Abstract
Mounting evidence suggests that the right cerebellum contributes to verbal working memory, but the functional role of this contribution remains unclear. In an established theory of motor control, the cerebellum is thought to predict sensory consequences of movements through an internal “forward model.” Here, we hypothesize a similar predictive process can generalize to cerebellar non-motor function, and that the right cerebellum plays a predictive role that is beneficial for rapidly engaging the phonological loop in verbal working memory. To test this hypothesis, double-pulse transcranial magnetic stimulation (TMS) was administered over either the right cerebellum or right occipital lobe (control site), on half the trials, to interrupt the rehearsal of a 6-letter sequence. We found that cerebellar stimulation resulted in greater errors in participants’ report of the letter in the current position. Additional analyses revealed that immediately after cerebellar TMS, participants were more likely to use out of date information to predict the next letter in the sequence. This pattern of errors is consistent with TMS causing a temporary disruption of state estimation and cerebellar forward model function, leading to prediction errors in the phonological loop.
Highlights
Previous neuroimaging studies of verbal working memory suggest that regions in left inferior temporal/parietal regions are associated with the phonological store, and the left inferior frontal regions are associated with articulatory control process (Paulesu et al, 1993; Awh et al, 1996; Fiez et al, 1996)
We examined whether the mean reaction time (RT) difference between transcranial magnetic stimulation (TMS) vs. non-TMS trials differed by probe position and stimulation site
We found that TMS administration to the right cerebellum, applied during covert rehearsal of a remembered sequence of letters, resulted in an interference with participants’ ability to identify whether a probe letter is in the correct position
Summary
It has become widely accepted in recent years that the human cerebellum contributes to motor function, and to a wide range of non-motor cognitive functions (for reviews, see Stoodley, 2012; Buckner, 2013; Schmahmann, 2019), such as verbal working memory (Chein and Fiez, 2001; Chen and Desmond, 2005; Justus et al, 2005; Ravizza et al, 2006; Hayter et al, 2007; Durisko and Fiez, 2010; Marvel and Desmond, 2010; Peterburs et al, 2010, 2016; Stoodley et al, 2012), executive function (Grafman et al, 1992; Rao et al, 1997; Schmahmann and Sherman, 1998; Karatekin et al, 2000; Neau et al, 2000; Bellebaum and Daum, 2007; Balsters et al, 2013; Wu et al, 2013; Rentiya et al, 2017), and language (Petersen et al, 1989; Desmond et al, 1998; Fulbright et al, 1999; Leggio et al, 2000; Lurito et al, 2000; Seger et al, 2000; Moretti et al, 2002; Xiang et al, 2003; Grönholm et al, 2005; Frings et al, 2006; Ben-Yehudah and Fiez, 2008; Rauschecker et al, 2008; Mariën et al, 2009, 2014; Stoodley and Schmahmann, 2009; Highnam and Bleile, 2011; Argyropoulos and Muggleton, 2013; Keren-Happuch et al, 2014). In the current study, we propose the right cerebellum plays a predictive role, similar to forward models in motor control, that is beneficial for rapidly engaging the phonological loop in verbal working memory.
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