The Basal Ganglia and Chunking of Action Repertoires

“The dorsolateral striatum is known to be engaged in stereotypical behaviors (Packard and McGaugh, 1992; Graybriel, 1998; Jog et al., 1999; Mizumori et al., 2004).”

“…autonomic and endocrine function (Parvizi & Damasio, 2001); to other areas of the basal ganglia that are associated with voluntary motor actions (Graybiel, 1998); to the central nucleus of amygdala, which directs attention (Holland & Gallagher, 1999); and to the ventromedial prefrontal cortex,…”

“For example, Poldrack and others (2001) found greater striatal activity during a probabilistic classification task, in which subjects learned to classify stimuli into categories, but greater hippocampal activity during a paired associates task, in which subjects memorized relationships between stimuli and category nam…”

“Habituation appears to be instantiated mainly in the mesostriatocortical circuits that ‘re-code’ the behavioral fate of repetitive actions [14,15] in a process that was aptly referred to as the ‘chunking’ of action repertoires [16 ].”

“Wide support exists for the key role of the BG, and the neurotransmitter dopamine (DA), in both Pavlovian and instrumental learning (Schultz et al., 1997; Schultz, 2007; O'Doherty et al., 2004), as well as habit learning (Graybiel, 1998; Yin and Knowlton, 2006).”

“…through repetition is closely related to its role in the chunking of action sequences; through repeated practice, behaviors that are composed of multiple actions can become unitized into a single action (Barnes et al., 2011; Graybiel, 1998; Jog, Kubota, Connolly, Hillegaart, & Graybiel, 1999).”

“In addition to its relationship to the apparent structural features of the striatum, this view is consistent with the current functional view of the striatum as engaged in action selection (Graybiel 1998; Gurney et al. 2001; Lawrence et al. 2000; Marsden 1984; Wickens 1993).”

“Specifically, corticothalamic loops are separably modulated by “Go” and “NoGo” striatal pathways, with activity in Go pathways acting to disinhibit each loop, and activity in NoGo pathways acting to inhibit that same loop (Mink et al., 1996; Graybiel, 1998; Frank et al., 2001; Hazy et al. 2007).”

“…Treves (1998) have argued that it is the cortex which imposes patterns on the underlying striatum (behavioral set, according to Hikosaka, 1993; state space according to Graybiel, 1998) and that we should not mis-attribute functions that have been computed at the cortical level to the basal ganglia.”

“Since the nonNMDA glutamate activity is necessary before the NMDA receptor can become activated, this selective amplification of the NMDA response should selectively boost already-strong input signals, as might be expected when the striatal cell receives converging excitatory input from a large number of cortical cell…”

“There are numerous indications that the basal ganglia can represent holistic action sequences that are subsequently implemented as performance units (Graybiel, 1998).”

“Tics are thought to result from dysfunctions in cortical and sub-cortical regions that are involved in habit formation, including the basal ganglia, thalamus, and frontal cortex (Graybiel 1998; Leck-man and Riddle 2000; Leckman 2002; Leckman et al. 2006; Graybiel 2008).”

“According to this view, during skill learning, multiple movement units can be combined into larger units by ‘chunking’, and multiple chunks can be formed to build up the whole behavioral sequence [13,17,18,88].”

“The BG select and coordinate cortical response patterns The studies described above demonstrate that the BG support not only motor functions (Graybiel, 1998; Yin et al., 2004; Keeler et al., 2014; Pidoux et al., 2018) but also limbic functions such as reward valuation and motivation (Yin et al.,…”

“…cortex; dual-label immunocytochemistry; electron microscopy; glutamate transmission; reward; striatum The caudate putamen nucleus (CPN) plays a major role in motor function and has also been implicated in certain types of learning and memory (White, 1997; Graybiel, 1998; Graybiel et al., 2000).”

