., 2012; Boyd et al., 2009; Graybiel, 1998).”

“, cueresponse-reward) are “chunked” into a sequence as a single entity (Graybiel, 1998), where only the beginning of the sequence generates a phasic dopamine response, unless the outcome varies from how the sequence is expected to end.”

“It has also been demonstrated experimentally that cortically derived information is re-coded within the striatum, chunking thereby the representations of motor and cognitive action sequences so that they can be implemented en bloc as performance Ž .units Graybiel, 1998 .”

“…Bunsey and Eichenbaum 1996; Burgess et al. 2002) and rigid, slowly acquired stimulus–response associations (which depend on the striatum; Graybiel 1998; Bayley et al. 2005;YinandKnowlton2006)hasbeentheorizedtocontributetodisorderslikeaddictionandpost-traumaticstressdisorder(PTSD;…”

“Routines and habits are established there (Graybiel, 1998; Knowlton, Mangels, & Squire, 1996; Lieberman, 2001).”

“This process may relate to the known role of the basal ganglia in motor chunking (Graybiel, 1998; Yin, 2010; Wymbs et al., 2012).”

“Animal studies have suggested that the role of the striatum in motor learning lies in the combination or the chunking of discrete movements into motor sequences ( Graybiel 1998; Jog et al. 1999).”

“The cortical-striatal pathways implicated in TTM appear to play a key role in the acquisition and retrieval of motor sequences (Graybiel, 1998).”

“Corticostriatal interactions ‘‘chunk’’ together these sequences of movement (Graybiel, 1998) and ‘‘gate’’ or select behavioral ‘‘candidates’’ on the basis of instrumental learning processes (Cockburn & Frank, 2011).”

“Instead, they reflect a domain-general sensori-motor chunking that links to working memory (Graybiel, 1998).”

“Such outcome would support our assumption that each sequence rendition was performed as a single executable unit (Graybiel, 1998), despite the continuous performance of the sequence renditions.”

“Findings from experimental studies in animals together with clinical studies in humans suggest that the striatum is involved in procedural or habit memory (Graybiel, 1998; Knowlton, Mangels, & Squire, 1996; Mishkin, Malamut, & Bachevalier, 1984; Packard & McGaugh, 1996).”

“An important component of this function of the basal ganglia is organizing material into unitary structures or chunks (Graybiel, 1998).”

“Because this integration is used to control reinforcement learning [67,74] and behavioural selection in the basal ganglia [69], its disturbance could explain why OCD patients select inappropriate programs for the circumstances.”

“The selective function of BG is not innate but is acquired through trial-and-error learning processes (Graybiel, 1998, 2005).”

“, and Graybiel, A. M. (1999).”

“On the motor level, for example, projections of the representation 534 of the hand and digits in the motor cortex (handknob; Yousry et al., 1997) interdigitate in the 535 striatum with the projections from the hand area in primary sensory cortex (S1; Graybiel, 1998; 536 Flaherty and Graybiel, 1991).”

“Indeed, cortico-basal ganglia loops are involved in the formation and automation of behavioural sequences [13]; a dysfunction of this system could, therefore, lock the patient into a specific behavioural program (i.”

“…Tang et al., 2007; Yin and Knowlton, 2006; Yin et al., 2004) that provide a substrate for skill learning (Benecke et al., 1987; Boyd et al., 2009; Graybiel, 1998;Hikosakaet al., 1999; Jog et al., 1999; Kermadi et al., 1993; Sakai et al., 2003; Seidler et al., 2007; Yin et al., 2009), especially…”

“The striatum is thought to play a critical role in habit learning and in the chunking of automated action sequences, according to tiers of evidence from animals and humans (Graybiel, 1998).”

“In addition to the importance of the DS in motor control, learning, memory [47,61,95,112], motivational processes (such as feeding behav-”

“It may act as a facilitating signal to focus cortico-striatal activity and reinforce the “motor readiness” and “release” functions that have been ascribed to the basal ganglia on the basis of clinical and experimental studies (Graybiel 1998).”

“These results suggest a specific role for cortical–basal ganglia circuits in organizing performance units (or ‘chunks’) of learned action sequences (Graybiel, 1998).”

“analysis of transition speeds (there are four unique key-press transitions: 4→1, 1→3, 3→2, and 2→4 in the sequence 4-1-3-2-4) led them to claim that sleep selectively improves the speed of the transitions that are the slowest before sleep. The change from a pre-sleep pattern of uneven speed across transition”

“Within a biological perspective, TERL architecture might be used to model basal ganglia-cortical loops [101], a brain hierarchical system playing a key role in trial-and-error learning in organisms [48], [96]–[98].”

“While the overlap between the neural correlates of chunking and WM gating (Graybiel 1998; Bor et al. 2003; Gruber et al. 2006; Tremblay et al. 2010; Murty et al. 2011; D’Ardenne et al. 2012; Wymbs et al. 2012; Jin et al. 2014) was considered as indirect evidence of their possible interactions, in…”

“[72]), our early studies with the rat SRTT were aimed at striatal dopamine, which we studied by testing the effects of systemic dopaminergic antagonists [42] and 6-OHDA lesions of ventral striatal or neostriatal dopamine [44,48,49].”

“the dorsal striatum, which supports response-driven behavior (Graybiel, 1998; Packard & McGaugh, 1992; Pych, Chang, ColonRivera, & Gold, 2005).”

“The critical role of the cortico-striato-thalamo-cortical loop in feed forward, automatized movements is well established in the literature (Graybiel, 1998).”

“Although an extensive review of the correlation between executive function and its anatomical substrates is beyond the scope of this article, it can be said that the location of executive functions implies activation of pre-frontal lobes, basal ganglia structures and other cortical and subcortical areas, including cor…”

“Studies aimed at finding the neural substrate for habitual behavior suggest an involvement of the subcortical structure, dorsolateral striatum (Hikosaka et al., 1999; Yin and Knowlton, 2006; Tricomi et al., 2009).”

“This proposal is supported by models of brain function, details of which are beyond the scope of this paper (Bever, 1975; Ullman, 2004; Graybiel, 1998, 2008; for a review see Van Lancker Sidtis, 2014).”

“…strongest anatomical contrast) using a 3 degrees of freedom (translation-only) linear FLIRT alignment, 2) using boundary-based registration (BBR; Greve and Fischl, 2009) to co-register the first EPI volume to the bias corrected, FNIRT-based brain extracted, and segmented structural image, 3)…”

“shown by previous work to be critical for instrumental learning, the learning of the relationship between action and outcomes (Graybiel, 1998; Yin et al., 2005a,b; Yin & Knowlton, 2006).”

“To learn to combine these individual movements into a well-programmed sequence of events is referred to as chunking and is also an important contribution of the basal ganglia (130, 174, 175).”

“The DS is important for behaviors such as: motor activity, memory and learning (Saint-Cyr et al., 1995; Lalonde and Botez-Marquard, 1997; Graybiel, 1998; Yin and Knowlton, 2006), drug addiction (Nestler, 2005; Kreitzer and Malenka, 2008), and for the nonhedonic response to food intake (Volkow et…”

“R learning (Graybiel, 1998), this opens the possibility that both visual and auditory sensory inputs participate in the shaping of the hierarchical organization of sequential sensorimotor behaviors.”