“Thus, when generating more striatal activity, controls were likely engaging an automatic response tendency based within that region.(53,54) In contrast, the more the patients activated both cortical and subcortical regions (ie, the less deficient their overall pattern of brain activation) (Figure 2), the fewer errors…”

“One possibility raised by our findings is that efficient proceduralization might accelerate probabilistic learning of language features (10) by chunking single speech and languagerelated actions into sequences, a chunking function that has been suggested to be a core property of the striatum in experimental work (48,…”

“…Hu, Jin, & Graybiel, 2005; Jog, Kubota, Connolly, Hillegaart, & Graybiel, 1999) code the beginnings and ends of action sequences, suggesting that frontostriatal circuits may participate in the “chunking” of action sequences (Graybiel, 1998) that characterizes hierarchical reinforcement learning.”

“Accordingly, Graybiel (1998) proposed that the basal ganglia may chunk the representations of motor and cognitive action sequences so that they can be implemented as performance units.”

“For example, learning about rewards involves the affective loop (including the ventral striatum and orbitofrontal cortex; Galvan and others 2005), and learning visual categories involves the visual loop (including the inferior temporal cortex and body/tail of the caudate; Ashby and Maddox 2005). An emerging area of re…”

“This result is open to a number interpretations, however, one possibility is that action selection by the basal ganglia may be involved in constructing new movement sequences which, following practice, then become available for selection as larger ‘chunks’ of behavior (Graybiel, 1998).”

“…with the basal ganglia may form part of the central control mechanism for motor sequencing (Brotchie et al. 1991; Cromwell and Berridge 1996; Graybiel 1995, 1998; Kermadi and Joseph 1995; Kimura et al. 1992; Marsden 1987; Miyachi et al. 1997; Mushiake and Strick 1996; Roland et al. 1980;…”

“The circuit, shown on Figure 5, is mainly based on data supporting the contribution of pre-SMA, SMA, Cer and BG in managing action sequences.(17,33,36,37) Different types of neurons in pre-SMA and SMA help to encode not only where in a sequence the action is but also the conditional links between the previous response…”

“The role of the more lateral systems in the patterning of behavior is better described by the concept of unconscious “habit strength” (Everitt & Robbins, 2005; Graybiel, 1998; Ikemoto & Panksepp, 1999; Yin & Knowlton, 2006).”

“be unwieldy (Graybiel, 1998; Miller, 1956).”

“…representations of sensory contexts and motor acts with mesencephalic reward signals, thus conveying behavioural context into cortical ensembles as a function of learning (Dominey et al., 1995; Houk and Wise, 1995; Dominey and Boussaoud, 1997; Graybiel, 1998; Hikosaka et al., 1999; Schultz, 2000).”

“The striatum is a main component of the basal ganglia involved in adaptive control of behavior and automation of action (Graybiel, 1998, 2005; Yin and Knowlton, 2006).”

“In particular, the basal ganglia have an important role in action sequencing (Lehericy et al., 2005, Graybiel, 1998) and in learning about the rewards associated with actions (Zink et al., 2004, Hollerman et al., 2000).”

“The BG may have a role in that they may be involved in habit and stimulusresponse learning [16].”

“Theorists view the corticostriatal system as an ideal substrate for the recognition of cortical states (Houk, 1995; Houk and Wise, 1995; Graybiel, 1998; Redgrave et al., 1999).”

“It is proposed that the basal ganglia control sequences of beha- viour via ‘chunking’ (Graybiel, 1998; Smith & Graybiel, 2014).”

“…adaptations rely on active maintenance of recent outcomes in the orbitofrontal cortex, whereas probabilistic learning depends on incremental synaptic weight changes in the basal ganglia (Knowlton et al., 1996; Packard and McGaugh, 1996; Graybiel, 1998, 2004; Delgado et al., 2000, 2005).”

“First, hierarchical representations of behavior can be more compact or efficient than non-hierarchical (flat) representations, allowing complex behaviors to be encoded more economically at the neural level [9].”

“…direct and indirect pathways in distinct ways and have important implications for a wide range of neurological diseases, from Parkinson’s disease to speech disorders (Brainard and Doupe, 2002; Graybiel, 1998; Hikosaka et al., 1999; Jin and Costa, 2015; Lai et al., 2001; Vinter and Gras, 1998).”

“Because the Pf has reciprocal connections with the basal ganglia (Berendse and Groenewegen, 1990), a region important for response selection (Graybiel, 1998; Balleine et al., 2009; Kimchi and Laubach, 2009), the Pf may relay information to the basal ganglia to signal a change in choice selection…”

“Our study was restricted to analysis of neocortical representation, and we did not address how sequences are represented in sub-cortical structures (Doyon et al., 2002; Graybiel, 1998; Graybiel and Grafton, 2015; Wymbs et al., 2012).”

“Lastly, intrinsic fluctuations in DA and ACh may help specify the timing of self-paced volitional actions 7,14,15,19 as well as sustain cognitive processes that lack overt behavioural correlates 48–50 .”

“1993; Salamone, 1992; Salamone and Correa, 2012), and in the role of the striatum in acquiring and executing skilled motor patterns (Graybiel, 1998; Hikosaka, 1991; Lacourse et al., 2005; Lovinger, 2010; Salamone, 1992; Yin et al., 2009).”

“Chunksplay a key role in the ability of the nervous system to efficiently learn, store and recall motor procedures such as walking, speaking or playing musical instruments 22 .”

“, 1999) and spatiotemporal sequencing of behaviors (Graybiel, 1998; Hikosaka et al., 1998; Jin and Costa, 2015).”

“The release of such cognitive and motor sequences or “action chunks” become pathological because they are no longer under the control of intent despite being consciously performed [55].”

“…and cognition-related signals through the cortico-basal ganglia loops in terms of the well-known direct and indirect pathways (biased control) (Alexander and Crutcher, 1990; Kimura, 1995; Mink, 1996; Boussaoud and Kermadi, 1997; Fukai and Tanaka, 1997; Graybiel, 1998; Hikosaka et al., 2000).”

“…difference of at least p < 0.05—see De Leonibus et al. (2011) for details. activity peaked only at choice points in the maze, consistent with a slow learning model-free system that learnt to associate differentiable intra-maze states with actions (Graybiel, 1998; Yin and Knowlton, 2006).”

“Taken together, the results are consistent with the proposal that cholinergic interneurons facilitate the coordination between different striatal input–output modules, but are not necessary for the expression of a choice pattern (Graybiel (1998)).”

“Application of nonlinear basal ganglia with the cortex and thalamus (Figure 1; Graybiel, 1998).”

“The basal ganglia have long been considered important for the organization of sequential behavior (Benecke et al., 1987; Graybiel, 1998).”

“The striatum is critical to forming such SR associations, including the chunking of motor and cognitive actions following a cue (e.g., Graybiel, 1998).”

“grams that are retained in the cortex [44, 59–62].”

“These effects of dopamine receptor antagonism on action sequence performance are consistent with findings from nonhuman primate studies showing that dopamine transmission is preferentially involved during the early stages of action sequence learning (53, 54), when discrete actions are being integrated into sequence-le…”

“…with the discovery of three distinct neural memory and learning systems for explicit or declarative memory, affective memory, and implicit or procedural memory – the latter related to habit learning (e.g. McDonald and White, 1993; Salmon and Butters, 1995; Knowlton et al., 1996; Graybiel, 1998).”

“Plasticity of cortical synaptic inputs to DLS is thought to play a role in striatum-dependent learning and memory (Graybiel, 1998; Gerdeman et al., 2003).”

“Graybiel (1998) has proposed that cholinergic interneurons modulate corticostriatal activity in different clusters of matrisomes, which leads to a recoding of corticostriatal information.”